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

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  • Sigurd Olson Environmental Institute

Abstract and Figures

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.
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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 naivete
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.
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10 Wildlife Society Bulletin 9999()
... Kellert et al 1996, Young et al 2015, Slagle et al 2017, Jones et al 2019. In this region there are areas with intact carnivore guilds and wilderness; yet rapid human development and polarizing debate about carnivore management threaten the future of these animals and can hinder effective policy-making (Bangs and Shivik 2001, Linnell et al 2001, Bruskotter 2013, Bradley et al 2015, Smith et al 2016. In particular, interaction between livestock and carnivores generates intense controversy in the American West (van Eeden et al 2018), where publically-owned grazing land is ubiquitous and livestock production is an important economic sector (Sarchet 2005). ...
... These obstacles require future research on how to build platforms for collaboration among key actors and institutions across states in order to foster coexistence in the American West. Although the formation of cross-state platforms might lead to new conflicts (Redpath et al 2017), such platforms could also represent new institutions by which multiple actors can embrace the challenge of coexisting with carnivores and engage to find shared solutions (see also Smith et al 2016, Hartel et al 2019. In this context, our proposal goes beyond including different sectors and actors since it aims to promote cross-state and transboundary collaborative management in the American West. ...
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Carnivore and humans live in proximity due to carnivore recovery efforts and ongoing human encroachment into carnivore habitats globally. The American West is a region that uniquely exemplifies these human-carnivore dynamics, however, it is unclear how the research community here integrates social and ecological factors to examine human-carnivore relations. Therefore, strategies promoting human-carnivore coexistence are urgently needed. We conducted a systematic review on human-carnivore relations in the American West covering studies between 2000 and 2018. We first characterized human-carnivore relations across states of the American West. Second, we analyzed similarities and dissimilarities across states in terms of coexistence, tolerance, number of ecosystem services and conflicts mentioned in literature. Third, we used Bayesian modeling to quantify the effect of social and ecological factors influencing the scientific interest on coexistence, tolerance, ecosystem services and conflicts. Results revealed some underlying biases in human-carnivore relations research. Colorado and Montana were the states where the highest proportion of studies were conducted with bears and wolves the most studied species. Non-lethal management was the most common strategy to mitigate conflicts. Overall, conflicts with carnivores were much more frequently mentioned than benefits. We found similarities among Arizona, California, Utah, and New Mexico according to how coexistence, tolerance, services and conflicts are addressed in literature. We identified percentage of federal/private land, carnivore family, social actors, and management actions, as factors explaining how coexistence, tolerance, conflicts and services are addressed in literature. We provide a roadmap to foster tolerance towards carnivores and successful coexistence strategies in the American West based on four main domains, (1) the dual role of carnivores as providers of both beneficial and detrimental contributions to people, (2) social-ecological factors underpinning the provision of beneficial and detrimental contributions, (3) the inclusion of diverse actors, and (4) cross-state collaborative management.
... The active reintroduction mandated by the ballot initiative would provide more flexibility as to where to initially restore wolves, preferably in regions with relatively lower risk of conflict with people. Restoring and managing a population of wolves in Colorado both greatly expands the occupancy within their historical range and establishes another viable protected population that could be managed within a regional framework in the Rocky Mountains (Smith et al., 2016;Mech, 2017). ...
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Reintroducing native carnivores risks creating conflict with people and consequently reducing support for coexistence and conservation efforts. Determining the interface between areas of ecological suitability and conflict risk can help enhance success of carnivore restoration, but this is often difficult because accurate data on risks and tolerance are lacking. Gray wolves (Canis lupus), a focus of reintroduction efforts in the US, require tolerance to persist in human-dominated landscapes but also catalyze societal-level conflicts throughout their global range. Via an unprecedented process to restore an apex predator, in November 2020, citizens in the state of Colorado, USA voted to reintroduce wolves to the state where they had been extirpated ~70 years prior. We leveraged voting records of over three million citizens to quantify and map an index of tolerance for wolves and combined it with spatially explicit data on livestock distributions and land ownership to create predictions of direct conflict risk between wolves and humans. Conflict risk was juxtaposed with estimates of wolf ecological suitability developed using seasonal prey densities along with environmental and anthropogenic features that influence wolf habitat use. Our social-ecological modeling approach predicted that ~56% of the West Slope of Colorado contained ecologically suitable habitat and relatively low conflict risk. Our models also delineated possible conflict hotspots where ecological suitability and conflict risk converge, thus facilitating targeted proactive management. We demonstrate how voting patterns can provide unique, spatially explicit insight on tolerance that can be integrated with other information to help facilitate human-carnivore coexistence and carnivore restoration success.
... encouraging species movement into the buffer zone could increase the flow of ecosystem services, but also the potential for human-wildlife conflict (Pascual-Rico et al., 2020). While this issue will be most important for rewilding projects which include carnivores or large herbivores (e.g., Smith et al., 2016), a recent attempt to introduce a breeding pair of beavers in the Knepp estate failed because the animals quickly moved out of the core rewilding site (Knepp, 2021). This highlights that monitoring environmental conditions in the area surrounding rewilded sites may play an important role in understanding (and responding to) ecological changes inside such sites. ...
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... Common The ongoing debate about preserving or eliminating dingoes in Australia mirrors the controversy about preserving or hunting other wild predators of domestic animals anywhere in the world. This includes wolves (Canis lupus) and bears (Ursus arctos) in Europe (Majić et al. 2011;Kuijper et al. 2019) and North America (Smith et al. 2016), but also felids such as lions (Panthera leo), tigers (Panthera tigris) and leopards (Panthera pardus) in Africa and India (Athreya et al. 2013;Krafte Holland et al. 2018). Even though the above mentioned carnivores are locally or globally endangered, and are not expected to attain large population sizes, or to become a direct threat to people lives, some lobbies (typicallyhunting and farmer associations) still advocate for their eradication or oppose reintroductions or other management actions (Fernández-Gil et al. 2016). ...
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Feral animals are those that live wild but are descendants of domesticated populations. Although in many cases, possibly the majority, these populations suppose a risk to the environment and may conflict with wild local species and human activities, there are feral populations that are considered worth preserving hold and, in some cases, already enjoy protection from interest groups and even pubic authorities. In this review, we aimed to separate those valuable populations using model cases classified by three main criteria of interest: (a) the genetic conservation value in case of extinct wild ancestors, (b) the niche occupancy criterion and, finally, (c) a cultural criterion. We propose a detailed analysis of feral populations under scrutiny, supporting control measures when necessary, but also allowing for international protection at the same level as wild animals for feral taxa of special concern. Feral taxa which are already in the focus of conservation efforts and may be awarded extended recognition and protection include ancient lineages of feral dogs, horses, camels, goats and bees (as pollinators) in different parts of the world.
... The Endangered Species Act (ESA) of 1973 provided federal protection for wolves throughout the US and enabled efforts for recovery of wolf populations across parts of their historical range. The debate over wolf recovery and management has been fraught with conflict (Smith et al. 2016). Nonetheless, reintroductions in the 1990s were successful in re-establishing wolf populations in the Northern Rocky Mountain region (Smith et al. 2003). ...
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Humans regularly exert a powerful influence on the survival and persistence of species, yet social‐science information is used only sporadically in conservation decisions. Using data obtained from a survey of 46,894 US residents, we developed and applied a spatially explicit “sociocultural index” to inform decision making through an understanding of public values toward wildlife. The classification is defined by opposing values of mutualism and domination, which have been previously shown to be highly predictive of attitudes on a wide range of policy issues. We developed state and county maps that can be used to represent public interests in policy decisions and inform management actions that target human behavior, such as education. To illustrate, we present findings indicating a supportive social context for gray wolf (Canis lupus) reintroduction in Colorado, an issue voted on and passed through a November 2020 citizen ballot initiative. Although the results are particularly relevant for the US, the technique is broadly applicable and its expansion is encouraged to better account for human factors in conservation decisions globally.
... Addressing transboundary mortality of large carnivores is challenging because of difficulties of reconciling different management paradigms, policies, and social pressures across park and non-park lands around the globe (Hebblewhite, 2007;Smith et al., 2016;Woodroffe and Ginsberg, 2001). Transboundary management agreements elsewhere provide an array of options ranging from the status quo to increased protection for wolves. ...
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Large carnivores are important ecological drivers of ecosystem dynamics when they occur at ecologically effective densities. They are also challenging to conserve, especially in transboundary settings such as along borders of parks and protected areas. Here, we tested for effects of transboundary movements on survival of 72 radiocollared gray wolves from 1987 to 2018 in and adjacent to Banff National Park, Canada. We fit Bayesian counting-process survival models to known-fate radiotelemetry data and tested for the influence of intrinsic covariates such as sex and age, time, and movements outside of protected areas on survival of wolves. We also estimated cause-specific mortality. Non-parametric survival was 0.733 (95% CI 0.622–0.816), and the top Bayesian survival model indicated that wolves outside the park had much lower annual survival rates (0.44, 95% BCI = 0.24–0.65) compared to wolves inside the park (0.84, 95% BCI = 0.73–0.91). The cumulative risk of mortality was on average 6.7 times higher (odds ratio 95% BCI = 2.2–21.4) for wolves outside the park, peaking during the winter hunting and trapping seasons. We found weak evidence for declining survival over time, opposite to patterns predicted by density-dependence. Bayesian cause-specific mortality indicated that the top three sources of mortality were trapping (rate = 0.080, 36% of mortality), followed by hunting (0.053, 18%), and highway (0.046, 18%) mortality. Surprisingly, we found no intraspecific mortality, and low dispersal from Banff National Park. This demographic profile is akin to other exploited populations across North America. While we were unable to combine survival rates with reproduction to estimate population trends, the overall mortality rates within our study area are consistent with a stable wolf population. Nonetheless, the long-term stability and ecological effectiveness of wolves likely differed inside and outside of protected areas, which highlights a challenge with managing transboundary carnivores exposed to different management regimes.
... Wolf harvest became seasonally legal in the autumn of 2009 and has been so since (with the exception of the winter of 2010-2011). Human harvest is permissible only outside of YNP and was often limited by a quota in the relevant geographic area (see Fig. 1c and Smith et al. 2016). Most harvest that affected wolves in our study occurred in Montana, where the harvest season generally began in September and lasted through March, unless the quota was filled. ...
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Imperfect detection is ubiquitous among wildlife research and is therefore commonly included in abundance estimation. Yet, the factors that affect observation success are largely unknown for rare and elusive species, such as large carnivores. Here, we took advantage of intensive ground‐based monitoring effort and an extensive GPS data set (2000–2018) and developed a winter sightability model for gray wolves (Canis lupus) in northern Yellowstone National Park, Wyoming, USA. Our resulting sightability model indicated that observation success was positively affected by the topographic nature of where wolves were in relation to observer locations (viewshed), areas being less forested (openness), and wolf group size, and negatively affected by distance from observer locations. Of these, viewshed had the strongest effect on the probability of observing a wolf, with the odds of observing a wolf being four times more likely when wolves were in the predicted viewshed. Openness was the next most influential covariate, and group size was the least influential. We also tested whether a wolf being harvested from a pack when they were outside of Yellowstone National Park had an effect on wolf sightability. We did not, however, find support for human‐induced mortality affecting wolf sightability inside of Yellowstone National Park. Our results indicate that the ability to observe wolves was greatly affected by ecological and landscape‐level factors, a finding that is likely to generally extend to other large carnivores. As such, our sightability model highlights the importance of considering landscape structure and variation in large carnivore use of the landscape when conducting observational‐based studies.
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Understanding the behavioral responses of large carnivores to human activity in protected areas is important for conserving top predators. Roads and associated vehicle traffic have a range of impacts on wildlife, including mortality from vehicle collisions and behavioral changes from increasing traffic levels. Roads concentrate human activities and may be particularly impactful when located adjacent to high‐quality habitat for wildlife. However, people often overlook road impacts in protected areas because of relatively low road densities. From 1979 to 2017, annual visits to Yellowstone National Park increased from 1.9 to 4.1 million, with many visitors in the last 25 yr focusing on the opportunity to view wild wolves (Canis lupus) in their natural habitat from the roadway. To better understand how human activity interacts with landscape attributes and prey availability to shape wolf habitat use, we developed seasonal and diel‐specific step selection functions (SSF) for wolves. Wolves responded to increased human activity by using areas farther from roadways during the day and during peak visitation in summer. Prey availability, as estimated by an elk SSF, did not significantly alter habitat selection patterns by wolves. The strength of habitat selection in relation to roads varied among wolf packs. The most heavily viewed wolf packs exhibited less road avoidance, suggesting increased tolerance, which could lead to increased vulnerability to human harvest if they leave the park. Federal and state managers have implemented several measures to mitigate disturbance effects to wolves and curtail habituation. These results may inform adaptive management strategies that seek to continue to conserve natural wolf behavior.
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A LARGE, DARK WOLF poked his nose out of the pines in Yellowstone National Park as he thrust a broad foot deep into the snow and plowed ahead. Soon a second animal appeared, then another, and a fourth. A few minutes later, a pack of thirteen lanky wolves had filed out of the pines and onto the open hillside. Wolf packs are the main social units of a wolf population. As numbers of wolves in packs change, so too, then, does the wolf population (Rausch 1967). Trying to understand the factors and mechanisms that affect these changes is what the field of wolf population dynamics is all about. In this chapter, we will explore this topic using two main approaches: (1) meta-analysis using data from studies from many areas and periods, and (2) case histories of key long-term studies. The combination presents a good picture-a picture, however, that is still incomplete. We also caution that the data sets summarized in the analyses represent snapshots of wolf population dynamics under widely varying conditions and population trends, and that the figures used are usually composites or averages. Nevertheless, they should allow generalizations that provide important insight into wolf population dynamics.
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In the Canadian Rocky Mountains, the gray wolf (Canis lupus) has experienced range contractions and expansions, which can greatly affect pack stability as well as population structure. In addition, this area has a highly heterogeneous landscape that may form barriers to dispersal. To understand factors affecting pack structure and large-scale gene flow across the Rocky Mountains, we examined wolf genetic structure using 1,981 noninvasive and invasively collected samples. We sampled over 44 packs in Alberta and British Columbia and, from these, identified 540 individuals based on 12 microsatellites. Relatedness of individuals within packs was greater than between packs, and female relatedness was greater than males suggesting strong pack structure and female philopatry. Relatedness within packs was greater near major roads suggesting decreased dispersal from natal packs with proximity to roads. Across the study area, 2 significantly differentiated genetic clusters were identified, corresponding to a north/south split. Landcover distance was a significant correlate for 2 of 4 genetic distance measures, where packs in the north were in areas of dense coniferous forest, while packs in the south were primarily in open coniferous forest. These landcover differences suggest natal associations or could relate to prey distribution. Fine-scale investigation of pack dynamics across this continuous distribution, together with large-scale estimators of population structure, highlights different drivers of gene flow at the pack and population level.
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Historical accounts, park records, and biologists' observations indicated that wintering elk in Yellowstone's northern range were present in low numbers prior to and at park establishment in 1872; increased to 20,000-35,000 by the early 1900s when they heavily impacted the northern-range ecosystem; and declined to 3,172 censused animals in 1968 due to park control efforts. In 1967, the park announced a politically coerced natural-regulation policy terminating park control; and in 1971 posed a natural-regulation ecological hypothesis stating that northern-range elk had been numerous prior to 1872, had not risen to 20,000-35,000 in the early 1900s, and would stabilize at moderate numbers following recovery from control efforts without significantly affecting the northern-range ecosystem. Archaeological and historic evidence, park records, and censuses begun in the 1920s indicate a northern herd of ~5,000-6,000 in 1872, increasing to ~20,000-35,000 in the early 1900s, declining to a censused number of 3,172 in 1968 in response to control efforts, increasing to a census-based number of 21,071-25,920 in the 1980s and 1990s, then declining somewhat after 2000. This book reviews critically the published and unpublished records to test the natural-regulation hypothesis and propose a conceptual model of the northern-range ecosystem with inferences from system changes associated with the four stages of elk abundance, inside-outside exclosure comparisons, and system comparisons inside and outside park boundaries.