CWD Response Plan

Proposal

Prevention. Surveillance. Response.

Last Updated: 09/12/2019

1
We Want Your Input
The Pennsylvania Game Commission is seeking your input on this response plan proposal. After
reading the information please provide comment at:
https://www.pgc.pa.gov/Wildlife/WildlifeRelatedDiseases/Documents/Draft%20CWD%20Response%20
Plan%20Public%20Comment%20form.pdf

Once complete, please send the document with responses to the Game Commission via email at
infocwd@pa.gov or via mail addressed to:
Attention: CWD Comments, 2001 Elmerton Avenue, Harrisburg, PA 17110

2
Living Document
This document is intended to be a living document that will continue to be updated as input
from the public, external collaborators, and Pennsylvania Game Commission staff necessitate
changes to previous versions. Expect changes to occur frequently.

3
Table of Contents
Title Page…………………………………………………………………………………………………………………………………..1
Public Input………………………………………………………………………………………………………………………………..2
Living Document Disclaimer…………………………………………………………………….…………………………………3
Acknowledgements……………………………………………………………………………………………………………………5
Executive Summary……………………………………………………………………………………………………………………6
Introduction………………………………………………………………………………………………….……………………………6
Disease Overview………………………………………………………………………………….……………………….6
Human Health……………………………………………………………………………………………………….……….8
Impact to Deer Populations………………………………………………………………………………….……..…8
Agency Responsibility………………………………………………………………………………………….…………9
Captive Cervid Facilities…………………………………………………………………………………….….………10
Stakeholder Responsibility…………………………………………….……………………….………….………..10
Impacts to Hunting and Communities…………………………………………………………………….…….11
History of Chronic Wasting Disease in Pennsylvania………………………………………….…………11
Strategies in Other States and Countries……………………………………………………………..……….14
Lessons Learned in Pennsylvania………………………………………………………………………………….16
Management Goals and Objectives………………………………………………………………………………………….16
Management Actions……………………………………………………………………………………………………………….20
CWD Control Units and CWD-Established Areas……………………………………………………………20
Management Actions in CWD Established Areas ………………………….………………………………21
Management Actions in CWD Control Units…………………………………………………………………24
Elk Herd Considerations……………………………………………………………………………………………….27
Conclusion……………………………………………………………………………………………………………………………….28
Literature Cited………………………………………………………………………………………………………………………..29
Appendix 1: Distance Sampling to Estimate Deer Abundance………….………………………………………35

4
Acknowledgments
We thank the many collaborators that contributed to this draft in a variety of capacities. Some
of these include: Dr. Krysten Schuler from Cornell University, Dr. Bryan Richards from USGS,
Nick Pinizzotto from the National Deer Alliance, Kip Adams from Quality Deer Management
Association, Dr. Michael Samuel now retired from University of Wisconsin-Madison, Dr.
Michelle Carstensen and Dr. Adam Landon from Minnesota Department of Natural Resources,
Dr. Tami Ryan from Wisconsin Department of Natural Resources, Dr. Kelly Straka from Michigan
Department of Natural Resources, Dr. Craig Miller from the University of Illinois, Dr. Lisa
Murphy and Dr. Julie Ellis from the University of Pennsylvania, Harris Glass and Kyle Van Why
from USDA-Wildlife Services, Dr. Kevin Brightbill and Dr. David Zellner from the Pennsylvania
Department of Agriculture, Emily Domoto, Ryan Reed and others from Pennsylvania
Department of Conservation and Natural Resources, Robb Miller-the Governor’s Advisor for
Hunting, Fishing, and Conservation, and many staff from the Pennsylvania Game Commission.

5
Executive Summary
Since Chronic Wasting Disease (CWD) was first discovered in Colorado in 1967, it has continued
to spread throughout North America and into other continents. CWD is a serious threat to the
deer and elk in Pennsylvania. Despite preventative measures and extensive monitoring, CWD
has continually increased in prevalence and geographic spread in Pennsylvania since its
discovery in 2012. Data from other states suggests that with no change, Pennsylvania will reach
a CWD prevalence over 30% in the area where CWD was first detected in the next 10-20 years.
The Pennsylvania Game Commission, hunters, landowners and others have a responsibility to
protect deer and elk populations for current and future generations. More aggressive
management actions are needed to effectively manage CWD. Control Units will be created
around new, isolated cases where CWD can be managed most effectively. Management
changes may include increased antlerless permit allocation, additional antlered permits,
expanded hunting seasons, removal of antler point restrictions, mandatory sampling in Control
Units, and consideration of incentive programs. The main goals of CWD management in
Pennsylvania are: 1) Reduce risks of spreading CWD, 2) Monitor for spatial distribution and
prevalence of CWD, 3) monitor for spatial distribution and prevalence of CWD, 4) prevent the
establishment of CWD in new areas, and 5) maintain prevalence below 5% in CWD-Established
Areas and slow the geographic spread of CWD. In Control Units, the initial management step is
to reach a minimum sampling goal via hunting. If management objectives are not met in CWD
Control Units or Established Areas, the Pennsylvania Game Commission will seek to reach those
objectives through agency-directed targeted removals. Hunters and landowners will be allowed
more opportunity to actively participate in CWD management.

Introduction
Disease Overview
Chronic Wasting Disease (CWD) is an always fatal brain disease, currently affecting white-tailed
deer and is threatening elk in Pennsylvania. CWD has been identified in all North American
cervid species, which includes deer, elk, caribou and moose (Williams et al. 2002). CWD was
first detected in a captive deer facility in Colorado in 1967. It wasn’t until the 1980s that CWD
was detected in an adjacent free-ranging elk. As of 2018, CWD has been detected in 26 states in
the US, 4 Canadian provinces, South Korea, and Scandinavian countries (USGS 2019).
CWD belongs to a family of diseases known as transmissible spongiform encephalopathies
(TSE). Other TSEs include scrapie in sheep, bovine spongiform encephalopathy (mad cow
disease) in cattle, and Creutzfeldt-Jakob disease in humans. TSEs are believed to be caused by
prions or misfolded proteins. Prions are a “proteinaceous particle” consisting solely of protein,
with no nucleic acid genome (Prusiner 1998). While it is not known what causes normal prions
to misfold, normal prions can convert to an abnormal or distorted form. These abnormal prions
replicate by binding to and misfolding normal prion proteins, they accumulate in lymphoid and

6
central nervous system tissues, eventually reaching the brain and causing clinical disease by
creating holes in the brain (Cohen 1999).

Figure 1. Distribution of Chronic Wasting Disease in North America as of 2019.

CWD transmission occurs through direct animal-to-animal contact or indirectly through prion
contaminated environments (Miller et al. 2004, Johnson et al. 2006). CWD-infected individuals
shed prions through saliva, urine, and feces (Mathiason et al. 2009). Despite research
conducted, there is currently no vaccine or treatment for CWD. Once in the environment, CWD
may remain infectious in the soil for several years (Miller et al. 2004). CWD prions can bind to
montmorillonite, a type of clay in soil, which may increase CWD infectivity (Johnson et al.
2006). Other laboratory research has shown that certain plants can absorb and uptake small
amounts of prions from contaminated soils, suggesting plants could also be a potential route of
infection (Rasmussen 2014, Pritzkow et al. 2015). While attempts to decontaminate infected
locations have been unsuccessful, a recent study found that humic acids extracted from soil
organic matter may decrease prion infectivity (Kuznetsova et al. 2018).
CWD, like other TSEs, is characterized by having long incubation periods before the animal
appears diseased. CWD-infected deer don’t show clinical signs of disease for up to 18 to 24
months (Tamgüney et al. 2009). Clinical symptoms of disease include lowered head, lowered

7
ears, progressive weight loss, rough hair coat, excessive salivation, excessive thirst, excessive
urination, and behavioral changes (Williams 2005). These clinical signs are not unique to CWD
and could be due to many other conditions, such as malnutrition or vehicle trauma. CWD-
infected individuals are more likely to die from predators, vehicle collisions, or hunters before
CWD causes death because of the increased vulnerability (Krumm et al. 2005, Edmunds et al.
2016, DeVivo et al. 2017).
Genetics plays a role in CWD susceptibility. Variations in the prion protein (PRNP) gene can
affect individual susceptibility to and progression of CWD (Robinson et al. 2012). Although
there is no genetic immunity, some rare genotypes are associated with lower rates of CWD
infection and slower progression of clinical disease (Hamir et al. 2006). However, the individuals
with the less susceptible genotype are not immune to CWD. Certain populations of deer in
Pennsylvania (Bedford, Blair, Huntingdon, Franklin, and Fulton) have a significantly higher
frequency of the more susceptible genotype than deer populations previously surveyed from
New Jersey, Wisconsin, Wyoming, and Canada (Blanchong et al. 2009, Robinson et al. 2012,
Miller and Walter 2019). Therefore, if not managed effectively, CWD might have a greater
impact on the deer population in portions of Pennsylvania compared to other regions.
Human Health
To date, there have been no known cases of CWD infecting humans. However, the Centers for
Disease Control and Prevention recommends no one consume CWD-infected meat (CDC, 2019).
Researchers continue to investigate the susceptibility of human infection to CWD, but no link
has been found to date. Experimental infections demonstrated that primates (squirrel
monkeys) could acquire CWD through oral consumption of infected meat or brain tissue (Race
et al. 2014, Waddell et al. 2018). Consequently, people should avoid exposure to CWD through
handling or consuming of infected animals. Hunters are advised to use gloves whenever
handling a deer carcass. Within Disease Management Areas (DMAs), hunters can receive free
CWD test results by submitting their deer head into a head collection bin. Locations of head
collection bins can be found on the Pennsylvania Game Commission website: www.pgc.pa.gov
Impacts to Deer Populations
CWD can have population-level impacts in areas of high prevalence. In Wyoming, where CWD
was first detected in 1985, there was a 10% annual decline in white-tailed deer populations in
an area with high CWD prevalence (Figure 2). CWD-infected white-tailed deer were 4.5 times
more likely to die annually than uninfected deer (Edmunds et al. 2016). Mule deer research also
showed annual population declines greater than 20% in an area with a high CWD prevalence
(DeVivo et al. 2017). In Colorado, the estimated mule deer population declined in a study area
of high CWD prevalence (>25 %). In the same area, estimated average life expectancy for
infected deer was an additional 1.6 years, compared to an additional 5.2 years for uninfected
deer (Miller et al. 2008). In some areas of Wisconsin, more than 55% of adult males and 35% of

8
adult females were infected with CWD; 75% of these infected deer died in the first year of study
(Wisconsin Department of Natural Resources 2019).

Figure 2. In the absence of disease management in Hunt Unit 65 in Southeastern Wyoming, the
CWD prevalence has increased since 2001, while CWD has contributed the decline of mule deer
(DeVivo et al. 2017) and white-tailed deer (not shown; Edmunds et al. 2016) over the same time
span. Source: Wyoming Game and Fish Department – Big Game Reports

Agency Responsibility
The Pennsylvania Game Commission is responsible for managing Pennsylvania’s free-ranging
mammals and birds for current and future generations. It would be irresponsible for the
Pennsylvania Game Commission not to take the threat of CWD seriously. The alternative option
of taking no further actions to combat CWD would also be irresponsible, given the current
knowledge of the disease and how it has impacted cervid populations in other states. State
wildlife agencies should manage for managing healthy, sustainable, free-ranging cervid
populations. The Pennsylvania Game Commission will continue to utilize the best available
science to make conscientious management decisions about CWD. It is imperative that the
Pennsylvania Game Commission take measures to prevent the disease from increasing in
prevalence and from spreading to other parts of the state. Doing this will ease CWD
management in the future if new, more acceptable and effective management strategies are
developed.

9
Captive Cervid Facilities
Since 2012, 16 captive cervid facilities (i.e. deer farms) have tested positive for CWD in
Pennsylvania. Eight captive herds that tested positive continue to have live animals on the
premise. All these facilities owned/own white-tailed deer. While the Pennsylvania Game
Commission has no regulatory authority over captive cervid facilities, the Pennsylvania Game
Commission will continue to work with the Pennsylvania Department of Agriculture (PDA; who
regulates the captive cervid industry) to create solutions that protect the deer and elk of
Pennsylvania. Some of those solutions include preventing contact between free-ranging and
captive deer (i.e. double fencing), prevent movement of captive cervids in high-risk CWD areas
(e.g. DMA 2) to areas outside DMAs to prevent introduction of CWD into previously uninfected
areas, mandatory CWD testing of all deer that die in captive cervid facilities, and mandatory
CWD sampling of a minimum number of animals tested per year in each captive cervid facility.
Stakeholder Responsibility
Hunters, Landowners, and other stakeholders occupy a critical role in CWD management.
Without their support and participation, we cannot be successful at managing this disease. The
Pennsylvania Game Commission will continue to engage hunters, landowners, other
governmental agencies, universities, and non-governmental organizations in the development
and implementation of a CWD response plan.
For over 100 years, hunters have been at the forefront of wildlife conservation success stories.
In the 1800s, hunters and anglers realized that to conserve rapidly disappearing wildlife they
would have to set limits and take responsibility for future management. Managing CWD in
Pennsylvania will be no different. Experiences from other states have shown when state wildlife
agencies and hunters do not work together, CWD is not successfully managed (Uehlinger et al.
2016). All stakeholders must stay informed, involved, and assist in managing CWD.
Particularly, hunters have an obligation to understand and abide by all current regulations and
best practices regarding CWD. For example, it is unlawful to transport high-risk parts outside of
DMAs.
High-risk cervid parts include the head (more specifically the brain, tonsils, eyes, and
retropharyngeal lymph nodes); spinal cord/backbone; spleen; skull plate with attached antlers,
if visible brain or spinal cord matter is present; upper canine teeth, if root structure or other
soft material is present; any object or article containing visible brain or spinal cord material;
unfinished taxidermy mounts; or brain-tanned hides.
Once the high-risk parts are removed, the following items can be brought into Pennsylvania, or
exported from a DMA: the remaining meat, on or off the bone; skull plate with antlers
attached, if no brain or spinal cord material is present; cleaned capes with no visible brain or
spinal cord material present; tanned or raw hide with no visible brain or spinal cord material

10
present; upper canine teeth, if no root structure or soft material is present; and finished
taxidermy mounts.
Movement of infected cervid carcasses, especially high-risk parts, is 1 of the known risks for
introducing CWD prions to new areas. Once infected, prions primarily accumulate in the central
nervous system (Kretzschmar et al. 1986), however prions have been detected in blood, saliva,
urine, and feces (Angers et al. 2006, Kramm et al. 2017). Since prions accumulate in the central
nervous system, the parts listed above are considered high-risk for transmitting disease.
It is best to dispose of high-risk parts in the commercial trash or a high-risk parts dumpster
provided by the Pennsylvania Game Commission. Within DMAs, it is unlawful to feed deer or
use urine-based lures. Not abiding by these regulations can increase the spread and severity of
the disease. Hunters can also take advantage of the increased role that the Pennsylvania Game
Commission wants hunters to play in CWD management. In part, this includes increased
hunting opportunity in critical areas, submitting samples for free CWD testing, and working
with the Pennsylvania Game Commission to develop solutions.
Landowners can also help by staying up to date with current information, educating others with
correct information, and allowing access for hunting or targeted removals on property.
For more information on what you can do to help control CWD, visit the Pennsylvania Game
Commission website: https://www.pgc.pa.gov
Impacts to Hunting and Communities
Declines in deer or elk populations can cause reduced hunter opportunity (DeVivo et al. 2017).
Even before populations decline from CWD, hunters may choose to hunt elsewhere or stop
hunting altogether. The first year after CWD was discovered in Wisconsin, hunting license sales
dropped 10% (Wisconsin Department of Natural Resources 2016). Other wildlife conservation
programs dependent on license sales, suffered as a result (Vaske et al. 2004). Over 1 in 5
hunters in Pennsylvania said their interest in deer hunting would be reduced if any deer were
detected with CWD in the area they hunt (Pennsylvania Game Commission, unpublished data,
2017). The potential economic losses of a decline in hunting participation in Pennsylvania would
have significant negative impacts on funding for conservation of all wildlife. In addition, CWD
could negatively affect the social and economic stability of communities that depend on
hunting for revenue.
History of Chronic Wasting Disease in Pennsylvania
The Pennsylvania Game Commission began monitoring for CWD in 2002, 10 years before it was
detected in Pennsylvania. As of 2019, nearly 80,000 deer and over 800 elk have been tested for
CWD in Pennsylvania. Surveillance efforts include testing road-killed deer, hunter-harvested
deer and elk, escaped captive deer or elk, and deer/elk showing clinical signs of CWD.

11
CWD was first detected in Pennsylvania in October 2012 in a captive white-tailed deer facility in
Adams County. Shortly after, CWD was detected in 3 free-ranging white-tailed deer in Bedford
and Blair county. Since, CWD prevalence has increased and CWD has spread geographically. As
of May 2019, a total of 250 free-ranging deer have tested positive for CWD in Pennsylvania
(Figure 3). Two hundred forty-six of these positives were detected in Disease Management Area
2, with the remaining 4 positives detected in Disease Management Area 3. Not only has the
number of deer detected continually increased over the years, the percentage of deer that test
positive for CWD (i.e. prevalence) has also continually increased (Figure 4).
When new CWD detections are found in free-ranging or captive deer in Pennsylvania, a 10-mile
radius is created around that detection to either expand or establish a Disease Management
Area. If a new CWD positive is detected near a DMA’s boundary or outside of an existing DMA,
the existing DMA could expand into a larger area or a new DMA may be created. A 10-mile
buffer around positive CWD detections is used because that encompasses the dispersal
distances of the majority of deer in Pennsylvania (Long et al. 2008; Lutz et al. 2015). Enhanced
CWD monitoring and specific regulations exist within DMAs to help slow the spread of CWD.

140
123
120
No. of known Positives

100
79
80

60

40
25
20 12
3 2 5
0
2012 2013 2014 2015 2016 2017 2018
Year
Figure 3. Number of positive free-ranging deer detected in Pennsylvania by year from statewide
CWD testing.

12
 5

3
Prevalence %

0
2012 2013 2014 2015 2016 2017 2018
Year
Figure 4. Prevalence (% of all samples that test positive) by year from all samples taken in Blair,
Bedford, and Fulton Counties, Pennsylvania. Note: the large drop in prevalence from 2012 to
2013 likely stems from “sampling error” because of a low sample size in 2012, not a true drop in
prevalence. Samples sizes from 2012-2018 were 142, 1006, 780, 1253, 1257, 2346, and 2382
samples, respectively.
DMA 1 was established in 2012 in Adams County after a captive cervid facility (i.e. deer farm)
tested positive for CWD. All deer were depopulated from the farm and DMA1 was dissolved in
2017 after no additional deer were detected with CWD for 5 consecutive years.

DMA 2 was established in 2012 after the detection of 3 free-ranging deer tested positive in Blair
and Bedford County. DMA 2 originally covered 892 square miles in Blair and Bedford counties.
Due to the discovery of new CWD cases every year, DMA 2 in 2019 now covers 6,715 square
miles in portions of 17 counties in south-central Pennsylvania. Eleven captive cervid facilities
have tested positive for CWD within DMA 2. Three farms have depopulated while the remaining
7 have live deer on the premises.

DMA 3 was established in 2014 after CWD was detected in 2 captive cervid facilities in Jefferson
county. Those farms were depopulated of all animals. In 2017, Disease Management Area 3
expanded to cover over 2,000 square miles due to the detection of CWD in 3 free-ranging
white-tailed deer in Clearfield and Jefferson county. DMA 3 was expanded further in 2019
because a deer within a captive cervid facility in Clearfield County tested positive after being
transported from a captive cervid facility in DMA 2 (Fulton County). Another free-ranging deer
(from a vehicle collision) tested positive in Jefferson County in 2019.

13
DMA 4 was established in 2018 after CWD was detected in a captive cervid facility in Lancaster
county. The farm was depopulated of all deer. Disease Management Area 4 covers 346 square
miles in Lancaster, Lebanon, and Berks counties. To date, no additional CWD detections have
been found in Disease Management Area 4.

Figure 5. Current Disease Management Areas (DMAs; blue polygons) in Pennsylvania. (Note:
DMA 1 that surrounded the captive cervid facility in Adams County was dissolved after 5 years
of no additional positive detections)

Strategies in Other States and Countries

Options to manage CWD where it has become established are currently limited, however where
success has occurred, states used a combination of increased hunter harvest and targeted
removals (i.e. agency-directed culling) to reduce CWD prevalence. For example, Illinois first
detected CWD in 2002 and actively began targeted removals of deer the following year.
Targeted removals in Illinois remove CWD-infected deer at 2.5 times the rate that hunters do in
the infected areas because targeted removals focus on removing deer near known CWD
detections (Dufford and McDonald 2018). Through consistent management actions for over 15
years, prevalence in the area it was originally detected is currently 3.3% (Dufford and McDonald
2018). In contrast, Wisconsin also detected CWD in 2002 and began targeted removals of deer
in 2003. Due to public pushback against Wisconsin Department of Natural Resources, Wisconsin
ceased targeted removals in 2007 (Uehlinger et al. 2016). Since that time, CWD has spread
geographically and increased in prevalence, with some areas showing over 55% of adult bucks

14
infected (Wisconsin Department of Natural Resources 2019). West Virginia has undertaken
similar management actions to that of Pennsylvania and continues to see prevalence increase
in their disease zones (Figure 6).

Figure 6. CWD Prevalence trajectory since detection for West Virginia, Wisconsin, and
Pennsylvania (Pennsylvania Game Commission, unpublished data, 2018;
West Virginia, unpublished data, 2018; Wisconsin DNR 2019).

Several examples exist where CWD was detected in free-ranging deer and did not become
established in the local deer population after aggressive management actions were taken. In
2005, New York detected CWD in a captive deer facility in Oneida county. Within weeks, deer
managers used a combination of hunter harvest and targeted removals to remove over 300
deer in the local area. Two additional free-ranging deer were detected during their initial
response. New York has not detected additional CWD positives since their initial response in
2005 (New York State Department of Environmental Conservation, 2019). Similarly, Minnesota
detected CWD in a free-ranging deer in Olmsted County in 2010, which was only 2 miles from a
CWD-positive captive deer facility. Minnesota implemented immediate management actions
within the local area via a combination of additional hunter harvest and agency-sponsored
targeted removals. This effort resulted in over 4,000 deer samples over 3 years. No additional
CWD positives were found in the local free-ranging deer population (Minnesota DNR, 2019).

These success stories from other states provide hope that increased hunter harvest and
targeted removals, if conducted quickly and effectively after CWD is detected in a new area,
can be used to successfully combat CWD. To date, reducing local deer populations is the only

15
management strategy that has shown any success at stabilizing or reducing prevalence of CWD
(Manjerovic et al. 2014). Reducing deer populations surrounding new CWD detections lowers
disease transmission through reduced contact among deer (Tosa et al. 2017). In addition,
reducing deer numbers around new CWD detections, reduces the number of deer shedding
prions into the environment. Norway has taken the most aggressive steps to eliminate CWD by
removing an entire reindeer herd, over 2,400 animals, in an area where CWD had been
detected. They plan to leave this area devoid of animals for 5 years to try to eliminate the
disease (the Pennsylvania Game Commission will not attempt to eradicate deer or elk in any
areas).

Lessons Learned in Pennsylvania

After more than 6 years of managing CWD in Pennsylvania, it is clear that controlling CWD in
Pennsylvania’s free-ranging white-tailed deer is extremely challenging. Managing CWD requires
substantial commitment of human and financial resources from the Pennsylvania Game
Commission over an extended period of time. If the trajectory of disease increases in
Pennsylvania as it has in other states, with no change in management actions, prevalence in the
Established Area (portions of Blair, Bedford, and Fulton county) will likely exceed 30% in the
next 10-20 years. Without a change in management, CWD will continue to increase and spread.

Although the public generally does not want to see the disease increase, reducing deer
populations to manage CWD is controversial. However, alternative options for managing CWD
in free-ranging deer are limited. Communication efforts will be critical to increase the public’s
understanding of, support for, and participation in CWD management. The Pennsylvania Game
Commission is committed to increasing stakeholder engagement and outreach to local
communities, hunters, and landowners affected by CWD. In addition to management of CWD,
the Pennsylvania Game Commission will conduct human dimensions studies to inform future
management decisions.

Management Goals and Objectives

Goal 1: Reduce Risks of Spreading CWD
Objective 1: Contain disease and stop introduction of prions via movement of high-risk
cervid parts.
When there is a new CWD detection in either a captive or free-ranging cervid, a 10-mile
radius DMA is established. If a new CWD positive is detected within 10-miles of a
current DMA boundary, the existing DMA may expand into a larger DMA. With DMAs,
specific rules apply to help prevent the spread of CWD into uninfected areas.
Within the state, it is unlawful to remove or export high-risk cervid parts from a DMA. In
addition, it is also unlawful to import high-risk cervid parts from CWD-positive states. As

16
 of May 2019, the importation of high-risk cervid parts are prohibited from 26 states and
3 Canadian provinces.
Objective 2: Reduce the risks of transmission between captive and free-ranging deer or
elk.
Work collaboratively with the Pennsylvania Department of Agriculture and the captive
cervid industry to minimize the risks to both free-ranging and captive cervids. Some of
the solutions include preventing contact between free-ranging and captive deer,
prevent movement of captive cervids in high risk CWD areas (DMAs) to areas outside
DMAs to prevent introduction of CWD into previously uninfected areas, mandatory
CWD testing of all deer that die in captive cervid facilities, and mandatory CWD
sampling of a minimum number of animals tested per year in each captive cervid facility.
Objective 3: Limit human-related activities that unnaturally congregate free-ranging
cervids.
Within DMAs, it is illegal to feed free-ranging cervids. Research suggests that CWD is
transmitted by direct contact between infected and uninfected individuals and indirectly
through prion contaminated environments. Prions can be shed onto the environment
through saliva, urine, feces (Mathiason et al. 2009), or through infected carcasses (Miller
et al. 2004). Once in the environment, prions can remain infectious for several years
(Johnson et al. 2006).
Feeding can negatively affect disease transmission by increasing contact rates and
promoting pathogen accumulation around feeding sites (Murray et al. 2016). Feeding,
including the use of mineral licks, causes deer and elk to congregate in ways they would
not do so naturally. In addition, deer and elk urinate and defecate in these same areas.
If 1 infected deer is in the area, increasing contact between individuals and increasing
environmental contamination only increases the risk of spreading CWD. For this reason,
feeding—including the use of mineral licks—is illegal in Pennsylvania’s DMAs.
Within DMAs, the use and possession of natural urine-based attractants is also
prohibited. As stated above, prions can be shed through saliva, urine, and feces
(Mathiason et al. 2009, Gough et al. 2010). While the amount of prions required to
cause clinical or subclinical infection of CWD is currently unknown, the use of natural
urine-based attractants still poses a potential risk of transmitting CWD. The
Pennsylvania Game Commission prohibits the use of urine-based attractants within
DMAs for the purpose of avoiding the unnatural congregation of deer or elk to a specific
area.
Goal 2: Monitor for Spatial Distribution and Prevalence of CWD
Objective 4: Continue state-wide surveillance efforts to enhance likelihood of detecting
new introductions of CWD.
17
Early detection of CWD is critical to implement management strategies that are aimed
at preventing the establishment of CWD in new areas. The Pennsylvania Game
Commission will continue statewide surveillance efforts for CWD. State-wide
surveillance efforts in Pennsylvania includes random sampling of hunter-harvested deer,
immediate removal and testing of escaped captive cervids, and collection and testing of
any free-ranging deer or elk behaving abnormally or displaying clinical signs of CWD.
Hunter-harvested deer are randomly sampled from cooperating meat processors and
across the state. Random surveillance of free-ranging white-tailed deer has a low
probability of detecting CWD at low (<1%) prevalence but can contribute to CWD
detection and monitoring (Pennsylvania Game Commission, unpublished data 2012).
Given limitations of active surveillance as a CWD detection program, free-ranging
surveillance is designed to: 1) train and test sample collection, extraction, and testing
procedures and personnel, 2) annually sample and test 4,000 hunter-killed deer 3)
provide scientific baseline data on presence and distribution of CWD in Pennsylvania, 4)
increase public awareness concerning CWD surveillance activities, and 5) potentially
provide a platform for additional disease monitoring.
Our active surveillance sampling design for deer is based on Wildlife Management Units
(WMUs). Quotas are created for each WMU based on the proportion of the state’s deer
population occurring in that WMU. Based on simulation modeling (Diefenbach et al.
2004), we expect a sample of 4,000 will provide a 62% chance of detecting CWD if
prevalence is 1% in any WMU. This sample size represents the maximum number that
can be collected by our personnel and infrastructure. Ongoing surveillance strengthens
our abilities for future CWD events.
All successful elk hunters are required to present their harvest at a designated check
station. At the check station, trained Pennsylvania Game Commission staff extract tissue
samples (i.e. obex and retropharyngeal lymph nodes) for testing from all harvests,
unless the hunter intends to have their harvest mounted. If the hunter intends to get
their harvest mounted, instructions and materials including postage-paid shipping
containers for transport of the caped head to the laboratory are provided to the hunter.
The objective for hunter-harvested elk is to collect samples from all harvested elk over 1
year of age.
All hunter-harvested elk are tested for CWD. As of May 2019, the Pennsylvania Game
Commission has tested 809 free-ranging elk for CWD and to date CWD has not been
detected in Pennsylvania’s free-ranging elk. Samples from hunter-harvested white-tailed
deer are collected in one of two ways: through the voluntary submission of hunter-
harvested deer heads in head collection bins provided by the Pennsylvania Game
Commission or through a random collection of hunter-harvested heads taken from meat
processors throughout the state.

18
 Objective 5: Continue surveillance efforts within Disease Management Areas to estimate
prevalence.
In addition to state-wide surveillance efforts noted above, increased surveillance occurs
within DMAs through sampling voluntary submissions of hunter-harvest deer, and
sampling road-killed deer. Additional samples within DMAs are used to estimate CWD
prevalence and spatial distribution.
The Pennsylvania Game Commission provides head collection bins for voluntary
submissions of hunter-harvested deer within DMAs. To encourage hunters to help
surveillance efforts, the Pennsylvania Game Commission tests each submitted head at
no cost to the hunter. Hunters are instructed to keep the harvest tag attached to the ear
for identification purposes and to double-bag all submitted heads to prevent
contamination from other submissions. All samples from head collection bins are sent
to the Pennsylvania Veterinary Laboratory in Harrisburg for CWD testing through
enzyme-linked immunosorbent assay (ELISA) or immunochemistry (IHC). All samples
that test positive are then tested again for confirmation via another IHC test.
Contractors are employed within DMAs, to collect road-killed deer from state and local
roads. Contractors are instructed to deliver carcasses to designated deer processing
locations, where trained Pennsylvania Game Commission staff extract samples (i.e. obex
and retropharyngeal lymph nodes) for testing. Remaining carcass parts are discarded
into dumpsters within these areas and disposed of in approved landfills to prevent
environmental contamination.
Currently the only federally approved CWD tests for cervids are IHC, ELISA, and Western
blot. As a general practice, all suspect positive ELISA and Western blot tests should be
confirmed with IHC (USDA 2013). The obex and retropharyngeal lymph nodes are the
preferred tissue samples for testing because data suggests these may be early CWD
detection sites (Miller and Williams 2002, Spraker et al. 2004). Since all federally
approved CWD tests require the removal of the obex and retropharyngeal lymph nodes,
all tests are done postmortem (i.e. after death). There are no tests that can be done in
the field because ELISA and IHC are both analyzed in the lab and require several steps
over the course of days and sterile environments to reduce the likelihood of false
positive results.
Goal 3: Prevent the Establishment of CWD in New Areas, Once Detected
Objective 6: Reduce potential for disease transmission and limit environmental
contamination of prions by quickly and effectively reducing local free-ranging cervid
populations around new CWD detections.
Goal 4: Maintain Prevalence below 5% in Established Areas and Slow the Geographic Spread
of CWD

19
 Objective 7: Reduce free-ranging cervid populations within areas where CWD is
established and at highest infections rates.
CWD prevalence should remain below 5% in the established area. Prevalence will be
calculated from hunter-harvested deer only. However, it could take several years for
management actions to lower CWD prevalence. Lowering deer densities in small areas
with the highest prevalence might reduce natural movements of infected cervids to
uninfected areas of the state (long et al. 2008, lutz et al. 2015).

Management Actions
CWD Control Units and CWD-Established Areas

CWD-Established Areas are characterized by a contiguous area where CWD positive cases are
within 10 miles of one another and where CWD prevalence has remained >1% for at least 2
consecutive years (Figure 7). Currently, there would only be 1 established area in Pennsylvania,
which would be centered in Blair, Bedford and Fulton counties. The Established Area
boundaries will be delineated by easily identifiable boundaries such as roads or rivers.

A CWD Control Unit is created when a new, isolated CWD positive case is detected in free-
ranging or captive cervid greater than 10 miles from another positive detection. A 3-mile radius
buffer is drawn around the new, isolated detection. This 3-mile buffer is the foundation to
create a CWD Control Unit. The CWD Control Unit boundaries are then delineated to follow
easily identifiable boundaries such as roads or rivers. Therefore, the buffer may sometimes fall
outside or slightly inside of the 3-mile radius buffer.

CWD Control Units are areas within DMAs where the status of the disease is yet uncertain;
CWD may or may not be established in the environment (e.g. in the soil). For this reason, they
are of particular concern for quick management action. CWD infections exhibit a clustered
distribution, with highest prevalence occurring near the point of introduction (Osnas et al.
2009). This suggests that deer or elk in closer proximity to CWD detections are more likely to
become infected with CWD (Joly et al. 2006). Therefore, areas directly surrounding CWD
detections are considered high risk for transmission and increased management is needed to
control the disease. Within Control Units, the main objective is to remove CWD positive deer
quickly before CWD becomes established in the local deer population and/or environment (e.g.
soil). Once established in the environment, there is no known way to eradicate CWD. CWD
Control Units will not be created in areas where CWD is established in the environment. In
Established Areas, the objective is to maintain the CWD prevalence below 5%.

20
Figure 7. Map of the CWD-Established Area (within bold black line) and examples of CWD
Control Units (oranges circles) around new, isolated cases.

Management Actions in CWD-Established Areas

Since detection in Blair and Bedford counties in 2012, CWD prevalence has continued to
increase until today and has spread outward every year. In this area, CWD may have become
established in the environment and with current science it is unrealistic to eradicate the
disease. However, it is possible to stop the increase or reduce prevalence in this area (Dufford
and McDonald 2018). That is the main management objective for this area; to lower
prevalence, with the goal of slowing the spread of CWD. To date, increasing hunter harvest and
conducting targeted removals is the only management strategy that has shown any success in
stabilizing CWD prevalence (Manjerovic et al. 2014, Dufford and McDonald 2018).

Increased hunting opportunities and reduced restrictions will be offered to hunters within
CWD-Established Areas. Increased ease and flexibility will give hunters the first opportunity to
harvest more deer and collect more CWD samples. Management actions within CWD-
Established Areas will include:

1. Maximize hunting opportunities: Changes within CWD Control Units may include:
a) Expanded hunting seasons:

21
 Dates Arms & License Holders Eligible
Ammunition
Oct. 17-26, 2020* All methods All
Nov. 28 - Dec. 12, 2020** All methods All
Dec. 26, 2020 – Jan. 30, 2021*** All methods All
*Early Firearm Season: This season combines what was previously the dates for
the muzzleloader and special firearms season. Antlered and antlerless deer may
be harvested with all arms and ammunitions during this season.
**Regular Firearm Season: All license holders may harvest antlerless deer
throughout this season; concurrent antlered and antlerless seasons like is
currently implemented in WMUs 2B, 5C, 5D.
***Extended Firearm Season: Antlerless only (except with the use of flintlock
during flintlock season), as currently implemented in Allegheny, Bucks, Chester,
Delaware, Montgomery, and Philadelphia Counties.
b) Removal of antler point restrictions within DMAs: In Pennsylvania, adults bucks are
nearly 3 times more likely to be infected with CWD than all other deer (Table 1).
Removing antler point restrictions will allow hunters to harvest more bucks. Preliminary
modeling by USGS-PA Cooperative Research Unit demonstrated that removal of antler
point restrictions would reduce the number of deer infected with CWD by 20%.
Removing antler point restrictions and increasing antlerless harvest would reduce the
number of deer infected with CWD by 40% or more (D.R. Diefenbach, U.S. Geological
Survey, personal communication).

Table 1. Infection for fawns, yearlings, and adults from 2016-2018 in Blair, Bedford, and
Fulton counties of Disease Management Area 2.
Fawn Yearling ADULT
Both sexes Female Male Female Male
2016 NA 0.9 1.0 1.2 3.8
2017 1.9 2.2 1.5 1.3 5.2
2018 1.8 1.7 2.4 2.9 6.6
Average 1.9 1.7 1.8 2.0 5.5

c) Increased allocation of antlerless harvest tags: Hunters can harvest antlerless deer using
DMAP permits during any portion of the hunting season. Currently, DMAP units
administered by Pennsylvania Game Commission within DMAs allocate 1 permit per 50
acres or 1 permit per 25 acres, depending on the unit. This allocation can be increased in
the future to allow more hunters the opportunity to harvest antlerless deer and to meet
management objectives.

22
2. Consider incentive programs: Incentive programs could be an effective tool to increase land
access for hunters and allow additional deer to be taken in areas where it is most needed to
control CWD. During 2019-2020, Pennsylvania Game Commission will survey hunters and
landowners to determine if there are incentive programs that would be effective and
supported.

3.Targeted removals (as needed): The Pennsylvania Game Commission wants hunters to have
the first opportunity to meet management objectives through increased deer harvest. We
also want hunters to be part of the solution and be successful at managing CWD. Nearly all
results from CWD samples collected during hunting seasons should be analyzed by Feb 1. If
prevalence is not maintained below 5% in the CWD-Established Area, Pennsylvania Game
Commission will strive to reach that objective through agency-directed targeted removals. It
may take more than 1 year of management actions to reach the objective of reducing and
stabilizing CWD prevalence below 5%.

Targeted removals are small-scale operations, that would remove deer from small areas with
the highest concentration of positive detections (Figure 8). Targeted removals reduce the
number of infected deer on the landscape, the number of potential dispersers that can
spread CWD to new areas (Long et al. 2008, Lutz et al. 2015), and can lead to stable or
reduced levels of infection (Manjerovic et al. 2014, Dufford and McDonald 2018).

Figure 8. Map showing where CWD positive deer were detected at highest concentrations
(yellow) in Pennsylvania.
23
The actual number of deer remaining on the landscape to achieve disease management
objectives may vary by location. The objective is not to remove all deer from the area. The main
objective is to lower CWD prevalence and slow the spread of CWD. Reducing local deer
numbers is the mechanism to achieve the main objectives, not the objective itself. If the CWD
prevalence is declining or remains low (<5%), then deer density, regardless of the number – is
acceptable for disease management purposes.

Management Actions in CWD Control Units

Increased hunting opportunities and reduced restrictions will be offered to hunters within
Control Units. Increased ease and flexibility will give hunters the first opportunity to harvest
more deer and collect more CWD samples to meet management objectives.

1. Deer Density Survey: Before hunting seasons begin, surveys will be conducted within each
Control Unit to estimate deer density. The surveys will be conducted by Pennsylvania Game
Commission staff via a road-based, Distance Sampling method (Appendix 1).

2. Sample Collection via Hunter Harvest: The estimated deer density will determine the
minimum sample quota that we will need to test to be 99% confident we would detect CWD
if at a 1% prevalence. Each Control Unit will have a minimum sample size quota of 350 deer.
If deer densities are significantly higher than 30 deer per square mile, then a minimum
sample size will need to be larger than 350 deer to be 99% confident that we can detect
CWD at a 1% prevalence (given certain assumptions, Belsare et al. 2018). The minimum
sample size will be slightly different for each Control Unit, based on the population estimate
of deer. Across Pennsylvania, hunters harvest an average of 8 deer/mi2. To reach the
minimum sample size quota needed in an average Control Unit, hunters would have to
harvest approximately 4 deer/mi2 more than normal and submit all heads for testing. Head
collection bins will be available in each Control Unit. If no additional detections of CWD
occur after 3 years of reaching the minimum sampling quota, the Control Unit would be
dissolved.

a.) Mandatory sampling in Control Units: Hunters must submit samples of all deer
harvested in a Control Unit within 72 hours of harvest. Hunters should submit heads to
Pennsylvania Game Commission for testing via head collection bins that will be placed
within each Control Unit. Hunters are responsible for filling out and securing a tag to the
ear of the deer, removing the head, bagging the head, and placing it in a collection bin
for testing. Antlers should be removed before submitting the head into a head collection
bin. If hunters harvest a buck that they want to mount they may have the cape and skull
cap with antlers removed before submitting the head for sampling.

3. Landowner Interaction: All landowners that own more than 5 acres within a Control Unit will
receive written communication before September each year with detailed information
including: a) CWD has been detected nearby, b) requirements associated with hunting in a

24
 Control Unit, c) Control Unit boundaries, d) instructions on how to obtain more deer tags, if
desired, d) locations of head collection bins, e) an invitation to attend a local public meeting
to discuss the upcoming hunting season, sample size quota, and management changes, and
f) other pertinent information.
4. Maximize hunting opportunities: Changes within CWD Control Units may include:

d) Expanded hunting seasons:

i. Option 1- Same season dates that are in effect in Special Regulation Areas
ii. Option 2 – Same season dates as proposed in the CWD-Established Areas:
Dates Arms & License Holders Eligible
Ammunition
Oct. 17-26, 2020* All methods All
Nov. 28 - Dec. 12, 2020** All methods All
Dec. 26, 2020 – Jan. 30, 2021*** All methods All

*Early Firearm Season: This season combines what was previously the dates for
the muzzleloader and special firearms season. Antlered and antlerless deer may
be harvested with all arms and ammunitions during this season.
**Regular Firearm Season: All license holders may harvest antlerless deer
throughout this season; concurrent antlered and antlerless seasons like is
currently implemented in WMUs 2B, 5C, 5D.
***Extended Firearm Season: Antlerless only (except with the use of flintlock
during flintlock season), as currently implemented in Allegheny, Bucks, Chester,
Delaware, Montgomery, and Philadelphia Counties.

e) Removal of antler point restrictions within DMA: In Pennsylvania, adults bucks are
nearly 3 times more likely to be infected with CWD than all other deer (Table 1).
Removing antler point restrictions will allow hunters to harvest more bucks. Preliminary
modeling by USGS-PA Cooperative Research Unit demonstrated that removal of antler
point restrictions would reduce the number of infected deer by 20%. Removing antler
point restrictions and increasing antlerless harvest would reduce the number of infected
deer by 40% or more (D.R. Diefenbach, U.S. Geological Survey, personal
communication).

25
 Table 1. Prevalence for fawns, yearlings, and adults from 2016-2018 in Blair, Bedford,
and Fulton counties of Disease Management Area 2.
Fawn Yearling ADULT
Both sexes Female Male Female Male
2016 NA 0.9 1.0 1.2 3.8
2017 1.9 2.2 1.5 1.3 5.2
2018 1.8 1.7 2.4 2.9 6.6
Average 1.9 1.7 1.8 2.0 5.5

f) Additional antlered tags: Within Control Units, hunters will be allowed to purchase (at
administrative cost of $2-3) an additional antlered deer tag (may harvest a total of 2
antlered deer). Similar to removing antler point restrictions, additional antlered deer
tags will allow hunters more flexibility to harvest additional deer in areas where we
need additional CWD samples.

g) Increased allocation of antlerless harvest tags: Hunters can harvest antlerless deer using
DMAP permits during any portion of the hunting season. Currently, DMAP units
administered by Pennsylvania Game Commission within DMAs allocate 1 permit per 50
acres or 1 permit per 25 acres, depending on the unit. This allocation can be increased in
the future to allow more hunters the opportunity to harvest antlerless deer.

5. Consider incentive programs: Incentive programs could be an effective tool to increase land
access for hunters and allow additional deer to be taken in specific areas. During 2019-2020,
Pennsylvania Game Commission will survey hunters and landowners to determine if there
are incentive programs that would be effective and supported.

6. Special season: Between February and April, if a new CWD case is detected and a Control
Unit created, a 3-week hunting season would be opened to all landowners and any hunters
with landowner permission. Additional antlered and antlerless tags will be available during
this special season and any legal arms and ammunition (e.g. firearm) may be used.

7. Targeted Removals (as needed): The Pennsylvania Game Commission wants hunters to
have the first opportunity to harvest more deer first. We also want hunters to be the
solution and be successful at managing CWD. If the predetermined, minimum sample size
quota is not achieved or additional positives are detected, agency-directed targeted
removals will occur in Control Units. An adequate number of deer sampled within a CWD
Control Unit is necessary to estimate prevalence and spatial distribution of CWD. Targeted
removals are small-scale operations, occurring within 3 miles of CWD detections and
focused as close to known CWD positive locations as possible. The number of deer removed
via targeted removals in each CWD Control Unit will vary depending on the number needed
to reach sampling quotas for that area.
a. Additional positive cases: If additional positive cases are detected within a Control
Unit, the management objective within that Control Unit will switch from collecting

26
 a minimum sample size to reducing deer densities immediately surround the
positive cases. Reducing local deer densities will be accomplished through increased
hunter harvest and agency-directed targeted removals.

The actual number of deer remaining on the landscape to achieve disease

management objectives may vary by location. The objective is not to remove all deer
from the area. The main objective is to stop the disease before it becomes
established in the environment (e.g. in soil). Reducing local deer numbers is the
mechanism to achieve the main objectives, not the objective itself. If no further
positives are detected in a 3-year period, then deer density, regardless of the
number – is acceptable for disease management purposes.

b. Prioritization for targeted removals locations: Factors that will be considered when
prioritizing Control Units for targeted removal operations include: spatial proximity
to other CWD detections, the number of CWD detections in local area, cervid density
in local area, time elapsed since detection, age of sample (CWD-infected cervid), sex
of sample, whether sample was a captive or free-ranging cervid, proximity to
geographic barriers for dispersal, proximity to Pennsylvania’s elk hunt zones, and
effect on DMAs.

Elk Population Considerations

The Pennsylvania Game Commission adapts its CWD management actions for elk hunt zones
based on the biology of cervid species. CWD can reduce elk survival (Monello et al. 2014). It is
the Pennsylvania Game Commission’s responsibility to act in ways to ensure sustainable deer
and elk populations for current and future generations. Efforts to reduce the risk of CWD in the
elk population help the Pennsylvania Game Commission to meet this responsibility. Our CWD
management actions are motivated by an effort to protect both deer and elk and to minimize
the impact on deer and elk hunters.

At present, the Pennsylvania Game Commission’s response to the discovery of a CWD positive
deer or elk within Pennsylvania’s elk hunt zones is to enclose all the elk hunt zones into a
singular DMA (~2500 sq. mi). The primary motivations for these action are 1) elk are a social
species and movement between generally distinct social units has been documented in
Pennsylvania and in other eastern elk populations (J.E. Banfield, Pennsylvania Game
Commission, personal communication; Rosatte 2017), 2) bull elk in Pennsylvania are known to
travel up to 20 miles for breeding during the annual rut (J.E. Banfield, Pennsylvania Game
Commission, personal communication), and 3) elk have relatively large home ranges compared
to white-tailed deer in Pennsylvania (Table 2; Walter et al. 2018). Thus, elk have the potential
capability of spreading CWD farther distances than white-tailed deer.

27
 Table 2. Average home ranges (95%) for
white-tailed deer and elk in Pennsylvania.
Species/Sex Mean (sq. mi) SD n
Female WTD1 0.74 0.51 49
Male WTD1 1.75 0.87 12
Female ELK 6.65 2.24 13
Male ELK 11.23 3.21 14
1 Data
summarized across 5 study areas in
Pennsylvania (Walter et al. 2018)

Additionally, if CWD is detected within the elk management area, management responses will
be similar as previously described. One notable difference is that elk will not be taken through
targeted removals. Rather, increased hunter opportunity for elk will be allowed in certain areas.
The increase in the number of elk tags would depend on the hunt zones where CWD was
detected, because population estimates among hunt zones vary. Any elk that disperse out of
the elk management area and into a DMA are euthanized to prevent potential spread of CWD
back into the elk management area. Additional factors considering our response to CWD
infections in elk are: a) elk are generally at much lower densities than deer, b) multiple
pregnancies (twins and triplets) are extremely rare in elk (<1%; Kittams 1953; Flook 1970;
Houston 1982), c) the population estimate of elk in Pennsylvania, about 1,000 elk (Pennsylvania
Game Commission, 2019), is relatively low and thus ecologically fragile compared to deer in
Pennsylvania.

Conclusion
CWD is a serious, long-term threat to deer and elk in Pennsylvania. Without change and further
management action, CWD will continue to spread and increase in Pennsylvania. It is the
responsibility of hunters, landowners, Pennsylvania Game Commission, and many others to
take action to manage CWD effectively. Despite the continued increase of CWD in Pennsylvania,
successes from other states provide hope and guidance. With swift and effective management
actions we can slow the spread and reduce prevalence of CWD. To be successful, it will take
support and participation from all interested stakeholders, with responsibility and effort from
landowners and hunters being crucial.

28
Literature Cited

Angers, R. C., S.R. Browning, T.S. Seward, C.J. Sigurdson, M.W. Miller, E.A. Hoover, and G.C.
Telling. 2006. Prions in skeletal muscles of deer with chronic wasting disease. Science.
311: 1117.
Belsare, A.V., M.E. Gompper, B. Keller, J. Sumners, L. Hansen, J.J. Millspaugh. 2018. An agent-
based framework for improving wildlife disease surveillance: A case study of chronic wasting
disease in Missouri white-tailed deer. BioRxiv https://doi.org/10.1101/478610
Blanchong, J. A., D.M. Heisey, K.T. Scribner, S.V. Libants, C. Johnson, J.M. Aiken, J.A.
Langenberg, and M.D. Samuel. 2009. Genetic susceptibility to chronic wasting disease in
free-ranging white-tailed deer: complement component c1q and Prnp polymorphisms.
Infection, Genetics, and Evolution 9(6): 1329-1335.
CDC. 2019. Chronic Wasting Disease. https://www.cdc.gov/prions/cwd/index.html. Accessed 11
July 2019.
Cohen F. 1999. Protein misfolding and prion diseases. Journal of Molecular Biology. 293: 313-
320.
Colorado Parks and Wildlife. 2019. Deer hunting statistics.
https://cpw.state.co.us/thingstodo/Pages/Statistics-Deer.aspx. Accessed 10 July 2019.
Dearing, D. M., C. Clay, E. Lehmer, and L. Dizney. 2015. The roles of community and contact
rates on pathogen prevalence. Journal of Mammalogy 96: 29-36.
Diefenbach, D. R., C. S. Rosenberry, and R. C. Boyd. 2004. Efficacy of detecting chronic
wasting disease via sampling hunter-killed white-tailed deer. Wildlife Society Bulletin 32:
267-272.
DeVivo, M. T., D.R. Edmunds, M.J. Kauffman, B.A. Schumaker, J. Binfet, T.J. Kreeger, B.J.
Richards, H.M. Schatzl, and T.E. Cornish. 2017. Endemic chronic wasting disease causes
mule deer population decline in Wyoming. PLoS One 12:
https://doi.org/10.1371/journal.pone.0186512
Dufford, D. and P. McDonald. 2018. Illinois chronic wasting disease: 2017-2018 surveillance and
management report. Illinois Department of Natural Resources, Wildlife Disease
Program.
Edmunds, D. R., M.J. Kauffman, B.A. Schumaker, F.G. Lindzey, W.E. Cook, T.J. Kreeger, R.G.
Grogan, and T.E. Cornish. 2016. Chronic Wasting Disease drives population decline of
white-tailed deer. PLoS One 11: https://doi.org/10.1371/journal.pone.0161127.

29
Flook, D.R. 1970. Causes and implications of an observed sex differential in the survival of
wapiti. Canadian Wildlife Service Report Series 11: 1-71.
Grear, D. A., M.D. Samuel, J.A. Langenberg, and D. Keane. 2006. Demographic patterns and
harvest vulnerability of chronic wasting disease infected white-tailed deer in Wisconsin.
Journal of Wildlife Management 70: 546–553.
Grear, D. A., M.D. Samuel, K.T. Scribner, B.V. Weckworth, and J.A. Langenberg. 2010. Influence
of genetic relatedness and spatial proximity on chronic wasting disease infection among
female white-tailed deer. Journal of Applied Ecology 47: 532–540.
Gross, J. E., and M.W. Miller. 2001. Chronic wasting disease in mule deer: disease dynamics
and control. Journal of Wildlife Management 65: 205–215.

Gough, K. C., and B.C. Maddison. 2010. Prion transmission. Prion 4: 275-282.

Habib, T. J., E.H. Merrill, M.J. Pybus, and D.W. Coltman. 2011. Modeling landscape effects on
density–contact rate relationships of deer in eastern Alberta: Implications for chronic
wasting disease. Ecological Modeling 222: 2722–2732.
Hamir, A.N., T. Gidlewski, T.R. Spraker, J.M. Miller, L. Creekmore, M. Crocheck, T. Cline, I.
O’rourke. 2006. Preliminary observations of genetic susceptibility of elk (Cervus elaphus
nelson) to chronic wasting disease by experimental oral inoculation. Journal of
Veterinary Diagnostic Investigation 18: 110-114.
Houston, D.B. 1982. The northern Yellowstone elk: ecology and management. Macmillan, New
York, New York, USA.
Johnson, C. J., K.E. Phillips, P.T. Schramm, D. McKenzie, J.A. Aiken, and J.A. Pederson. 2006.
Prions adhere to soil minerals and remain infectious. PLoS Pathogens. 2(4): e32.
https://doi.org/10.1371/journal.ppat.0020032.
Joly, D.O., M.D. Samuel, J.A. Lagenberg, J.A. Blanchong, C.A. Batha, R.E. Rolley, D.P. Keane, and
C.A. Ribic. 2006. Spatial epidemiology of chronic wasting disease in Wisconsin white-
tailed deer. Journal of Wildlife Diseases. 42: 578-588.
Kittmas, W.H. 1953. Reproduction of Yellowstone Elk. Journal of Wildlife Management 17(2):
177-184.
Kramm, C., S. Pritzkow, A. Lyon, T. Nichols, R. Morales, and C. Soto. 2017. Detection of prion in
blood of cervids at the asymptomatic stage of chronic wasting disease. Science Reports.
7: 1-8.
Krumm C.E., M.M. Conner, M.W. Miller. 2005. Relative vulnerability of chronic wasting disease
infected mule deer to vehicle collisions. Journal of Wildlife Disease 41: 503–511.

30
Kretzschmar, H. A., S.B. Prusiner, L.E. Stowring, and S.J. DeArmond. 1986. Scrapie prion proteins
are synthesized in neurons. The American journal of pathology: 122(1): 1–5.
Kuznetsova, A., C. Cullingham, D. McKenzie, and J. M. Aiken. 2018. Soil humic acids degrade
CWD prions and reduce infectivity. PLoS Pathogens. 14(11): e1007414.
Loehle, C. 1995. Social barriers to pathogen transmission in free-ranging animal populations.
Ecology 76: 326–335.
Long, E. S., D.R. Diefenbach, C.S. Rosenberry, and B.D. Wallingford. 2008. Multiple proximate
and ultimate causes of natal dispersal in white-tailed deer. Behavioral Ecology. 19: 1235-
1242.
Long, E. S., D.R. Diefenbach, C.S. Rosenberry, and B.D. Wallingford. 2010. Influence of roads,
rivers, and mountains on natal dispersal of white-tailed deer. Journal of Wildlife
Management. 74: 1242-1249.
Lutz, C. L., D.R. Diefenbach, and C.S. Rosenberry. 2015. Population density influence dispersal in
female white-tailed deer. Journal of Mammalogy 96 (3): 494-501.
Lutz, C. L., D.R. Diefenbach, and C.S. Rosenberry. 2016. Proximate influences on female
dispersal in white-tailed deer. Journal of Wildlife Management. 80: 1218-1226.
Manjerovic, M.B., M.L. Green, N. Mateus-Pinilla, J. Novakofski. 2014. The importance of
localized culling in stabilizing chronic wasting disease prevalence in white-tailed deer
populations. Preventive Veterinary Medicine 113 (1): 139-145.
Mathiason, C. K., S.A. Hays, J. Powers, J. Hayes-Klug, J. Langenberg, J. Dahmes, D.A. Osborn, K.V.
Miller, R.J. Warren, G.L. Mason, and E.A. Hoover. 2009. Infectious prions in pre-clinical
deer and transmission of chronic wasting disease solely by environmental exposure.
PLoS ONE 4(6): e5916. https://doi.org/10.1371/journal.pone.0005916
Miller, M. W., and M. Conner. 2005. Epidemiology of CWD in free-ranging mule deer: spatial,
temporal, and demographic influences on observed prevalence patterns. Journal of
Wildlife Diseases. 41(2): 275-290.
Miller, M. W., and E. S. Williams. 2002. Detection of PrP (CWD) in mule deer by
immunohistochemistry of lymphoid tissues. Veterinary Record 151: 610-612.
Miller, M. W., and E.S. Williams. 2003. Horizontal prion transfer in mule deer. Nature 425: 35-
36.
Miller, M.W., E.S. Williams, N.T Hobbs, and L.L. Wolfe. 2004. Environmental Sources of Prion
transmission in mule deer. Emerging Infectious Diseases. 10(6): 1003-1006.

31
Miller, M.W., H.M. Swanson, L.L. Wolfe, F.G. Quartarone, S.L. Huwer, C.H. Southwick, P.M.
Lukacs. 2008. Lions and prions and deer demise. PLoS ONE 3(12): e4019.
https://doi.org/10.1371/journal.pone.000401.
Miller, W.L., and W.D. Walter. 2019. Spatial heterogeneity of prion gene polymorphisms in an
area recently infected by chronic wasting disease. Prion 13(1): 65-76.
Minnesota DNR. 2019. Chronic Wasting Disease Management.
https://www.dnr.state.mn.us/cwd/index.html. Accessed 11 July 2019.
Monello, R. J., J. G. Powers, N. T. Hobbs, T. R. Spraker, M. K. Watry, and Free-ranging, M. A.
2014. Survival and population growth of a free-ranging elk population with a long
history of exposure to chronic wasting disease. The Journal of Wildlife Management 78:
214-223.

Murray, M. H., Becker, D. J., Hall, R. J., and Hernandez, S. M. 2016. Wildlife health and
supplemental feeding: a review and management recommendations. Biological
Conservation. 204: 163-174.
Nathan, R. 2001. Dispersal biogeography. Pages 127-152 in S. A. Levin, editor. Encyclopedia of
biodiversity. Academic Press, San Diego, California, USA.
New York DEC. 2019. Status of CWD. https://www.dec.ny.gov/animals/33220.html. Accessed
11 July 2019.
Osnas, E. E., G. Wasserberg, R.E. Rolley, and M.D. Samuel. 2009. Host culling as an adaptive
management tool for chronic wasting disease in white-tailed deer: a modeling study.
Journal of Applied Ecology. 46(2): 457-466.
Prusiner, S. B. 1998. Prions. PNAS. 95(23): 13363-13383.
Rasmussen, J., B.H. Gilroyed, T. Reuter, S. Dudas, N.F. Neumann, A. Blachandran, N.N.V. Kav, C.
Graham, and T.A. McAllister. 2014. Can plants serve as a vector for prions causing
chronic wasting disease. Prion. 8(1): 136-142.
Race, B., K.D. Meade-White, K. Phillips, J. Striebel, R. Race, and B. Chesebro. 2014. Chronic
wasting disease agents in nonhuman primates. Emerging Infectious Diseases. 20(5): 833-
837.
Robinson, S. J., M.D. Samuel, K.I. O’Rourke, and C.J. Johnson. 2012. The role of genetics in
chronic wasting disease of North American cervids. Prion. 6(2): 153-162.
Rosatte, R. 2017. Home ranges and movements of elk (Cervus canadensis) restored to southern
Ontario, Canada. The Canadian field-naturalist 130(4): 320-331.

32
Schauber, E. M., D.J. Storm, and C.K. Nielsen. 2007. Effects of joint space use and group
membership on contact rates among white-tailed deer. Journal of Wildlife Management
71: 155–163.

Sigurdson, C.J. E.S. Williams, M.W. Miller, T.R. Spraker, K.I. O’Rourke, and E.A. Hoover. 1999.
Oral transmission and early lymphoid tropism of chronic wasting disease PrPres in mule
deer fawns (Odocoileus hemionus). Journal of General Virology 80: 2757–2764.
Spitznagel, E. 2012. Odd jobs: deer urine farmer. Bloomberg. August 31.
https://www.bloomberg.com/news/articles/2012-08-31/odd-jobs-deer-urine-farmer.
Spraker, T. R., A. Balachandran, D. Zhuang, and K.I. O’Rourke. 2004. Variable patterns of
distribution of PrP (CWD) in the obex and cranial lymphoid tissues of Rocky Mountain
elk (Cervus elaphus nelson) with subclinical chronic wasting disease. Veterinary Record
155(10): 295-302.
Strickland, B.K., S. Demarais. 2000. Age and regional differences in antlers and mass of white-
tailed deer. Journal of Wildlife Management 64(4): 903-911.
Tamgüney, G., M.W. Miller, L.L. Wolfe, T.M. Sirochman, D.V. Glidden, C. Palmer, A. Lemus, S.J.
DeArmond, and S.B. Prusiner. 2009. Asymptomatic deer excrete infectious prions in
faeces. Nature. 461: 529-532.
Tosa, M.I., E.M. Schauber, C.K. Nielsen. 2017. Localized removal affects white-tailed deer space
use and contacts. Journal of Wildlife Management 81(1): 26-37.
Uehlinger, F. D., A.C. Johnston, T.K. Bollinger, and C.L. Waldner. 2016. Systematic review of
management strategies to control chronic wasting disease in free-ranging deer
populations in North America. BMC Veterinary Research 12: 173.
USDA. 2013. Chronic wasting disease: revised program standards. USDA, Washington, DC.
USGS. 2019. Distribution of Chronic Wasting Disease in North America.
https://www.usgs.gov/media/images/distribution-chronic-wasting-disease-north-
america-0. Accessed 11 July 2019.
Vaske, J. J., N.R. Timmons, J. Beaman, and J. Petchenik. 2004. Chronic wasting disease in
Wisconsin: hunter behavior, perceived risk, and agency trust. Human dimensions of
wildlife 9(3): 193-209.
Waddell, L., J. Greig, M. Mascarenhas, A. Otten, T. Corrin, and K. Hierlihy. 2018. Current
evidence on the transmissibility of chronic wasting disease prions to humans— A
systematic review. Transboundary and Emerging Diseases 61(1): 37-49.
Walter, W. D., T.S. Evans, D Stainbrook, B.D. Wallingford. 2018. Heterogeneity of a landscape
influences size of home range in a North American cervid. Scientific reports 8(1): 14667.

33
Williams, D. M., A.C.D. Quinn, and W.F. Porter. 2014. Informing Disease Models with Temporal
and Spatial Contact Structure among GPS-Collared Individuals in Free-ranging
Populations. PLoS ONE 9(1): e84368.
Williams, E. S. 2005. Chronic wasting disease. Veterinary Pathology. 42(5): 530-549.
Williams, E. S., M.W. Miller, T.J. Kreeger, R.H. Kahn, and E.T. Thorne. 2002. Chronic wasting
disease of deer and elk: a review with recommendations for management. The Journal
of Wildlife Management. 66(3): 551-563.
Wisconsin Department of Natural Resources. 2016. A chronology of Wisconsin deer hunting:
from closed seasons to record harvests.
https://dnr.wi.gov/topic/hunt/documents/deer4page.pdf Accessed 2 August 2019.
Wisconsin Department of Natural Resources. 2018. Southwest Wisconsin CWD, Deer &
Predator Study Newsletter.
https://dnr.wi.gov/topic/research/articles/february2018.html#articleOne. Accessed 2
August 2019.
Wisconsin DNR. 2019. Prevalence and Surveillance.
http://dnr.wi.gov/topic/wildlifehabitat/prevalence.html Accessed 11 July 2019.

34
Appendix 1: Distance Sampling to Estimate Deer Abundance
The Pennsylvania Game Commission and the Pennsylvania Cooperative Fish and Wildlife
Research Unit (PACFWRU) at Penn State have studied different methods of estimating
abundance of white-tailed deer over the past 30 years. Methods vary in time and money
required and every method has assumptions that need to be met before it will provide accurate
estimates of abundance.

Distance sampling is a method of estimating deer abundance that has been used for many
species (Buckland et al. 2001) and is being used on current studies in Pennsylvania
(https://ecosystems.psu.edu/research/projects/deer/news/2018/distance-sampling). The
Game Commission and PACFWRU are studying the use of distance sampling from roads using
window-mounted Forward-Looking infrared (FLIR) units. The FLIR units increase the probability
of observing deer, but still are affected by the amount of vegetation and distribution of deer
(Figure 1).

The premise of distance sampling is that no matter how deer are distributed on the landscape,
if you randomly place transects across the landscape deer will be uniformly distributed with
respect to the transects. In other words, if you randomly place transects across the study area,
you will just as likely find a deer 10 feet from the transect as 200 feet from the transect.

Figure 1. Infrared images of white-tailed deer in Bald Eagle State Forest, May 2019, obtained
during a study to develop methods of using vehicle-based FLIR surveys to estimate deer
abundance. Detection probabilities of deer can be increased using FLIR, but the further a deer is
from a road the less likely it will be detected. Distance sampling methods can adjust for this
problem.

35
The problem, however, is that a deer 200 feet from the transect is less likely to be detected
(because of trees, topography, etc.) than a deer 10 feet from the transect. This problem is a
detection issue and distance sampling estimates a detection probability and adjusts the
observed count of deer to estimate the total number of deer on the study area. In the graph
below (Figure 2), the vertical bars represent the relative number of deer observed at different
distance intervals and the line represents the detection probability for a given distance from
the transect. Notice how detection probability is 1.0 (100%) on the transect line (distance = 0.00
km) and declines rapidly until no deer were observed beyond 60 meters from the transect line
(distance = 0.06 km).

Figure 2. Relative number of deer observed (bars) and the calculated detection probability (line)
of deer by distance (km) from the transect line.

To conduct a distance sampling survey, there are 4 pieces of information that are needed:
(1) the size of the study area (A),
(2) the length of transects surveyed (L)
(3) the perpendicular distances from the transect each deer group was detected, and
(4) the number of deer in each group detected.

The length of the transect and the greatest distance a deer was observed (w) is used to
calculate the area surveyed (2 × L × w = a) and the distances deer were detected are used to
estimate the probability of detecting a deer (p) within the area surveyed. If the total number of
deer observed is n, then an estimate of the number of deer in the study area (N) is:

𝑛𝑛
� = 𝐴𝐴
𝑁𝑁
𝑎𝑎𝑝𝑝̂

36
An important assumption of distance sampling cannot be met using roads to survey deer. This
assumption is violated because deer are less likely to be observed near roads (Figure 3).
Estimates of abundance can still be obtained, but they will underestimate deer abundance
(Marques et al. 2013).

Additional information on distance sampling can be found at http://distancesampling.org/

Figure 3. Locations of deer fitted with GPS collars relative roads on the chronic wasting disease
management area in south-central Pennsylvania, April 2018. Deer are less likely to be observed
within 50 meters of a road.

Literature Cited

Buckland, S.T., D. R. Anderson, K. P. Burnham, J. L. Laake, D. L. Borchers, and L. Thomas. 2001.

Introduction to distance sampling: estimating abundance of biological populations.
Oxford University Press, Oxford, U.K.

Marques, T. A., S. T. Buckland, R. Bispo, and B. Howland. 2013. Accounting for animal density
gradients using independent information in distance sampling surveys. Statistical
Methods and Applications 22:67-80.

Stainbrook, D. P. 2011. Methods of estimating white-tailed deer abundance at Gettysburg

National Military Park: testing assumptions of distance sampling. M.S. Thesis,
Pennsylvania State University, University Park, USA.

37