Managing wolves in the Yellowstone area: Balancing goals across jurisdictional boundaries: Managing Wolves Around Yellowstone

Abstract

Gray wolf (Canis lupus) restoration in the Greater Yellowstone Ecosystem began in 1995 with a small founder population in Yellowstone National Park, USA, which increased and contributed to a fully restored population in the northern Rocky Mountains by 2003. Upon removal as a federally listed, threatened species, wolf management outside the park was conveyed to Idaho, Montana, and Wyoming (USA) during 2009–2012; though wolves were relisted in Wyoming in 2014. Subsequent harvests elicited substantial negative reactions from wolf advocates when wolves that lived primarily in the park moved into surrounding states and were lawfully harvested. Conversely, many game managers and hunters advocated for larger harvests of wolves, including those in the park. These divergent viewpoints merit consideration when developing management plans for wolves that move between preserves and areas where hunting is permitted. We describe the history of wolf restoration and hunting in the ecosystem, contrast National Park Service and state management objectives, and characterize the risk to wolves living primarily in the park from hunting in surrounding areas. We recommend a framework for trans-boundary wolf management that considers population, social structure, and ecosystem objectives on public lands, potential influences of harvests on population growth, depredation risks to livestock, and opportunities for hunters and wildlife watching. We also consider human attitudes and social norms, thereby allowing for differences in values in our prescriptions. This framework has been implemented in a key hunting area along the northern boundary of Yellowstone and is broadly applicable elsewhere for the management of wildlife species held in the public trust.

Full text

Emerging Issues
Managing Wolves in the Yellowstone Area:
Balancing Goals Across Jurisdictional
Boundaries
DOUGLAS W. SMITH,
1
Yellowstone National Park, P.O. Box 168, Mammoth, WY 82190, USA
P. J. WHITE, Yellowstone National Park, P.O. Box 168, Mammoth, WY 82190, USA
DANIEL R. STAHLER, Yellowstone National Park, P.O. Box 168, Mammoth, WY 82190, USA
ADRIAN WYDEVEN,
2
Wisconsin Department of Natural Resources, 875 S 4th Avenue, Park Falls, WI 54552, USA
DAVID E. HALLAC, Cape Hatteras National Seashore, 1401 National Park Drive, Manteo, NC 27954, USA
ABSTRACT Gray wolf (Canis lupus) restoration in the Greater Yellowstone Ecosystem began in 1995 with a
small founder population in Yellowstone National Park, USA, which increased and contributed to a fully
restored population in the northern Rocky Mountains by 2003. Upon removal as a federally listed, threatened
species, wolf management outside the park was conveyed to Idaho, Montana, and Wyoming (USA) during
2009–2012; though wolves were relisted in Wyoming in 2014. Subsequent harvests elicited substantial
negative reactions from wolf advocates when wolves that lived primarily in the park moved into surrounding
states and were lawfully harvested. Conversely, many game managers and hunters advocated for larger
harvests of wolves, including those in the park. These divergent viewpoints merit consideration when
developing management plans for wolves that move between preserves and areas where hunting is permitted.
We describe the history of wolf restoration and hunting in the ecosystem, contrast National Park Service and
state management objectives, and characterize the risk to wolves living primarily in the park from hunting in
surrounding areas. We recommend a framework for trans-boundary wolf management that considers
population, social structure, and ecosystem objectives on public lands, potential influences of harvests on
population growth, depredation risks to livestock, and opportunities for hunters and wildlife watching. We
also consider human attitudes and social norms, thereby allowing for differences in values in our
prescriptions. This framework has been implemented in a key hunting area along the northern boundary of
Yellowstone and is broadly applicable elsewhere for the management of wildlife species held in the public
trust. Ó2016 The Wildlife Society.
KEY WORDS Canis lupus, jurisdictional boundaries, national parks, wolves.
Wolves (Canis lupus) from Canada were released in
Yellowstone National Park (YNP) and central Idaho,
USA, during 1995–1997 (Bangs and Fritts 1996, Phillips
and Smith 1996). These wolves increased in abundance and
distribution to such an extent that they were considered
biologically recovered in Idaho, Montana, and Wyoming
(USA) by 2003 and removed from the federal threatened and
endangered list under the Endangered Species Act by 2012
(USFWS 2011). As a result, management authority outside
of national parks, national wildlife refuges, and tribal lands
was turned over to the states. Each state instituted
management plans including hunting as the primary method
of managing wolf abundance and distribution—similar to
other wildlife species across North America (see www.
westerngraywolf.fws.gov). Wolves were hunted in 2009 and
2011–2013 in Idaho and Montana, and in 2012 and 2013 in
Wyoming. There was no hunting in 2010 and in Wyoming
after 2013 as a result of legal and regulatory actions related to
delisting. Hunting is prohibited in YNP (16 U.S.C. I, V §
26), but wolves that primarily live in YNP (locations inside
YNP ¼96%, n¼53 GPS-tagged wolves, 2001–2015) can be
hunted when they move outside the park.
Elk (Cervus canadensis) are the primary prey of wolves in
YNP, comprising >85% of kills in most portions of the park
(Metz et al. 2012). Many elk that use high-elevation
grasslands in YNP during summer migrate to lower
elevations outside the park that have less snow during
winter. For the most part, wolves do not follow elk migrating
from YNP because of their resident, territorial social
structure (Smith and Bangs 2009). However, wolf packs
living primarily in YNP may make seasonal movements
Received: 12 August 2015; Accepted: 30 April 2016
1
E-mail: doug_smith@nps.gov
2
Present address: Timber Wolf Alliance, Northland College, 1411 Ellis
Avenue, Ashland, WI 54806, USA
Wildlife Society Bulletin; DOI: 10.1002/wsb.677
Smith et al. Managing Wolves Around Yellowstone 1

outside the park based on prey distribution and vulnerability.
Initiation of these movements typically corresponds to the
autumn hunting season when elk are widely distributed and
in good physical condition, which causes wolves to make
more expansive movements in search of vulnerable prey
(Fig. 1). Also, remains from hunter-killed elk outside the
park create scavenging opportunities that attract wolves from
the park (Ruth et al. 2003).
Wolf restoration efforts have generated controversy
regarding the relative value of large carnivores as components
of natural ecosystems. A spectrum of opinions exists, ranging
from advocates for complete protection with no harvest to
advocates for no protection. Management of large carnivores,
such as wolves, has advanced from an era in which they were
actively limited to low numbers to one in which there is
increasing social tolerance and valuation of these animals as
part of wild ecosystems. Thus, there is increasing pressure on
wildlife agencies to balance the desires of hunters and
nonhunters (Schwartz et al. 2003). However, livestock
depredation and substantial decreases in numbers of elk that
migrate outside YNP during autumn and winter have also
increased pressure on state wildlife officials to limit wolf
numbers through hunting and trapping. In turn, park
officials have been asked to control numbers of wolves and
other wildlife in YNP. As a result, there is still vigorous
debate regarding the management of wolves in and near
YNP, including 1) the National Park Service (NPS)
philosophy of allowing wildlife within national parks to
fluctuate with minimal human intervention; 2) the increasing
value society places on wildlife for nonconsumptive purposes
(e.g., wildlife watching) compared with their traditional
values for sport and subsistence hunting; and 3) how to
balance the social value of predators with their effects on
domestic livestock production and wildlife species highly
valued for hunting. These topics are not mutually exclusive.
For example, preserves without hunting can serve as
sources of migratory or dispersing wildlife that benefit
nearby areas where hunting occurs, while hunting outside
preserves can serve to limit the abundance of wildlife that
may otherwise increase to high densities and cause
substantial changes to ecosystem processes and vegetation
communities before density-dependent regulatory processes
decrease their numbers.
Public opinion surveys indicate sport harvest and the
removal of wolves that kill livestock may increase social
tolerance for wolves in some cases, but not others (Mech
1995, Bangs et al. 2005, Treves and Martin 2011, Treves
et al. 2013). Wolf restoration occurred, in part, because
human attitudes changed and people supported their
restoration, with the prospect of hunting being a part of
this acceptance (Mech 1995, Clark et al. 1996, Fritts et al.
1997). Hunting is a key component of the North American
model of conservation that is used by all state wildlife
agencies and provides economic and social benefits (Leopold
1933, Arnett and Southwick 2015). This model is based on
the premise that wildlife are held in the public trust and
managed responsibly for wise use by people (Leopold 1933,
1949). Since 1968, the NPS has taken a somewhat different
approach known as ecosystem process management (or
natural regulation) in which population trajectories and
associations of native wildlife species are, to the extent
practicable, determined by nature with minimal intervention
by humans inside park units (Boyce 1998, Sinclair 1998,
Cole and Yung 2010, White et al. 2013a). More recent
management policies also contain provisions designed to
ensure the nonconsumptive enjoyment of wildlife by visitors
(National Park Service 2006). There is tremendous overlap
in these stewardship and naturalness paradigms, with both
approaches managing wildlife species as public trust
resources for the benefit of present and future generations
(The Wildlife Society 2007). Both paradigms also focus on
conserving habitats and ecological processes that sustain
viable populations of wildlife.
Despite these similarities, differences between the stew-
ardship and naturalness management approaches in terms of
consumptive and nonconsumptive uses and the extent of
human intervention have been a source of conflict between
national park units, state wildlife agencies, and other
stakeholders for decades (e.g., National Research Council
2002; Wagner 2006; White et al. 2013a,b). Furthermore,
there are vocal minorities that often highlight differences in
mandates and policies among the NPS and state wildlife
agencies, which tends to exacerbate differences rather than
provide support for a commonly held trust. The reintroduc-
tion of wolves into the Yellowstone area added to this
chronic, contentious debate (Smith and Bangs 2009). Our
objective was to summarize trans-boundary policy differ-
ences and develop a framework for sustainable and adaptable
interagency solutions that balance hunter harvest with the
conservation of watchable wildlife (Borg 2015), thereby
reducing conflicts about wolf management in the Yellow-
stone area, and possibly, other places where large carnivores
have been, or will be, restored.
Figure 1. Percent of Global Positioning System (GPS)-tagged wolf
locations by month outside Yellowstone National Park (YNP), WY, USA,
for 53 wolves from January 2001 to March 2015. This includes only GPS
locations from wolves living in packs (excludes loners). Sample sizes (no. of
wolves sampled with 5 locations/month) are above each bar. “Backcountry”
and “Rifle” bars denote adjacent backcountry wilderness and general rifle-
hunting seasons for elk, respectively. Error bars represent the standard error.
2 Wildlife Society Bulletin 9999()

Contrasting Wildlife Management Policies
The mission of state wildlife agencies is to provide for the
stewardship of wildlife resources for present and future
generations of people. State policies generally emphasize
the wise use and consumptive benefits of wildlife, while also
providing for a broad array of nonconsumptive uses for the
public. Public fishing, hunting, and trapping are used to
regulate wildlife populations of species valued for food, fur,
or other purposes at sustainable levels that provide
ecological and recreational benefits, while also reducing
conflicts with people. Most state wildlife agencies depend
on revenue generated by sales of licenses to fish and hunt,
leveraged with money from federal excise taxes on the sale
of sporting equipment such as ammunition and firearms.
Many states also have nongame programs funded by
voluntary contributions or sales of vanity license plates and
educational materials, but these funds are typically minimal
compared with those generated by license fees (J. Hammill,
Michigan Department of Natural Resources, personal
communication).
The mission of the NPS has a somewhat different focus
that emphasizes the preservation of native wildlife resources
and the processes that sustain them. Hunting is prohibited in
most parks, including YNP (544 U.S.C. 100101(a), 100301
et seq.). In other words, nonconsumptive benefits and
enjoyment by visitors are emphasized over consumptive uses
(National Park Service 2006). Wildlife can be intensively
managed when necessary, but the current management
approach, which evolved over time, emphasizes minimizing
human intervention, maintaining ecological integrity and
resiliency, and maintaining natural disturbances and dynam-
ics (Leopold et al. 1963; National Park Service 2006; White
et al. 2013a,b). Thus, parks can sometimes serve as ecological
baselines or benchmarks for assessing the effects of more
intense human activities in surrounding areas (Leopold et al.
1963, Sinclair 1998, National Park Service Advisory Board
Science Committee 2012).
Wolf Recovery and Management in the Northern
Rockies
Predator control was conducted in YNP through the 1930s,
which eliminated or greatly reduced the abundance of most
large carnivores (Schullery and Whittlesey 1992). However,
predators began increasing in numbers and distribution during
the late 1900s because of changes in human attitudes and
modern wildlife management practices, both inside and
outside of preserves such as YNP. Currently, YNP is more
predator-rich than at any time in the park’s history because all
of the large carnivores have been restored and exist at densities
commensurate with available prey and administrative pro-
tections (Olliff et al. 2013, White et al. 2013b).This recovery is
not without issues because large carnivores prey on livestock,
compete with human hunters for ungulates, and, rarely, attack
humans (McNay 2002, Bradley and Pletscher 2005, Unger
2008). Thus, this recovery is celebrated by some and detested
by others (Mech 1995, Smith and Bangs 2009).
Wolf recovery to Idaho, Montana, and Wyoming was
guided by the Northern Rocky Mountain Wolf Recovery
Plan, which established a goal of 3 populations of 100 wolves
with some level of genetic connectivity, or alternatively, 10
breeding pairs with 2 pups at year’s end in each of the 3
recovery areas for 3 successive years (USFWS 1987). This
objective was achieved in 2003 with 761 wolves and 51
breeding pairs across the northern Rocky Mountains
(USFWS 2006). However, delisting was delayed by
litigation over what constituted wolf recovery and acceptable
state management plans. Resolution of these issues was
achieved in 2009 for Montana and Idaho, and these states
initiated wolf hunting that autumn. Wolves were relisted in
2010 because of litigation, and public hunting was closed. In
2011, wolves were again delisted in Idaho and Montana, and
hunting occurred in these states. In 2012, wolves were also
delisted in Wyoming, and public hunting occurred in all 3
states during 2012 and 2013. However, wolves were relisted
in Wyoming in September 2014.
Prior to delisting, each state developed a management plan
with a minimum population objective of 15 breeding pairs or
approximately 150 wolves, which exceeds the recovery
requirements (see www.westerngraywolf.fws.gov for state
wolf plans). Each plan also contained provisions to manage
for connectivity among the 3 populations in the northern
Rocky Mountains and use public hunting as the primary tool
to adjust numbers (vonHoldt et al. 2010). Evaluating the
success of management by the states required assigning wolf
packs to a state based on where each pack denned and what
proportion of its territory occurred (usually >50%) within a
particular state. Wolves living primarily in YNP are counted
by Wyoming (Fig. 2).
During the period of federal oversight provided by the
Endangered Species Act, as well as litigation (1995–2009)
arising from interpretations of this Act, the wolf population in
the northern Rocky Mountains grew from approximately
103–113 wolves in 1995 to approximately 1,700 wolves in
2009 (USFWS 2011). Management primarily focused on
removing livestock-killing wolves, with 2,100 wolves removed
during control actions during 1995–2013 (USFWS et al.
2014). There was no public hunting until wolf delisting in the
years mentioned previously. In YNP, numbers increased
rapidly to 174 wolves in 14 packs by 2003, but decreased to 99
wolves in 10 packs by 2016 primarily because of intraspecific
aggression, disease, and possibly, food stress (Almberg et al.
2010, Cubaynes et al. 2014, Smith et al. 2014).
Human-Tolerant Wolves in YNP
One unintentional outcome of wolf restoration in YNP is
that some wolves become habituated to people after frequent,
nonthreatening encounters with visitors along road corri-
dors. Greater than 3 million visitors each year seek wildlife
viewing experiences in YNP. The park is one of the best
places in the world to view wild wolves because of their
visibility and tolerance of humans (Smith and Bangs 2009).
Wolves are now one of the primary reasons people visit YNP
and wolf watchers contribute approximately US$35 million
annually to local economies (Duffield et al. 2008). Wildlife-
viewing companies, educational associations, and loosely
organized groups of wolf watchers track and communicate
Smith et al. Managing Wolves Around Yellowstone 3

wolf viewing opportunities, so that when a wolf is sighted
tens to hundreds of people quickly assemble to view it. This
regular exposure to people has produced some wolves that
have become habituated and 2 wolves had to be removed.
This tolerance to humans likely makes them more vulnerable
to hunting, although some question this (Borg 2015).
Another inadvertent effect of the visibility and tolerance of
wolves in YNP was the attention and celebrity status given to
certain animals or packs by advocates, which led to the
naming and anthropomorphization or personification of
many animals. Although this tendency is common in a zoo
environment, it creates unrealistic expectations and issues for
managers of wild animals tasked with sustaining viable
populations rather than protecting individuals. Park biolo-
gists and interpreters have been discouraging this practice,
but with limited success (Reid 2015).
Research on the Effects of Wolf Restoration
Given long-standing debates about wolf–prey dynamics
and use of wolf control to increase prey populations, a
fundamental objective of research in the Yellowstone area
has been to assess the effects of wolves on elk (National
Research Council 1997, Hayes et al. 2003). Elk monitoring
in YNP and the surrounding area was initiated long before
wolf reintroduction; intensive efforts to monitor radio-
tagged elk to evaluate the effects of wolf restoration have
persisted for >18 years (Houston 1982, Taper and Gogan
2002, Garrott et al. 2009, Hebblewhite and Smith 2010,
White and Garrott 2013). Several National Science
Foundation grants have supported this research, in part,
with collaboration by scientists from numerous federal and
state agencies, universities, and nongovernmental orga-
nizations.
Figure 2. Distribution of wolf pack territories in Yellowstone National Park, WY, USA, in 2014. Polygons were created using >30 locations/pack and
represent 95% of the locations for each pack so does not represent most of the movements made outside the park. One territory is approximated by a circle due to
no radio collar data locations available to create a polygon.
4 Wildlife Society Bulletin 9999()

To facilitate this monitoring and research, approximately
25–30% of the adult wolves in YNP are fitted with
radiotransmitters. Telemetry enables biologists to maintain
contact with each pack and facilitate the collection of
information on abundance, behavior, demographics, disease,
distribution, genetics, livestock depredation, and predator–
prey relationships (Smith et al. 2014). This information is
shared with, and benefits, multiple federal and state agencies,
universities, and stakeholders (e.g., see www.westerngraywolf.
fws.gov). As a result, radio-tagged wolves are an extremely
valuable resource. In addition, monitoring radio-tagged
wolves across jurisdictions and management paradigms is
extremely important because the effects of wolves on elk
population dynamics vary considerably across the region on
account of differences in elk migratory patterns, habitat,
human harvests, land use, local weather, and predator densities
and management (Garrott et al. 2005, Hamlin et al. 2009).
Wolf Hunting in the Yellowstone Area, 2009–2015
In 2009, 4 wolves that primarily lived in YNP (99% of
locations) were harvested 1–5 km north in Montana
(Fig. 3). These wolves constituted 3% of the 137 wolves
living primarily in YNP, and as a result, their deaths had no
effect on the growth rate of the population. However, the
harvested wolves were all from the Cottonwood Creek pack,
including the breeding pair and 2 wolves fitted with
radiotransmitters. Also, the remaining 6 pack members
were never located again and their vacated territory was
usurped by existing, adjacent packs (Smith et al. 2010). As a
result, these harvests generated considerable negative media
coverage nation-wide.
There were no wolf-hunting seasons in states surrounding
YNP during 2010 as a result of litigation and wolf relisting.
However, the Montana Fish and Wildlife Commission
reduced the quota from 15 to 3 wolves north of the park
where the Cottonwood Creek wolves were harvested.
During 2011, only 2 wolves (1 radio-tagged) from YNP
were shot close to the park boundary in Montana (Fig. 3), but
there was little media attention; this was likely due to the
small number harvested.
In 2012, 12 wolves that primarily lived within YNP were
harvested in Idaho (2), Montana (7), and Wyoming (3;
Fig. 3). These wolves constituted 12% of the 98 wolves living
primarily in YNP, but their removal had little to no apparent
effect on the population growth rate (Smith et al. 2014).
However, 7 of 10 packs in YNP had 1 wolves removed
during this harvest. Also, 6 of the harvested wolves were
fitted with radiotransmitters, the loss of which hampered
federal and state biologists from tracking packs for
management and scientific purposes. Furthermore, 2 of
the harvested wolves from the Lamar Canyon pack were
recognizable and popular among park visitors. Even though
these wolves were harvested >24 km east of the park
boundary in Wyoming, their deaths generated considerable
negative, world-wide, media attention and thousands of
letters, electronic mail messages, and phone calls from angry
people (e.g., Schweber 2012). In turn, some individuals and
groups ceased their financial support to donor organizations
Figure 3. Harvest locations of wolves that primarily lived in Yellowstone National Park (YNP), WY, USA, during 2009–2015, but ventured outside the
boundary during hunting seasons in the states of Idaho, Montana, and Wyoming. Each circle represents the location of a harvested wolf, whereas different
colors represent the years they were harvested. The wolf hunting season was closed in 2010 and no known wolves from YNP were harvested during 2013.
Smith et al. Managing Wolves Around Yellowstone 5

that support management and research within and near
YNP, and the National Science Foundation questioned the
validity of ongoing wolf–elk research they were financially
supporting given the level of harvest of wolves living
primarily in YNP. In contrast, the states received positive
feedback for the hunting of wolves following this negative
publicity.
No wolves that primarily used YNP were harvested in the
surrounding states during the 2013–2014 hunting season,
while 5 wolves (2 tagged with radiotransmitters) from YNP
were harvested north in Montana during the 2014–2015
season (Fig. 3). These harvests again generated outcry from
wolf advocates and counter-responses from hunters and
other stakeholders. In response, the Montana Fish and
Wildlife Commission reduced the number of wolves that can
be killed north of YNP from 3 to 2 for the 2015–2016 season.
This brief history indicates the Montana Fish and Wildlife
Commission responded positively to stakeholders’ requests
to consider nonconsumptive values placed on wolves in the
region by substantially reducing quotas in these areas; this
management action helped to alleviate future conflicts.
However, the potential still exists for high harvests of wolves
exiting the park in other areas, where wolf harvest quotas are
substantially higher or unlimited, including in Idaho and
Wyoming. This probability may be heightened by the
habituation of wolves to visitors in YNP, their likely naivete
when they initially move outside the park, and current
temporal patterns of increased movement outside the park
that coincide with the initiation of wolf hunting season
(Fig. 1). To garner ideas for resolving these issues, we
investigated how federal and state management agencies in
other areas have dealt with trans-boundary wolf movements
between preserves and areas with hunting.
Trans-Boundary Management Paradigms Employed
Elsewhere
Other parks where the cross-boundary movements of wolves
are common include Denali National Park in Alaska, USA;
Algonquin Provincial Park in Ontario, Canada; Kluane
National Park in the YukonTerritory, Canada; Banff National
Park in Alberta, Canada; and Riding Mountain National Park
in Manitoba, Canada. In each situation, different approaches
and rationales for trans-boundary management occur, with no
universally applicable solution and controversy in each
situation.
A 233-km
2
area northeastof Denali National Park (called the
“wolf townships”) was recently reopened after being closed to
wolf hunting and trapping from 2000 to 2010 (Borg 2015).
The closure near the park was implemented because the most
visible wolf packs along the road in summer used a small area
outside the park in winter, where they were subject to harvest.
Reopening this area to harvest sparked considerable contro-
versy because management went from some protection of
wolves that primarily used the park, but migrated out
occasionally, to no protection outside the park (Borg 2015).
Denali is a large park (24,280 km
2
), with a large portion of the
wolf population protected inside the park for most of the year.
In 2012, following removal of the hunting closure, a breeding
female of a commonly viewed pack was harvested, which
caused the pack not to den. An index of wolf sightings by
visitors decreased by approximately 40%, which adversely
affected the park goal for visitor enjoyment (Borg 2015). Two
more wolves that primarily used the park were harvested in this
area in 2015; one was a pregnant female, which again led to
reproductive failure and no denning of the well-known pack.
The following year, thewolf sighting index for visitors dropped
again (B. Borg, Denali National Park and Preserve, personal
communication).
In Algonquin Provincial Park, 80–90% of the wolf packs
using the eastern portion of the park followed migrating
white-tailed deer (Odocoileus virginianus) out of the park,
which subjected them to harvest and contributed to a
decreasing wolf population. There was also a decrease in
within-pack kinship, indicating deviations from natural
social and genetic structure found in wolves (Forbes and
Theberge 1996, Grewal et al. 2004, Theberge and Theberge
2004). Therefore, a 10-km buffer was established around the
park, within which wolves could not be harvested (Theberge
and Theberge 2004). This buffer was effective in preventing
the likely extirpation of wolves in the park given the trend in
human-caused mortality rates (56–66%; Forbes and The-
berge 1996), with an initial effect of increasing wolf densities.
This was followed by a stabilization of wolf density as
human-caused mortality was offset by natural causes, as well
as an increase in within-pack genetic kinship and restoration
of natural social structure (Rutledge et al. 2010).
Similarly, in Kluane National Park wolves that dennedin and
used the park more than one-half the time were protected by a
buffer zone outside the park (Carey et al. 1994). This decision
was intended to minimize human influence on wolves in the
park (Carey et al. 1994). Management included radio-tagging
wolves to better understand their movements, which led to a
clearer definition of what constituted a pack that lived in the
park and creation of a buffer outside the park to protect them
(Carey et al. 1994). This buffer was effective in reducing
harvest on park wolves while wolf control was occurring
outside of the park.
In Banff National Park, no closures or reduced harvest
levels were implemented for wolves adjacent to the park
boundary because the park wolves were considered secure
and a source population (Thiessen 2007). Banff is a relatively
small park (9,000 km
2
), and as a result, there is frequent wolf
movement across the boundary. In turn, wolves that live
primarily in the park are frequently harvested close to the
boundary (Callaghan 2002, Hebblewhite 2006). To date,
this mortality has not noticeably affected population growth
(Thiessen 2007).
Riding Mountain National Park is also a small park
(2,974 km
2
), but unlike Banff is surrounded mostly by
agricultural land and supports approximately 70–75 wolves
(Stronen et al. 2007). Until 1980, wolves were considered
predators outside of the park and reductions under the
Predator Control Act, which sometimes removed entire park
packs, negatively affected wolf numbers inside the park (L.
Carbyn, University of Alberta, personal communication).
Thereafter, wolves were classified as big game under revisions
6 Wildlife Society Bulletin 9999()

to the Manitoba Provincial Wildlife Act; in 2001, wolf
hunting was closed in areas that surround the park
(3,200 km
2
area of variable width surrounding entire park;
Stronen et al. 2007), which likely lessened effects to park and
dispersing wolves. This hunting ban was recently overturned,
but it is too soon to know the effects of this policy change.
However, dispersal into the park is probably reduced (A.
Stronen, Aalborg University, personal communication).
Framework for Trans-Boundary Wolf Management in
the Yellowstone Area
We recommend a model for the management of wolves
transitioning (i.e., moving) among areas containing clusters
of relatively unexploited wolf packs to areas with harvest
goals that consider federal and state ecosystem and
population objectives. Recognizing and coalescing the
somewhat different missions and management approaches
used by the NPS and wildlife agencies in surrounding states
are important first steps in developing effective, coordinated
management of wolves in the region. Public hunting will
certainly be a key tool used to manage wolves in the northern
Rocky Mountains, with the North American model of
wildlife conservation serving as the guiding philosophy for
achieving scientifically based population objectives. Also,
there is a limit to tolerance for wolves in human-dominated
landscapes of this region given social and economic concerns
related to livestock depredation and ungulate population
levels for sport hunting, which is a major source of seasonal
revenue and wildlife agency support (du Toit et al. 2004,
Gordon et al. 2004).
The NPS goals of preserving natural systems and visitor
enjoyment do not preclude harvest of wildlife outside of parks
and, in fact, ungulates that migrate outside YNP during
autumn and winter have been harvested in a responsible and
sustainable manner for decades. In the past, Montana Fish,
Wildlife and Parks developed and refined regulations that
precluded overharvest or unethical hunting practices for bison
(Bison bison) and elk near the park boundary, and as a result,
mitigated public objections and negative effects to tourism and
local economies (Lemke et al. 1998, Bidwell 2010). Also, the
Montana Fish, Wildlife & Parks Commission created the
relatively small Wolf Management Unit 313 with a modest
quota to limit harvests in an intensely hunted area along the
northern park boundary. We suggest similar regulations could
be implemented in other key locations to allow wolves that
primarily live in YNP, but sometimes move outside the park
during hunting seasons, to transition from a protected to a
hunted environment without being exposed to liberal harvests
near the park boundary. The area of conservative harvests near
the park boundary could be defined usingpast radiolocations of
wolves in vulnerable packs, while also giving consideration to
socioeconomic and human dimensions factors. Lower harvests
in this transition zone would conserve wolves so that they are
available for public viewing and maintain naturalness and wolf
social structure. Such a plan would not significantly affect
hunting opportunities, the control of livestock depredation, or
the ability of state wildlife agencies to regulate wolf numbers
elsewhere.
Numerous studies have reported that harvest rates between
15% and 48% did not suppress wolf population growth
(Fuller et al. 2003, Adams et al. 2008, Creel and Rotella
2010, Gude et al. 2012). One possible explanation for this
finding is that the numerical response of wolves to harvest
depends on whether they are colonizing or have saturated an
area. In areas where wolf numbers are well below carrying
capacity, a larger portion of the harvest may be additive
because wolves will generally be in good physical condition,
with high rates of reproduction and survival, and would not
typically die from starvation (Kie et al. 2003, Murray et al.
2010). Where wolf numbers are near the (food or social)
carrying capacity of an area to support them, however, a
larger portion of the harvest will likely be compensatory, or a
substitute for other forms of mortality (e.g., starvation),
because there are not enough resources or capacity in the
environment to sustain all the wolves (Fuller et al. 2003). The
number of wolves in YNP appears to be near saturation,
which would allow for some compensation due to harvest
(Cubaynes et al. 2014).
However, numerical assessments of the effects of harvest
(or other types of mortality) on wolves do not consider
possible effects to their social structure. The death of a wolf
of high social rank will likely cause more disruption to a pack
than the death of a low-ranking wolf (Brainerd et al. 2008,
Rutledge et al. 2010). Thus, the harvest of a breeding female
could disrupt reproduction in a pack that year (Stahler et al.
2013), whereas the harvest of a mature male could affect the
pack’s hunting ability or competitive interactions with other
packs (MacNulty et al. 2009, Cassidy et al. 2015). Social
openings in wolf packs created by any cause are often readily
filled by dispersing wolves, but some of this depends on
timing and density (Fuller et al. 2003). Therefore, avoiding
some social disruption of wolf packs due to harvest is difficult
because removing one or more high-ranking wolves could
have significant effects for some unknown length of time,
which could include the disbanding of the pack (e.g.,
Cottonwood Creek pack during 2009; Lamar Canyon pack
during 2012). In turn, viewing opportunities for visitors to
YNP are diminished because a predictable resident pack is
eliminated (Borg et al. 2015). Reducing the harvest rate of
wolves near the boundary of YNP would reduce the
likelihood of social disruption to wolf packs, as well as
effects to high-ranking wolves that are recognizable and
well-known to the public.
Managing wolves with conservative harvests close to the
boundary of YNP and liberal harvests farther from the park
could lessen much of the controversy surrounding the harvest
of wolves that primarily live in the park. It is not possible for
this transition zone to be inside YNP because Congress
prohibited hunting therein (16 USC 26). Even if hunting
were allowed, however, such a proposal would likely remove
many breeding pairs from the northern and southern
portions of the park—thereby jeopardizing the number of
pairs assumed in the recovery plan (USFWS 1987). For
example, in 2014, there were 11 packs that substantially used
portions of YNP. If a 16-km (10-mile) transition zone with
conservative hunting were established inside the park
Smith et al. Managing Wolves Around Yellowstone 7

boundary, then 9 of these packs would have been exposed,
including 7 packs with almost their entire ranges within the
hunting area. Thus, the number of breeding pairs of wolves
in Wyoming could be substantially reduced below the level
assumed in the recovery plan (USFWS 1987).
Furthermore, there is value in having refuges from hunting
across regions where wolf populations need to be regulated.
The current levels of wolf harvest in the 3 recovery areas of
the northern Rockies represent case studies with alternative
forms of wolf population management (i.e., regulation). For
comparison, there also should be control or reference areas
where wolves are not harvested. In addition, refuges from
hunting could help prevent overharvest within regions of
high quality wolf habitat, while maintaining areas where
more normal pack dynamics and ecological effects are
expected. In areas with few parks or preserves, this concept
could be implemented in a zone management framework,
ranging from no or minimal harvest to areas with flexible
controls, while recognizing that culling may at times need to
occur in protected areas and restraint may sometimes be
necessary in liberal harvest areas.
Given these considerations, we propose that no >5–7% of
the prehunt number of wolves living primarily in YNP be
harvested each year. In addition, we recommend harvests
remove no >20% of the wolves in any given pack and no
>15% of the radio-tagged wolves living in YNP. This
harvest rate is the level recommended to preserve social
structure and pack size and minimize behavioral disruption
(Brainerd et al. 2008, Rutledge et al. 2010, Borg et al. 2015).
Such a harvest rate would sustain the long-term demo-
graphics of wolves living in YNP, minimize effects to wolf
sociality, preserve research and public viewing opportunities,
allow wolves to behaviorally transition from NPS to state
management paradigms, and not unduly affect regional
population management objectives for wolves. A flexible,
closely monitored approach to harvest management through
the hunting season would be necessary should any of these
benchmarks be exceeded (e.g., emergency season closures).
The use of a 24-hour closure technique is currently
implemented for some ungulate species (e.g., bighorn sheep
[Ovis canadensis]) once harvest quotas are reached; and, thus
far, park and state managers have detailed enough
information to regulate such a hunt (Canfield 2014, Smith
et al. 2014). In addition, we recommend wildlife managers
surrounding preserves implement quotas in small, but
flexible management units so that management agility is
provided to help avoid excessive harvest on one particular
pack or areas of wildlife viewing importance inside preserves.
Smaller units would provide the spatial framework for
intermediate management paradigms, versus having no
quotas and large hunting units that can result in unpredict-
able and undesirable effects.
A policy of this nature is flexible (based on proportions, not
fixed amounts), meets the management objectives of the
NPS and state wildlife agencies, is defensible because it is
based on scientific research and public input, and would
reduce the current level of conflict about appropriate wolf
management near the world’s most famous and scrutinized
national park. This approach to management focuses on
the population sustainability of wolves, rather than
personal values and the survival of individual animals,
which will result in a better outcome for wolves and people.
By incorporating human-dimensions thinking into the
decision-making process, this approach can serve as a
template for similar situations in different political climates.
Thus, it is a way to strengthen and modernize the North
American model of wildlife conservation (J. Hammill,
Michigan Department of Natural Resources, personal
communication). Similar challenges exist worldwide where
protected preserves and national parks are critical to
effectively conserving other large carnivore species (e.g.,
African lions [Panthera leo]; leopards [P. pardus]), but where
adjacent trophy hunting, livestock protection, and conflict
resolution are management elements that can threaten
species conservation (Woodroffe and Ginsberg 1998,
Loveridge et al. 2007, Balme et al. 2010). Consequently,
understanding the effects of various management actions and
stakeholder values on species populations in and around
national parks benefits many, and can guide implementation
of conservation management worldwide.
ACKNOWLEDGMENTS
We thank the Associate Editor, reviewers, R. Garrott
(Montana State University), J. Hammill (Michigan Depart-
ment of Natural Resources), and wildlife staff from the states
of Idaho, Montana, and Wyoming for their insightful
comments and recommendations on previous versions of this
manuscript. We also thank B. Borg (Denali National Park),
L. Carbyn (University of Alberta), A. Stronen (Aalborg
University), P. Paquet, and R. Hayes (retired Yukon
government) for review of some sections and conversations
about wolf management in their respective areas. We also
appreciate help with figures from K. Cassidy M. Metz, and
E. Stahler. Wolf monitoring and management in Yellow-
stone National Park is supported by the National Park
Service and the Yellowstone Park Foundation. We would
like to especially thank an anonymous donor, the Perkin-
Protho Foundation, and the Tapeats Foundation for funding
this research.
LITERATURE CITED
Adams, L. G, R. O. Stephenson, B. W. Dale, R. T. Ahgook, and D. J.
Demma. 2008. Population dynamics and harvest characteristics of wolves
in the central Brooks Range, Alaska. Wildlife Monographs 170.
Almberg, E., P. Cross, and D. Smith. 2010. Persistence of canine distemper
virus in the Greater Yellowstone Ecosystem’s carnivore community.
Ecological Applications 20:2058–2074.
Arnett, E. B., and R. Southwick. 2015. Economic and social benefits of
hunting in North America. International Journal of Environmental Studies
72:734–745.
Balme, G. A., R. Slotow, and L. T. B. Hunter. 2010. Edge effects and the
impact of non-protected areas in carnivore conservation: leopards in the
Phinda-Mkhuze Complex, SouthAfrica. Animal Conservation13:315–323.
Bangs, E. E., J. A. Fontaine, M. D. Jimenez, T. J. Meier, E. H. Bradley,
C. C. Niemeyer, D. W. Smith, C. M. Mack, V. Asher, and J. K. Oakleaf.
2005. Managing wolf–human conflict in the northwestern United States.
Pages 340–356 in R. Woodroffe, S. Thirgood, and A. Rabinowitz, editors.
People and wildlife: conflict or coexistence? Cambridge University Press,
New York, New York, USA.
8 Wildlife Society Bulletin 9999()

Bangs, E. E., and S. H. Fritts. 1996. Reintroducing the gray wolf to central
Idaho and Yellowstone National Park. Wildlife Society Bulletin
24:402–413.
Bidwell, D. 2010. Bison, boundaries, and brucellosis: risk perception and
political ecology at Yellowstone. Society and Natural Resources 23:14–30.
Borg, B. L. 2015. Effects of harvest on wolf social structure, population
dynamics and viewing opportunities in National Parks. Dissertation,
University of Alaska, Fairbanks, USA.
Borg, B. L. S. M. Brainerd, T. J. Meier, and L. R. Prugh. 2015. Impacts of
breeder loss on social structure, reproduction and population growth in a
social canid. Journal of Animal Ecology 84:177–187.
Boyce, M. S. 1998. Ecological-process management and ungulates: Yellow-
stone’s conservation paradigm. Wildlife Society Bulletin 26:391–398.
Bradley, E. H., and D. H. Pletscher. 2005. Assessing factors related to wolf
depredation of cattle in fenced pastures in Montana and Idaho. Wildlife
Society Bulletin 33:1256–1265.
Brainerd, S. M., H. Andren, E. E. Bangs, E. H. Bradley, J. A. Fontaine, W.
Hall, Y. Iliopoulos, M. D. Jimenez, E. A. Jozwiak, O. Liberg, C. M.
Mack, T. J. Meier, C. C. Niemeyer, H. C. Pedersen, H. Sand, R. N.
Schultz, D. W. Smith, P. Wabakken, and A. P. Wydeven. 2008. The
effects of breeder loss on wolves. Journal of Wildlife Management
72:89–98.
Callaghan, C. J. 2002. The ecology of gray wolf (Canis lupus) habitat use,
survival, and persistence in the central Rocky Mountains of Canada.
Dissertation, University of Guelph, Ontario, Canada.
Canfield, J. 2014. Northern Yellowstone cooperative wildlife working group
2013 annual report (October 1, 2012–September 30, 2013). Montana
Fish, Wildlife and Parks, National Park Service—Yellowstone National
Park, U.S. Forest Service—Gallatin National Forest, and U.S. Geological
Survey—Northern Rocky Mountain Science Center.
Carey, J., R. Hayes, R. Farnell, R. Ward, and A. Baer. 1994. Aishihik and
Kluane caribou recovery program: November 1992 to October 1993.
Yukon Fish and Wildlife Branch Report PR-94-2. Yukon Government,
Whitehorse, Canada.
Cassidy, K. A., D. R. MacNulty, D. R. Stahler, D. W. Smith, and L. D.
Mech. 2015. Group composition effects on aggressive inter-pack
interactions of gray wolves in Yellowstone National Park. Behavioral
Ecology 26:1352–1360.
Clark, S., P. C. Paquet, and A. P. Curlee. 1996. General lessons and positive
trends in large carnivore conservation. Conservation Biology
10:1055–1058.
Cole, D. N., and L. Yung, editors. 2010. Beyond naturalness: rethinking
park and wilderness stewardship in an era of rapid change. Island Press,
Washington, D.C., USA.
Creel, S., and J. J. Rotella. 2010. Meta-analysis of relationships between
human offtake, total mortality and population dynamics of gray wolves
(Canis lupus). PLoS ONE 5:e12918.
Cubaynes, S., D. R. MacNulty, D. R. Stahler, K. A. Quimby, D. W. Smith,
and T. Coulson. 2014. Density-dependent intraspecific aggression
regulates survival in northern Yellowstone wolves (Canis lupus). Journal
of Animal Ecology 83:1344–1356.
Duffield, J. W., C. J. Neher, and D. A. Patterson. 2008. Wolf recovery in
Yellowstone: park visitor attitudes, expenditures and economic impacts.
Yellowstone Science 16:21–25.
du Toit, J. T., B. H. Walker, and B. M. Campbell. 2004. Conserving tropical
nature: current challenges for ecologists. Trends in Ecology and Evolution
19:12–17
Forbes, G. J., and J. B. Theberge. 1996. Cross-boundary management of
Algonquin Park wolves. Conservation Biology 10:1091–1097.
Fritts, S. H., E. E. Bangs, J. A. Fontaine, M. R. Johnson, M. K. Phillips,
E. D. Koch, and J. R. Gunson. 1997. Planning and implementing
reintroduction of wolves to Yellowstone National Park and central Idaho.
Restoration Ecology 5:7–27.
Fuller, T., L. D. Mech, and J. Fitts Cochrane. 2003. Wolf population
dynamics. Pages 161–191 in L. D. Mech and L. Boitani, editors. Wolves:
behavior, ecology and conservation. University of Chicago Press, Chicago,
Illinois, USA.
Garrott, R. A., J. A. Gude, E. J. Bergman, C. Gower, P. J. White, and K. L.
Hamlin. 2005. Generalizing wolf effects across the greater Yellowstone
area: a cautionary note. Wildlife Society Bulletin 33:1245–1255.
Garrott, R. A., P. J. White, and J. R. Rotella. 2009. The Madison
headwaters elk herd: transitioning from bottom-up regulation to top-
down limitation. Pages 489–517 in R. A. Garrott, P. J. White, and F. G.
R. Watson, editors. The ecology of large mammals in central Yellowstone:
sixteen years of integrated field studies. Elsevier, San Diego, California,
USA.
Gordon, I. J., A. J. Hester, and M. Festa-Bianchet. 2004. The management
of wild large herbivores to meet economic, conservation and environmen-
tal objectives. Journal of Applied Ecology 41:1021–1031.
Grewal, S. K., P. J. Wilson, T. K. Kung, K. Shami, M. T. Theberge, J. B.
Theberge, and B. N. White. 2004. A genetic assessment of the eastern
wolf (Canis lycaon) in Algonquin Provincial Park. Journal Mammalogy
85:625–632.
Gude, J. A., M. S. Mitchell, R. E. Russell, C. A. Sime, E. E. Bangs, L. D.
Mech, and R. R. Ream. 2012. Wolf population dynamics in the U.S.
northern Rocky Mountains are affected by recruitment and human-caused
mortality. Journal of Wildlife Management 76:108–118.
Hamlin, K. L., R. A. Garrott, P. J. White, and J. A. Cunningham. 2009.
Contrasting wolf–ungulate interactions in the Greater Yellowstone
Ecosystem. Pages 541–577 in R. A. Garrott, P. J. White, and F. G. R.
Watson, editors. The ecology of large mammals in central Yellowstone:
sixteen years of integrated field studies. Elsevier, San Diego, California,
USA.
Hayes, R. D., R. Farnell, R. M. Ward, J. Carey, M. Dehn, G. W. Kuzyk,
A. M. Baer, C. L. Gardner, and M. O’Donoghue. 2003. Experimental
reduction of wolves in the Yukon: ungulate responses and management
implications. Wildlife Monographs 152.
Hebblewhite, M. 2006. Predator–prey management in the National Park
context: lessons from transboundary wolf, elk, moose and caribou systems.
Transactions of the North American Wildlife and Natural Resources
Conference 72:348–365.
Hebblewhite, M., and D. W. Smith. 2010. Wolf community ecology:
ecosystem effects of recovering wolves in Banff and Yellowstone National
Parks. Pages 69–120 in M. Musiano, P. Paquet, and L. Boitani, editors.
The world of wolves. University of Calgary Press, Calgary, Alberta,
Canada.
Houston, D. B. 1982. The northern Yellowstone elk herd. Macmillan, New
York, New York, USA.
Kie, J. G., R. T. Bowyer, and K. M. Stewart. 2003. Ungulates in western
coniferous forests: habitat relationships, population dynamics, and
ecosystem processes. Pages 296–339 in C. J. Zabel and R. G. Anthony,
editors. Mammal community dynamics. Cambridge University Press,
Cambridge, England, United Kingdom.
Lemke, T. O., J. A. Mack, and D. B. Houston. 1998. Winter range
expansion by the northern Yellowstone elk herd. Intermountain Journal of
Sciences 4:1–9.
Leopold, A. 1933. Game management. Charles Scribners’s Sons, New York,
New York, USA.
Leopold, A. 1949. A Sand County almanac. Oxford University Press, New
York, New York, USA.
Leopold, A. S., S. A. Cain, D. M. Cottam, I. N. Gabrielson, and T. L.
Kimball. 1963. Wildlife management in the national parks. Transactions
of the North American Wildlife and Natural Resources Conference
28:28–45.
Loveridge, A. J., A. W. Searle, F. Murindagomo, and D. W. MacDonald.
2007. The impact of sport-hunting on the population dynamics of an
African lion population in a protected area. Biological Conservation
134:548–558.
MacNulty, D. R., D. W. Smith, L. D. Mech, and L. E. Eberly. 2009. Body
size and predatory performance in wolves: is bigger better? Journal of
Animal Ecology 78:532–539.
McNay, M. 2002. Wolf–human interactions in Alaska and Canada: a review
of the case history. Wildlife Society Bulletin 30:831–843.
Mech, L. D. 1995. The challenge and opportunity of recovering wolf
populations. Conservation Biology 9:270–278.
Metz, M. C., D. W. Smith, J. A. Vucetich, D. R. Stahler, and R. O.
Peterson. 2012. Seasonal patterns of predation for gray wolves in the
multi-prey system of Yellowstone National Park. Journal of Animal
Ecology 81:553–563.
Murray, D. L., D. W. Smith, E. E. Bangs, C. Mack, J. K. Oakleaf, J.
Fontaine, D. Boyd, M. Jimenez, C. Niemeyer, T. J. Meier, D. Stahler, J.
Holyan and V. J. Asher. 2010. Death from anthropogenic causes is
partially compensatory in recovering wolf populations. Biological
Conservation 143:2514–2524.
Smith et al. Managing Wolves Around Yellowstone 9

National Park Service. 2006. Management policies 2006. U.S. Department
of the Interior, Washington, D.C., USA.
National Park Service Advisory Board Science Committee. 2012. Revisiting
Leopold: resource stewardship in the national parks. National Park Service
Advisory Board Science Committee, Washington, D.C., USA.
National Research Council. 1997. Wolves, bears, and their prey in Alaska:
biological and social challenges in wildlife management. National
Academy Press, Washington D.C., USA.
National Research Council. 2002. Ecological dynamics on Yellowstone’s
northern range. National Academy Press, Washington, D.C., USA.
Olliff, S. T., P. Schullery, G. E. Plumb, and L. H. Whittlesey. 2013.
Understanding the past: the history of wildlife and resource management
in the Greater Yellowstone Area. Pages 10–28 in P. J. White, R. A.
Garrott, and G. E. Plumb, editors. Yellowstone’s wildlife in transition.
Harvard University Press, Cambridge, Massachusetts, USA.
Phillips, M. K., and D. W. Smith. 1996. The wolves of Yellowstone.
Voyageur Press, Stillwater, Minnesota, USA.
Reid, C. 2015. What is anthropomorphism and what’s wrong with it?
Yellowstone Science 23:47.
Ruth, T. R., D. W. Smith, M. A. Haroldson, P. C. Buotte, C. C. Schwartz,
H. B. Quigley, S. Cherry, K. M. Murphy, D. Tyers, and K. Frey. 2003.
Large-carnivore response to recreational big game hunting along the
Yellowstone National Park and Absaroka-Beartooth Wilderness bound-
ary. Wildlife Society Bulletin 31:1150–1161.
Rutledge,L. Y., B. R. Patterson, K. J. Mills, K. M. Loveless,D. L. Murray, and
B. N. White. 2010. Protection from harvesting restores the natural social
structure of eastern wolf packs. Biological Conversation 143:332–339.
Schullery, P., and L. Whittlesey. 1992. The documentary record of wolves
and related wildlife species in the Yellowstone National Park area prior to
1882. Pages 1–4 to 1–174 in J. D. Varley and W. G. Brewster, editors.
Wolves for Yellowstone? A report to the United States Congress. Volume
IV: research and analysis. National Park Service, Yellowstone National
Park, Mammoth, Wyoming, USA.
Schwartz, C. C., J. E. Swenson, and S. D. Miller. 2003. Large carnivores,
moose, and humans: a changing paradigm of predator management in the
21st century. Alces 39:41–63.
Schweber, N. 2012. Famous wolf is killed outside Yellowstone. New York
Times, December 8, 2012.
Sinclair, A. R. E. 1998. Natural regulation of ecosystems in protected areas
as ecological baselines. Wildlife Society Bulletin 26:399–409.
Smith,D. W., and E. E. Bangs. 2009. Reintroduction ofwolves to Yellowstone
National Park: history, values and ecosystem restoration. Pages 92–125 in
M. W. Hayward and M. J. Somers, editors. Reintroduction of top-order
predators. Wiley-Blackwell, Oxford, England, United Kingdom.
Smith, D. W., D. R. Stahler, E. Albers, R. McIntyre, M. Metz, K. Cassidy,
J. Irving, R. Raymond, H. Zaranek, C. Anton, and N. Bowersock. 2010.
Yellowstone wolf project: annual report YCR-210-06, 2009. National
Park Service, Yellowstone National Park, Wyoming, USA.
Smith, D. W., D. R. Stahler, E. Stahler, M. Metz, K. Quimby, R. McIntyre,
C. Ruhl, and M. McDevitt. 2014. Yellowstone wolf project: annual report
YCR-2012-02, 2013. National Park Service, Yellowstone Center for
Resources, Yellowstone National Park, Wyoming, USA.
Stahler, D. R., D. R. MacNulty, R. K. Wayne, B. vonHoldt, and D. W.
Smith. 2013. The adaptive value of morphological, behavioral and life-
history traits in reproductive female wolves. Journal of Animal Ecology
82:222–234.
Stronen, A. V., R. K. Brook, P. C. Paquet, and S. Mclachlan. 2007. Farmer
attitudes toward wolves: implications for the role of predators in managing
disease. Biological Conservation 135:1–10.
Taper, M. L., and P. J. P. Gogan.2002. The northern Yellowstone elk: density
dependence and climatic conditions. Journal of Wildlife Management
66:106–122.
Theberge, J. T., and M. T. Theberge. 2004. The wolves of Algonquin Park:
a 12 year ecological study. Department of Geography, University of
Waterloo, Publication Series No. 56, Waterloo, Ontario, Canada.
The Wildlife Society. 2007. Final TWS position statement. The North
American model of wildlife conservation. The Wildlife Society, Bethesda,
Maryland, USA.
Thiessen, C. D. 2007. Population structure and dispersal of wolves (Canis
lupus) in the Canadian Rocky Mountains. Thesis, University of Alberta,
Alberta, Canada.
Treves, A., and K. A. Martin. 2011. Hunters as stewards of wolves in
Wisconsin and the northern Rocky Mountains, USA. Society and Natural
Resources 24:984–994.
Treves, A., L. Naughton-Treves, and V. Shelley. 2013. Longitudinal
analysis of attitudes toward wolves. Conservation Biology 27:315–323.
Unger, K. 2008. Managing a charismatic carnivore. Wildlife Professional
2:30–33.
U.S. Fish and Wildlife Service [USFWS]. 1987. Northern Rocky Mountain
wolf recovery plan. U.S. Fish and Wildlife Service, Denver, Colorado,
USA.
U.S. Fish and Wildlife Service [USFWS]. 2006. 12-Month finding on a
petition to establish the northern Rocky Mountain gray wolf population
(Canis lupus) as a distinct population segment to remove the northern
Rocky Mountain gray wolf distinct population segment from the list of
endangered and threatened species. Federal Register 71:43410–43432.
U.S. Fish and Wildlife Service [USFWS]. 2011. Reissuance of final rule to
identify the northern Rocky Mountain population of gray wolf as a distinct
population segment and to revise the list of endangered and threatened
wildlife. Federal Register 76:25590–25592.
U.S. Fish and Wildlife Service [USFWS], Idaho Department of Fish and
Game, Montana Fish, Wildlife & Parks, Wyoming Game and Fish
Department, Nez Perce Tribe, National Park Service, Blackfeet Nation,
Confederated Salish and Kootenai Tribes, Wind River Tribes, Confeder-
ated Colville Tribes, Spokane Tribe of Indians, Washington Department
of Fish and Wildlife, Oregon Department of Fish and Wildlife, Utah
Department of Natural Resources, and U.S. Department of Agriculture
Wildlife Services. 2014. Northern Rocky Mountain Wolf Recovery
Program 2013 Interagency Annual Report. M. D. Jimenez and S. A.
Becker, editors. Ecological Services, Helena, Montana, USA.
vonHoldt, B. M., D. R. Stahler, E. E. Bangs, D. W. Smith, M. D. Jimenez,
C. M. Mack, C. C. Niemeyer, J. P. Poliinger, and R. K. Wayne. 2010.
A novel assessment of population structure and gene flow in grey wolf
populations of the northern Rocky Mountains of the United States.
Molecular Ecology 19:4412–4427
Wagner, F. 2006. Yellowstone’s destabilized ecosystem: elk effects, science,
and policy conflict. Oxford University Press, New York, New York, USA.
White, P. J., and R. A. Garrott. 2013. Predation: wolf restoration and the
transition of Yellowstone elk. Pages 69–93 in P. J. White, R. A. Garrott,
and G. E. Plumb, editors. Yellowstone’s wildlife in transition. Harvard
University Press, Cambridge, Massachusetts, USA.
White, P. J., R. A. Garrott, and G. E. Plumb. 2013a. Ecological process
management. Pages 3–9 in P. J. White, R. A. Garrott, and G. E. Plumb,
editors. Yellowstone’s wildlife in transition. Harvard University Press,
Cambridge and London, United Kingdom.
White, P. J., R. A. Garrott, and G. E. Plumb. 2013b. The future of
ecological process management. Pages 255–266 in P. J. White, R. A.
Garrott, and G. E. Plumb, editors. Yellowstone’s wildlife in transition.
Harvard University Press, Cambridge and London, United Kingdom.
Woodroffe, R., and J. R. Ginsberg. 1998. Edge effects and the extinction of
populations inside protected areas. Science 280:2126–2128.
Associate Editor: Glenn.
10 Wildlife Society Bulletin 9999()