Via Email and Federal Express
June 30, 2015
Joanna Eide, Legal Services
Washington Department of Fish and Wildlife
P.O. Box 43144
Olympia, WA 98504-3144
Joanna.eide@dfw.wa.gov
Cc:

Washington Fish & Wildlife Commission
600 Capitol Way N.
Olympia, WA 98501
Governor Jay Inslee
Office of the Governor
P.O. Box 40002
Olympia, WA 98504-0002

Re: Petition to Amend Washington Administrative Code Section 23228-297
The Humane Society of the United States (HSUS), Conservation
Northwest, Center for Biological Diversity, Mountain Lion Foundation, Wolf
Haven International, The Cougar Fund, The Lands Council, Predator
Defense, Kettle Range Conservation Group, and Gary Koehler, Ph. D.,
collectively “Petitioners,” hereby petition the Washington Fish & Wildlife
Commission (the “Commission”), pursuant to RCW § 34.05.340(3) and RCW §
34.05.330, to amend the Commission’s May 1, 2015 regulation that
unjustifiably and unsustainably increased the hunting quotas for cougars.
On February 2, 2015, the Commission published notice of a proposed
rulemaking establishing regulations for, in relevant part, the 2015-2016,

1

 2016-2017 and 2017-2018 cougar hunting seasons. 1 The proposal appeared to
carefully consider the scientific recommendations of the Washington
Department of Fish and Wildlife (the “Department” or “WDFW”) and other
leading cougar experts. It proposed maintaining the existing harvest rate of
12-16 percent of total population, on the grounds that “[r]ecently published
studies suggest that a 12-16 percent harvest rate of a local cougar population
is the maximum harvest rate that still has a high probability for maintaining
a stable cougar population along with stable adult male territorial behavior.” 2
Public comment was solicited (and submitted by Petitioners) and a public
hearing on the Proposed Rule was held on March 21, 2015.
But at the April 10 Commission meeting where the Proposed Rule was
docketed for finalization, the Commission abruptly jettisoned this reasoning,
proposing and approving a last-minute amendment increasing cougar harvest
rates in select areas, the zones where wolf packs overlap with cougars, to 1721 percent of total population. 3 This amendment, which nearly doubled
harvest rates in some areas, was made without providing any prior notice to
the public. Many stakeholders with serious concerns as to the lack of
scientific support for the change and its potential for adverse ecological
consequences were made aware of the amendment for the first time after it
passed, and were consequently left without opportunity to comment in
opposition to the regulation.
Proposed Rule Making, WSR 15-04-085 (Feb 2, 2015) (“Proposed Rule”).
Id., at 10 (emphasis added).
3 Rule Making Order, WSR 15-10-066 (May 1, 2015) (“Final Rule”).
1
2

2

 To ensure that this significant policy change receives the careful
deliberation and public input required by law and sound principles of wildlife
management, Petitioners hereby petition the Commission and Department to
amend WAC 232-28-297 to revert cougar harvest guidelines in the 14 hunting
areas altered in the Final Rule to the science-based levels originally
submitted in the Proposed Rule. For the reasons outlined below, the best
available science strongly supports the petitioned action and the Commission
therefore has a legal duty to reconsider this arbitrary regulation.
This petition is submitted pursuant to RCW § 34.05.340(3) on the
grounds that the Final Rule is substantially different than the Proposed
Rule, and as a citizen petition pursuant to RCW § 34.05.330 to amend WAC
232-28-297. It is filed within sixty days of the May 1, 2015 publication of the
Final Rule by the Code Reviser. 4 Proposed amended rule language is
attached herewith as Attachment A.
I.

Interests of the Petitioners
The Humane Society of the United States (“The HSUS”) is a nonprofit

organization that promotes the protection of all animals. It is the largest
animal protection organization in the United States, with several staff and
thousands of members in Washington. Since its inception in 1952, The HSUS
has worked to foster the humane treatment of wildlife through various
programs, including initiatives to protect native carnivores.
RCW § 34.05.340(3) (60-day timeframe for filing petitions based on variance between final
and proposed rule).
4

3

 Conservation Northwest, based in Bellingham, Washington, has been
working to conserve the wildlife and wildlands of the Pacific Northwest for
more than 25 years. It focuses on protecting viable populations of large
carnivores, including cougars, bears, wolves, wolverine, and lynx, and
ensuring that they have large, connected landscapes with high-quality
habitat in which to thrive, thereby ensuring that smaller species important to
maintaining biodiversity and ecosystems are also healthy.
The Center for Biological Diversity is a national, nonprofit
conservation organization with more than 900,000 members and online
activists dedicated to the protection of wildlife and wild places. The Center
submits this petition in support of sound, science-based cougar management
on behalf of our thousands of Washington members.
The Mountain Lion Foundation is a 501(c)3 non-profit conservation
organization founded in 1986 to protect cougars and their habitat. It is
registered to do business in Washington, and has been involved with
protecting Washington’s cougars since the Initiative 655 campaign in 1996.
Wolf Haven International disagrees with using wolves as an excuse to
pass laws to implement scientifically unsound kill rates of another predator
just because wolves exist in the same landscape. It believes that wildlife
officials should start with agreed-upon scientific parameters, and set kill
rates based on public tolerance within those parameters.

4

 The Cougar Fund is a national organization with many constituents in
the State of Washington. Its mission is to protect mountain lions and
other large carnivores through an effective trident of education, advocacy,
and the promotion of current best science as the basis for wildlife
management.
The Lands Council has been working to protect wildlife, including
important predators such as cougar, for over 30 years. It has over 1500
members and supporters in Washington State.
Predator Defense is extremely concerned about the senseless killing of
cougars and other territorial apex predators across the country. It is a
national nonprofit organization with members in Washington.
Kettle Range Conservation Group is a non-profit organization founded
in Republic, Washington in 1976 and whose mission is to defend wilderness,
protect biodiversity, and restore ecosystems of the Columbia River Basin.
Kettle Range Conservation Group staff and members regularly recreate
throughout Washington for the purposes of hiking, recreation, bird watching,
observing wildlife such as cougars, wolves and other recreational and
professional pursuits.
Gary M. Koehler, Ph. D. is a retired Research Scientist for Carnivore
Investigations at Washington Department of Fish and Wildlife, and the
current Scientific Advisor for Snow Leopard Trust and Save China’s Tigers.
As WDFW’s principal investigator for the studies of cougars in Washington in

5

 his position with WDFW, and co-author of many of the publications cited in
the petition, he has concerns for the Commission’s lack of scientific and
biological support for their decision to increase harvest for cougars in the
selected portions of the state.
II.

Factual and Scientific Support for Petitioned Action
From 2000 to the present, Washington State University, University of

Washington, and the Department have collaborated on a series of research
projects on Washington cougar management. 5 These studies, conducted by
experts - including the Department’s own biologists - have formed the
scientific basis for the Department’s cougar management program in
previous years. 6 Yet the quota increase in the Final Rule represents a sudden
and unsubstantiated shift away from the nearly unanimous conclusions
reached by these researchers. It departs dramatically from the harvest
thresholds recommended by cougar biologists, 7 and ignores strong evidence of

See, e.g., Kaylie A. Peebles et al., “Effects of Remedial Sport Hunting on Cougar Complaints
and Livestock Depredations,” PLoS ONE 8, no. 11 (Nov. 2013); Richard A. Beausoleil et al.,
“Research to Regulation: Cougar Social Behavior as a Guide for Management,” Wildlife
Society Bulletin 37, no. 3 (June 2013); Hilary S. Cooley et al, “Does hunting regulate cougar
populations? A test of the compensatory mortality hypothesis,” Ecology 90, no. 10 (2009)
(“Cooley et al. 1”); Hilary S. Cooley et al., “Source Populations in Carnivore Management:
Cougar Demography and Emigration in a Lightly Hunted Population,” Animal Conservation
12, no. 4 (2009) (“Cooley et al. 2”); see also H.S. Robinson et al., “Sink Populations in
Carnivore Management: Cougar Demography in a Hunted Population,” Ecological
Applications 18, no. 4 (2008); C.M.S. Lambert et al., “Cougar Population Dynamics and
Viability in the Pacific Northwest,” Journal of Wildlife Management 70 (2006).
6 See Washington Department of Fish and Wildlife, Final Supplemental Environmental
Impact Statement for the 2015-2021 Game Management Plan” (July 31, 2014) (“Department
EIS”) (“The scientific findings from [these] projects…have been incorporated into how the
Department currently manages cougar.”) .
7 See, e.g. Beausoleil et al. at 684, 686 (14 percent harvest threshold); Department EIS at
109-10 (12-16 percent harvest threshold).
5

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 deleterious consequences for cougars and humans alike. It is imperative that
the Commission open a public comment period in order to fully consider the
quota increase in light of the best available science, and to engage
stakeholders in a dialogue about the rationale animating this change in
course.
a. The Best Available Science Concludes that Increased
Hunting Quotas Will Harm Cougar Social Structures,
Increase Human-Cougar Conflict, and Impede the
Department’s Cougar Management Goals
i. Population Management and Dynamics
Studies specific to Washington cougars have consistently demonstrated
that increased sport hunting does not result in a net change in total cougar
population within a given area. 8 Cougars display long-distance immigration
and emigration patterns that compensate for and offset the population effects
of sport hunting (the “source-sink” model). 9 Even heavily hunted areas have
shown no net reduction in the population. 10 Because immigrants will
inevitably replace cougars removed from the landscape by hunters, sport
hunting is an ineffective means of controlling total population size.
But even though increased hunting will not reduce the total
population, it will nevertheless have potentially harmful effects on cougar
social dynamics and population structure. As the Department itself
recognized in its Proposed Rule:

See Peebles et al., at 6; see also Cooley et al. 1 at 2919.
See Cooley et al. 1, at 2913-14; see also Peebles et al., at 6.
10 Id.
8
9

7

 Recently published studies suggest that a 12-16 percent harvest rate of
a local cougar population is the maximum harvest rate that still has a
high probability for maintaining a stable cougar population along with
stable adult male territorial behavior. Harvest rates in excess of 16
percent can result in declines in core populations of breeding females,
and excessive male harvest rates result in the loss of adult male
territorial behavior, which acts as a regulatory mechanism for local
male cougar numbers. 11
The Department’s reading of the leading studies is correct. They have
consistently shown that removal of adult males through sport hunting
adversely affects the sex and age demographics of local cougar populations by
creating territorial vacancies that attract subadult, dispersing males. 12 This
compensatory immigration by subadult males causes increased infanticide
and a decline in adult female and kitten survival rates, undermining the local
population’s ability to stabilize under heavier hunting pressure. 13
ii. Human-Cougar Conflict and Social Tolerance
The benefits of a stable social structure are not limited to the cougars
themselves. Indeed, studies conducted by Department biologists and other
experts have shown time and time again that the disruption in demographic
structure caused by hunting in excess of the recommended 12-16 percent
range generates more complaints and livestock depredations. Leading studies
relied on by the Department suggest that “increased harvest up to 24% of the
estimated population had no effect on reducing the overall number of cougarProposed Rule, at 10.
Cooley et al. 1, at 2918-9; see also Cooley et al. 2, supra n. 5; Lambert et al., supra n. 5;
Robinson et al., supra n. 5; D. Stoner, M. , M.L. Wolfe, and D. Choate, "Cougar Exploitation
Levels in Utah: Implications for Demographic Structure, Population Recovery, and
Metapopulation Dynamics," Journal of Wildlife Management 70, (2006);
13 Cooley et al. 1, at 2919. R. B. Wielgus et al., "Effects of Male Trophy Hunting on Female
Carnivore Population Growth and Persistence," Biological Conservation 167 (2013).
11
12

8

 human conflicts” 14 and rather “[t]he odds of increased complaints and
livestock depredations increased dramatically…with increased cougar
harvest.” 15 This is because the demographic shifts wrought by increased
harvest attract subadult male immigrants who are far more likely to engage
humans and livestock, and who push adult females into hunting suboptimal
prey or entering human-inhabited areas in search of food. 16 The best
available science therefore demonstrates that, despite its intentions, the
Final Rule stands to worsen human-cougar conflict. 17
b. No Evidence Exists Supporting the Purported Rationale for
Increasing Quotas
Per the explanatory statement prepared by the Department, the Final
Rule seeks to “increase the harvest guideline from 12-16% of the cougar
population…to 17-21% in hunt areas that overlap known wolf packs.” 18 While
that statement does not specifically address the rationale underpinning the
increased quotas for these areas, statements at Commission meetings
indicate that the purpose of this change was to improve social tolerance of
predators in areas with both cougar and wolf populations.
There is simply no evidence suggesting that the increased cougar
quotas will achieve improved social tolerance of wolves. But the premise—
that hunting carnivores will increase tolerance—has been empirically proven
Department EIS at 108, citing generally Peebles et al.
Peebles et al, at 1.
16 Id.; see also B.N. Kertson, “Cougar ecology, behavior, and interactions with people in a
wildland-urban environment in western Washington,” Dissertation, University of
Washington (2010).
17 See generally Peebles et al.
18 Final Rule, Attachment B.
14
15

9

 as false. 19 Moreover, ample evidence suggests that increasing the hunting of
cougars will have the opposite effect. First, the notion that hunting additional
cougars will ease the social tension created by an unacceptable level of
human conflict with wolves — an entirely different species — is facially
nonsensical and is not supported by any scientific evidence. 20 By its plain
terms the Final Rule does not address wolves, and it is unreasonable to
expect that social tolerance of wolves will improve when the same number of
wolves will be managed in the same way after the Final Rule is implemented.
There is, however, credible reason to believe that the Final Rule will
worsen social tolerance of native carnivores. As discussed above, the social
destabilization created by heavier cougar hunting raises the risk of increased
levels of livestock depredation and other forms of human-cougar conflict. 21 To
the extent that social tolerance concerns stem from negative human
interactions with wildlife, research indicates that increased harvest
thresholds can harm social tolerance by generating more complaints and
more depredations. 22 If local human communities are truly unable to
differentiate between cougar and wolf-related conflict, then the expected
uptick in cougar depredations caused by the Final Rule may have an adverse
spillover effect on the social tolerance of wolves as well.
Adrian Treves, "Hunting for Large Carnivore Conservation," Journal of Applied Ecology
46, (2009).
20 Wolves, cougars, and bears can limit each other’s’ populations through competition and
predation, a notion call intraguild predation. Toni Ruth and Kerry Murphy, "Competition
with Other Carnivores for Prey," in Cougar: Ecology and Conservation, ed. Maurice
Hornocker and Sharon Negri (Chicago and London: University of Chicago Press, 2010).
21 See, e.g., Peebles et al. at 6.
22 Id.
19

10

 Finally, it should be noted that the Department’s own research
illustrates that social tolerance of cougars among Washingtonians is
presently very high. 23 A statewide agency survey found that 92% of
respondents “agree with the statement that cougars are an essential
component of Washington ecosystems, and 92% agree with the statement
that cougars have an inherent right to live here.” 24 Conversely, “87% of
Washington residents disagree with the statement that cougars are a
nuisance animal damaging rural economies.” 25 And the overwhelming social
tolerance for cougars is not limited to urban areas; rather “the positive value
of cougars is a predominant sentiment in both urban and rural
communities.” 26 In sum, the best available evidence proves that raising
hunting quotas seeks to fix something that isn’t broken, and risks birthing a
social tolerance problem where one does not currently exist.
II.

Legal Support for Petitioned Action
a. The Petition Should Be Granted Because the Final Rule is
Substantially Different From the Proposed Rule
The Washington Administrative Procedure Act (“WAPA”), RCW §

34.05 et seq., governs Commission rulemaking 27 and protects the right of
stakeholders and the general public to a robust notice and comment process.
To prevent post-notice amendments to the substance of a proposed rule from
Washington Department of Fish and Wildlife, “Cougar Outreach and Education in
Washington State” (Nov. 30, 2010).
24 Id., at 17.
25 Id.
26 Id., at 52.
27 RCW § 77.04.130 (“Rules of the commission shall be adopted by the commission or a
designee in accordance with chapter 34.05 RCW.”).
23

11

 depriving interested parties of these procedural rights, it provides that “an
agency may not adopt a rule that is substantially different from the rule
proposed in the published notice of proposed rule adoption or a supplemental
notice in the proceeding.” 28 When, like in this case, no such supplemental
notice is provided, WAPA allows “any interested person” to “petition the
agency to amend any portion of the adopted rule that is substantially
different from the proposed rule.” 29 This Petition seeks to amend the
increased harvest guidelines contained in the Final Rule and restore them to
the levels proposed in the Proposed Rule (and does not otherwise seek to
amend the Final Rule). Because the Final Rule meets two out of three
statutory factors – out of which any one is sufficient to make a finding of
substantial difference – the Commission should initiate a rulemaking process
implementing the petitioned amendment. 30
i. Difference in Effect
One of the factors to be considered in determining whether “an
adopted rule is substantially different from the proposed rule on which it is
based” is “[t]he extent to which the effects of the adopted rule differ from the
effects of the published proposed rule.” 31 Here, the effects of the Final Rule
differ from the Proposed Rule because the Final Rule substantially increased

RCW § 34.05.340.
RCW § 34.05.340(3). Notably, the agency may not reject petitions submitted pursuant to
this section and instead must initiate rulemaking proceedings if substantial variance is
shown.
30 RCW § 34.05.340(2) (three factors for substantial difference are difference in effect,
difference in scope, inadequate notice to persons affected).
31 Id.
28
29

12

 cougar quotas in fourteen areas while the Proposed Rule followed the
recommendation of the Department’s biologists and other experts and kept
quotas at status quo levels. And far from being a mere routine calibration of
harvest guidelines, the negative ecological effects of this change – which
allows for a doubling of cougar take in some GMUs 32 – will be substantial as
well. As described above, the scientific evidence predicts that the Final Rule
will destabilize cougar social structures in the fourteen areas subject to the
increase, resulting in imbalanced age and sex distribution, increased humancougar conflict, and increased livestock depredation. 33
ii. Inadequate Notice to Stakeholders
A second factor to be considered in determining whether a final rule is
“substantially different” than a proposed rule is “the extent to which a
reasonable person affected by the adopted rule would have understood that
the published proposed rule would affect his or her interests.” 34 This factor is
meant to ensure that stakeholders are put on adequate notice of a possible
rule change affecting their interests so that they can exercise their public
input rights.
Here, many stakeholders, including non-consumptive users like
Petitioners and their members, were not put on adequate notice (or any

For instance, the new maximum harvest guideline for GMUs 113, 145, 166, 175, 178 is
double the prior minimum guideline. See Final Rule, sec. 2.
33 These impacts were not considered in the Department EIS, which only analyzed the
environmental impact of the Department’s recommended 12-16% harvest guideline. See
Department EIS, supra n. 6, at 109-10.
34 RCW § 34.05.340(2).
32

13

 notice at all) of the possibility of a quota increase. Not only did the Proposed
Rule maintain harvest guideline numbers at status quo levels, it expressly
set out in its statement of purpose that “[t]he harvest guideline represents a
sustainable 12-16 percent harvest rate for each hunt area.” 35 Absent
supplemental notice, which was not provided in this rulemaking, a
reasonable party would not have predicted that such a proposal would be
stretched, at the eleventh hour, into an increase to 17-21 percent harvest
rates. Stakeholders like Petitioners and other conservation groups were
consequently denied their right to meaningful public input on the quota
increase. And other interested parties – including Washingtonians who may
now be subject to heightened wildlife conflict and livestock depredation
because they happen to live near the increased harvest areas – were denied
their right to notice and comment on a policy change that will concretely
affect their interests.
b. Even if the Final Rule Did Not Substantially Differ From the
Proposed Rule, the Commission Has the Authority and
Mandate to Amend the Final Rule
WAPA also provides “any person” the right to “petition an agency
requesting the…amendment…of any rule.” 36 If the Commission determines
that the Final Rule is not substantially different from the Proposed Rule
The proposal continued, “Recently published studies suggest that a 12-16 percent harvest
rate of a local cougar population is the maximum harvest rate that still has a high
probability for maintaining a stable cougar population along with stable adult male
territorial behavior. Harvest rates in excess of 16 percent can result in declines in core
populations of breeding females, and excessive adult male harvest rates result in the loss of
adult male territorial behavioral, which acts as a regulatory mechanism for local cougar
numbers.” Proposed Rule, at 10.
36 RCW § 34.05.330(1).
35

14

 pursuant to RCW § 34.05.340, Petitioners request in the alternative that the
Commission consider this Petition to amend WAC 232-28-297.
The petitioned amendment is plainly within the authority of the
Commission and Department to implement pursuant to its broad rulemaking
power. 37 Indeed, the Department is not only authorized but mandated to
“conserve the wildlife… in a manner that does not impair the resource” and
may only “authorize the taking of wildlife… in manners or quantities” that
“[do] not impair the supply of these resources.” 38 These mandates counsel the
calibration of hunting quotas in consultation with experts and in accordance
with the best available science in order to avoid type of deleterious
consequences described above. The Final Rule violates the legislature’s intent
by raising cougar quotas far beyond the level recommended by expert
biologists, risking well-documented “impairment” 39 of a wildlife resource held
in the public trust. Furthermore, the scientific documentation militating
against the Final Rule is so overwhelming and the Commission’s justification
so weak that the Final Rule may be considered “arbitrary and capricious” or
“not supported by evidence that is substantial” in violation of WAPA. 40
In addition to satisfying the Commission’s statutory mandates under
its authorizing statute and WAPA, the proposed amendment would also bring
Department regulations into alignment with its prior reasonable policy

RCW §§ 77.04.012; 77.04.090.
RCW § 77.04.012.
39 Id.
40 RCW § 34.05.570(2)(c).
37
38

15

 positions. The Department has long prioritized using the best available
science in its wildlife-management decisions, as evidenced by the Proposed
Rule and the 2015-2021 Game Management Plan Environmental Impact
Statement, both of which proposed harvest guidelines of 12-16 percent
premised on a careful analysis of the studies referenced in this Petition. 41
Adopting the petitioned amendment would restore consistency to a
management program that had previously been defined by rational, sciencebased decision-making.
IV. Conclusion
The Final Rule was undoubtedly a procedural and scientific misstep
and the Commission is legally required to take action to bring cougar hunting
quotas back to a sustainable level. To protect the right of Washingtonians to
meaningful input on the management of their wildlife, and ensure the
scientific integrity of the Department’s cougar program, Petitioners
respectfully urge the Commission to initiate a rulemaking process to amend
WAC 232-28-297.
Petitioners remain available to address specific questions or provide
additional documentation as requested.
Sincerely,

Supra, n. 6. While this Petition does not allege specific violations of the Washington State
Environmental Policy Act (“SEPA”), it does note a major and legally actionable discrepancy
between the statewide 12-16% harvest guidelines whose environmental impacts were
discussed in the Department EIS and the content of the Final Rule.
41

16

 Anna Frostic
Senior Attorney
The Humane Society of the United
States

Mitch Friedman
Executive Director
Conservation Northwest

Tim Dunbar
Executive Director
Mountain Lion Foundation

Collette Adkins
Senior Attorney
Center for Biological Diversity

Diane K. Gallegos
Executive Director
Wolf Haven International

Penelope Maldonado
Managing Director
The Cougar Fund

Brooks Fahy
Executive Director
Predator Defense

Mike Petersen
Executive Director
The Lands Council

Gary M. Koehler, Ph. D.
Research Scientist for Carnivore
Investigations (Retired)
Washington Department of Fish
and Wildlife

Timothy J. Coleman
Executive Director
Kettle Range Conservation
Group
17

 Enclosure: Proposed amendment to WAC Section 232-28-297; CD
containing each of the studies cited herein and which are expressly submitted
into the administrative record.
References Cited
Cooley, H. S., R. B. Wielgus, G. M. Koehler, H. S. Robinson and B. T.
Maletzke. "Does Hunting Regulate Cougar Populations? A Test of the
Compensatory Mortality Hypothesis." Ecology 90, no. 10 (2009): 29132921.
Cooley, H. S., R. B. Wielgus, G. Koehler and B. Maletzke. "Source
Populations in Carnivore Management: Cougar Demography and
Emigration in a Lightly Hunted Population." Animal Conservation 12,
no. 4 (2009): 321-328.
Peebles, Kaylie A., Robert B. Wielgus, Benjamin T. Maletzke and Mark E.
Swanson. "Effects of Remedial Sport Hunting on Cougar Complaints
and Livestock Depredations." Plos One 8, no. 11 (2013).
Robinson, H. S., R. B. Wielgus, H. S. Cooley and S. W. Cooley. "Sink
Populations in Carnivore Management: Cougar Demography and
Immigration in a Hunted Population." Ecological Applications 18, no. 4
(2008): 1028-1037.
Ruth, Toni and Kerry Murphy. "Competition with Other Carnivores for
Prey." In Cougar: Ecology and Conservation, edited by Maurice
Hornocker and Sharon Negri, 163-172. Chicago and London:
University of Chicago Press, 2010.
Stoner, D., M. , M.L. Wolfe and D. Choate. "Cougar Exploitation Levels in
Utah: Implications for Demographic Structure, Population Recovery,
and Metapopulation Dynamics." Journal of Wildlife Management 70,
(2006): 1588-1600.
Treves, Adrian. "Hunting for Large Carnivore Conservation." Journal of
Applied Ecology 46, (2009): 1350-1356.

18

 Wielgus, R. B., D. E. Morrison, H. S. Cooley and B. Maletzke. "Effects of Male
Trophy Hunting on Female Carnivore Population Growth and
Persistence." Biological Conservation 167, (2013): 69-75.

19