Endangered and Threatened Wildlife and Plants; Withdrawal of the Proposed Rule for the North American Wolverine, 64618-64648 [2020-19538]
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Federal Register / Vol. 85, No. 198 / Tuesday, October 13, 2020 / Proposed Rules
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R6–ES–2016–0106;
FF09E21000 FXES11110900000 201]
RIN 1018–BB78
Endangered and Threatened Wildlife
and Plants; Withdrawal of the
Proposed Rule for the North American
Wolverine
Fish and Wildlife Service,
Interior.
ACTION: Proposed rule, withdrawal.
AGENCY:
We, the U.S. Fish and
Wildlife Service, withdraw the
proposed rule to list the distinct
population segment (DPS) of the North
American wolverine (Gulo gulo luscus)
occurring in the contiguous United
States as a threatened species under the
Endangered Species Act of 1973, as
amended (Act). This withdrawal is
based on our conclusion that the factors
affecting the species as identified in the
proposed rule are not as significant as
believed at the time of the proposed
rule. We base this conclusion on our
analysis of current and future threat
factors. We also find that North
American wolverines occurring in the
contiguous United States do not qualify
as a DPS. Therefore, we are withdrawing
our proposal to list the wolverine within
the contiguous United States as a
threatened species.
DATES: The proposed rule that
published February 4, 2013 (78 FR
7864), to list the distinct population
segment of the North American
wolverine occurring in the contiguous
United States as a threatened species is
withdrawn on October 13, 2020.
ADDRESSES: The withdrawal of our
proposed rule, comments, and
supplementary documents are available
on the internet at https://
www.regulations.gov at Docket Nos.
FWS–R6–ES–2012–0107 and FWS–R6–
ES–2016–0106.
FOR FURTHER INFORMATION CONTACT: Jodi
Bush, Field Supervisor, U.S. Fish and
Wildlife Service, Montana Ecological
Services Office, (see ADDRESSES).
Persons who use a telecommunications
device for the deaf may call the Federal
Relay Service at 800–877–8339.
SUPPLEMENTARY INFORMATION:
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SUMMARY:
Executive Summary
Why we need to publish a rule. Under
the Act, if we determine that a species
may be an endangered or threatened
species throughout all or a significant
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portion of its range, we are required to
promptly publish a proposal in the
Federal Register and make a
determination on our proposal within 1
year. To the maximum extent prudent
and determinable, we must designate
critical habitat for any species that we
determine to be an endangered or
threatened species under the Act.
Listing a species as an endangered or
threatened species and designation of
critical habitat can only be completed
by issuing a rule.
What this document does. We
withdraw the proposed rule to list the
DPS of the North American wolverine
occurring in the contiguous United
States as a threatened species under the
Act.
The basis for our action. Under the
Act, we may determine that a species is
an endangered or threatened species
because of any of five factors: (A) The
present or threatened destruction,
modification, or curtailment of its
habitat or range; (B) overutilization for
commercial, recreational, scientific, or
educational purposes; (C) disease or
predation; (D) the inadequacy of
existing regulatory mechanisms; or (E)
other natural or manmade factors
affecting its continued existence. We
have determined that the factors
affecting the species as identified in the
proposed rule (loss of habitat due to
climate change) are not as significant as
believed at the time of the proposed
rule. We also find that North American
wolverines occurring in the contiguous
United States do not qualify as a DPS.
Peer review. In accordance with our
July 1, 1994, peer review policy (59 FR
34270; July 1, 1994), the Service’s
August 22, 2016, Director’s Memo on
the Peer Review Process, and the Office
of Management and Budget’s December
16, 2004, Final Information Quality
Bulletin for Peer Review (revised June
2012), we sought the expert opinions of
four appropriate specialists regarding
the species status assessment report. We
received responses from four specialists,
which informed this proposed rule. The
purpose of peer review is to ensure that
our listing determinations are based on
scientifically sound data, assumptions,
and analyses. The peer reviewers have
expertise in the biology, habitat, and
threats to the species. Results of this
structured peer review process can be
found at https://www.fws.gov/mountainprairie/science/peerReview.php. A draft
analysis was also submitted to our
Federal, State, and Tribal partners for
scientific review. In preparing this
withdrawal, we incorporated the results
of these reviews in the final SSA report,
as appropriate.
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During the reopening of the public
comment periods for the proposed
listing rule, we requested any new
information and announced that we
initiated a new and comprehensive
status review of the North American
wolverine to determine whether the
species meets the definition of an
endangered or threatened species under
the Act, or whether the species is not
warranted for listing. The wolverine
SSA report provides the scientific basis
for the decision to withdraw the
proposed listing rule for the DPS of
wolverine occurring in the contiguous
United States. Both new and updated
information and analyses presented in
the wolverine SSA report, summarized
below in support of our listing
determination, along with public
comment, have also prompted us to
reevaluate our previous assessment of
the DPS with respect to wolverine in the
contiguous United States.
Supporting Documents
A team prepared a Species Status
Assessment (SSA) for the North
American Wolverine (Gulo gulo luscus)
(Service, 2018) (hereafter referred to as
the wolverine SSA report). The SSA
team was composed of U.S. Fish and
Wildlife Service biologists, who
consulted with other species experts.
The wolverine SSA report represents a
compilation of the best scientific and
commercial data available concerning
the status of the species, including the
impacts of past, present, and future
factors (both negative and beneficial)
affecting the wolverine. The wolverine
SSA report underwent independent
peer review by scientists with
experience with mesocarnivores and
their conservation and management,
genetics, population modeling, and
climate change. The wolverine SSA
report and other materials relating to
this proposal can be found on the
Mountain-Prairie Region website at
https://www.fws.gov/mountain-prairie/
and at https://www.regulations.gov
under Docket No. FWS–R6–ES–2016–
0106.
Previous Federal Actions
Please refer to the proposed listing
rule for the wolverine (78 FR 7864;
February 4, 2013) for a detailed
description of previous Federal actions
concerning the wolverine prior to 2013.
On February 4, 2013, we published a
proposed rule to list the DPS of
wolverine occurring in the contiguous
United States as threatened, under the
Act, with a proposed rule under section
4(d) of the Act that outlines the
prohibitions necessary and advisable for
the conservation of the wolverine (78 FR
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7864). We also published a proposed
rule on February 4, 2013, to establish a
nonessential experimental population
(NEP) area for the North American
wolverine in the Southern Rocky
Mountains of Colorado, northern New
Mexico, and southern Wyoming (78 FR
7890). On October 31, 2013, we
reopened the comment period on the
proposed listing rule for an additional
30 days (78 FR 65248).
Following publication of the 2013
proposed rules, there was scientific
disagreement and debate about the
interpretation of the habitat
requirements for wolverines and the
available climate change information
used to determine the extent of threats
to the DPS. Based on this substantial
disagreement regarding the sufficiency
or accuracy of the available data
relevant to the proposed listing, on
February 5, 2014 (79 FR 6874), we
announced a 6-month extension of the
final determination of whether to list
the wolverine DPS as a threatened
species. We also reopened the comment
period on the proposed rule to list the
contiguous United States DPS of the
North American wolverine for 90 days.
On August 13, 2014, we withdrew the
proposed rule to list the DPS of the
North American wolverine as a
threatened species under the Act (79 FR
47522). This withdrawal was based on
our conclusion that the factors affecting
the DPS as identified in the proposed
rule were not as significant as believed
at the time of the proposed rule’s
publication in 2013. As a result, we also
withdrew our associated proposed rule
under section 4(d) of the Act contained
in the proposed listing rule and
withdrew the proposed NEP designation
under section 10(j) of the Act for the
southern Rocky Mountains.
In October 2014, three complaints
were filed in the District Court for the
District of Montana by Defenders of
Wildlife, WildEarth Guardians, Center
for Biological Diversity, and other
organizations challenging the
withdrawal of the proposal to list the
North American wolverine DPS.
Numerous parties intervened in the
litigation. These three cases were
consolidated, and on April 4, 2016, the
court issued a decision. The court
granted plaintiff’s motion for summary
judgment with respect to the Service’s
determination regarding (1) the threat
posed to the wolverine by the effects of
climate change at the reproductive
denning scale, (2) the threat posed to the
wolverine by small population size and
lack of genetic diversity, and (3) the
application of the significant portion of
its range policy to the wolverine. As a
result of the court order, the August 13,
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2014, withdrawal (79 FR 47522) was
vacated and remanded to the Service for
further consideration consistent with
the order. As documented in the SSA
report, the Service conducted additional
analyses and reviewed new literature
regarding climate change effects at the
denning scale (see pages 73–99 of the
SSA report) and included additional
life-history information relevant to this
potential stressor (see pages 25–39).
With regard to population size, we also
provide in the SSA report an analysis of
information on wolverine population
abundance and distribution (to date)
and have included a discussion of
population structure (genetics, effective
population size) in the context of the
species’ known genetic variability (see
pages 44–50). Finally, in this
withdrawal, we have provided an
updated significant portion of its range
analysis (see discussion below).
In effect, the court’s action returned
the process to the proposed rule stage,
and the status of the wolverine under
the Act reverted to that of a proposed
species for the purposes of consultation
under section 7 of the Act. On October
18, 2016, we published a notice (81 FR
71670) reopening the comment period
on the February 4, 2013, proposed rule
(78 FR 7864) to list the DPS of
wolverine occurring in the contiguous
United States as threatened, under the
Act. We also requested new information
and announced that we initiated a new
and comprehensive status review of the
North American wolverine, to
determine whether the species meets
the definition of an endangered or
threatened species under the Act, or
whether the species is not warranted for
listing. The wolverine SSA report
provides the scientific basis for the
decision to withdraw the proposed
listing rule for the DPS of wolverine
occurring in the contiguous United
States. Both new and updated
information and analyses presented in
the wolverine SSA report, summarized
below in support of our listing
determination, along with public
comment, have prompted us to
reevaluate our previous assessment of
the DPS (presented in our 2013
proposed listing rule, which in turn
relied on the DPS analysis completed in
our 2010 12-month finding) with respect
to wolverine in the contiguous United
States.
Summary of Comments and
Recommendations
As stated above in the Previous
Federal Actions section, on October 18,
2016 (81 FR 71670), we opened a public
comment period on our February 4,
2013, proposed rule (78 FR 7864) to list
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the DPS of wolverine occurring in the
contiguous United States as threatened.
We also contacted appropriate Federal
and State agencies, scientific experts
and organizations, Tribes, and other
interested parties and invited them to
comment on the proposed rule. Many of
the comments we received from State
agencies during our notice for reopening
the comment period (81 FR 71670) were
similar to those received for the
previously proposed rule (78 FR 7864).
All substantive information provided
during comment periods has either been
incorporated directly into this final
determination or is addressed below.
Public Comments
(1) Comment: We received several
public comments claiming that the
North American wolverine faces
increasing threats from the effects of
climate change, particularly habitat loss
due to declining snow pack.
Our Response: As discussed in the
wolverine SSA report, we recognize that
current climate trends and future (2055
and later) climate model projections
indicate warming temperatures for
much of western North America and
changes to snow pack conditions. Our
review of the literature found that,
overall, higher elevation areas (e.g.,
Rocky Mountains, Sierra Nevada
Mountains) are more resilient to
projected changes in temperature and
precipitation as compared to lower
elevations (Wobus et al. 2017, p. 12). In
general, models indicate higher
elevations, where documented historical
wolverine denning has occurred, will
retain more snow cover than lower
elevations, particularly in early spring
(April 30/May 1). We present in the
wolverine SSA report a summary of
new, fine-scale analysis of future snow
persistence in two regions of the
western United States, Glacier National
Park and Rocky Mountain National
Park. The two regions studied include a
high-latitude area near tree line within
Glacier National Park, where tree line
occurs at (∼1,800 to 2,100 meters (m)
(5,906 to 6,890 feet (ft))) that is currently
occupied by wolverines; and a lower
latitude area within Rocky Mountain
National Park (occupied by a single
male wolverine from 2009 to at least
2012, but not known to be currently
occupied), where tree line occurs at
higher elevation (∼3,500 m (11,483 ft))
(Ray et al. 2017, p. 2). These sites were
chosen to bracket the range of latitude
and elevation wolverines currently
occupy in the contiguous United States
(Ray et al. 2017, p. 2). This effort built
upon previous model projections
presented in McKelvey et al. (2011), but
with significant differences such as finer
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spatial resolution, incorporation of
slope and aspect, snow depth estimates,
additional years of historical data, and
wider temporal analyses of snow
persistence (April–June). Details of this
modeling exercise are presented in Ray
et al. (2017), and summarized in the
SSA report. That analysis indicates
significant areas (several hundred
square kilometers (km2)/square miles
(mi2) for each study area) of future snow
(greater than 0.5 m (20 inches (in) in
depth) will persist on May 1 at
elevations currently used by wolverines
for denning. This is true, on average,
across the range of climate models used
out to approximately year 2055.
(2) Comment: We received several
public comments during our request for
information claiming that low
population size (and small effective
population size) warrant listing of the
North American wolverine as
threatened or endangered.
Our Response: Small populations in
and of themselves do not constitute a
threat such that a species would be
endangered or threatened. When
evaluating species status, we take into
consideration the species’ life history,
population dynamics, and other impacts
to populations and species to determine
if small population dynamics increases
the species’ vulnerability to extinction
such that listing as threatened or
endangered is warranted. Wolverines
are difficult animals to survey, and
populations occur in naturally low
densities across their North American
range due, in large part, to their need for
large, exclusive territories. At the
present time, there is no reliable
estimate of the number of wolverines
that currently occupy or previously
occupied the contiguous United States,
nor are there reliable quantitative
estimates of wolverine population
trends in the contiguous United States.
The often-cited population estimate of
318 wolverines (range: 249–626) in the
contiguous United States is derived
from a habitat modeling exercise
presented in Inman et al. (2013). That
publication also provided a model
estimate of potential wolverine capacity
of 644 (range: 249–626). However, both
of these estimates did not consider
important spatial considerations related
to wolverine behavior, such as
territoriality, relative to wolverine
populations. Despite the paucity of
information regarding wolverine
populations, the SSA analysis is a
thorough examination of all of the
available population information.
As discussed in the wolverine SSA
report, preliminary field results from a
recent (2016–2017) occupancy study in
four western States (Idaho, Montana,
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Washington, and Wyoming) and from a
pilot occupancy study in Wyoming
(2015–2016) indicate detections of
wolverines in areas where they would
be expected to be found, but also no
detections in areas where they are
known to occur (e.g., areas within
Glacier National Park) (see Service 2018,
Appendix B for a descriptive map). To
date, this study reports a total of 86
photographic detections through
camera-trapping and 157 wolverine hair
samples collected for genetic analysis. It
has not yet been determined from the
camera-trap images or hair samples how
many of the detections are unique
individuals. Preliminary analysis of the
study results indicates an average
estimated probability of occupancy of
0.42 suggesting that wolverines used
nearly half of all sites during the study
period (Montana FWP, pers. comm.,
2017); however, the study did not
encompass all potential wolverine
habitat in the western United States
(Service 2018, Appendix B). For
example, wolverines have also been
recently detected in northeastern
Oregon (as of 2017) and in parts of
Grand Teton National Park (two records
during the winter of 2017), which were
not included in the surveyed study
cells. Our SSA report presents a visual
summary of these recent detections
(Service 2018, Figure 3). Although the
sum of these reports cannot confirm
previous estimates of population size or
verify population trends, they offer
recent evidence that wolverines
continue to be observed across a large
area of the western United States.
The 2013 proposed rule presented an
effective population size estimate from
a publication by Schwartz et al. (2009),
which estimated a summed effective
population size of 35, with credible
limits from 28 to 52 (Schwartz et al.
2009, p. 3,226). As described in the SSA
report, the study used wolverine
samples from the main part of the Rocky
Mountain wolverine populations and
did not include subpopulations from
two other mountain regions in Montana,
and samples were missing from other
parts of the wolverine range in Idaho
and other areas of Montana. Thus, the
analysis missed both wolverine
subpopulations and individuals, which
would underestimate the results for this
type of analysis.
In the wolverine SSA report, we
provide a contextual discussion of the
effective population size concept,
particularly in the context of genetic
studies related to the phylogeographic
history of the North American
wolverine that were not well described
in the 2013 proposed rule. In sum, the
spatial distribution of genetic variability
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currently observed in wolverines in
North America appears to be a reflection
of a complex history in which
population abundance has fluctuated
since the time of the last glaciation with
insufficient time passing since human
persecution since at least the 1700s to
allow for full recovery of wolverine
densities (Cardinal 2004, pp. 23–24;
Zigouris et al. 2012, p. 1,554). This
history and the fact that wolverines in
the contiguous United States occupy the
southern periphery of the species’ entire
North American range are important
considerations in estimating and
interpreting current wolverine
distribution and abundance. The
wolverine SSA report also presents
information from genetic and
observational studies that provide
support for wolverine movement across
the international border of the
contiguous United States and Canada. In
the 2013 proposed rule, we stated there
is an apparent lack of connectivity
between wolverine populations in
Canada and the United States based on
genetic data (78 FR 7864; February 4,
2013). We now consider wolverines that
occupy the contiguous United States to
be genetically continuous with
wolverines in adjacent Canadian
provinces. A small effective population
size would be more of a concern if the
population was in isolation; however,
wolverines in the contiguous United
States are not genetically isolated from
wolverines in Canada. For more
information, see the Small Total
Population Size and Effective
Population Size sections under Distinct
Population Segment below.
(3) Comment: We received several
public comments during our request for
information claiming that North
American wolverine face threats from
indiscriminant trapping in the
contiguous United States, or are
threatened by incidental trapping.
Our Response: At the present time,
trapping or hunting of wolverine is not
allowed in any State within the range of
the wolverine (with the exception of
Alaska). Legal protections for
wolverines are codified in western State
laws and regulations and include:
Endangered in Colorado, threatened in
California and Oregon, candidate
species in Washington, non-game
species protections in Idaho and
Wyoming, a species of concern and a
furbearer with a closed season in
Montana, and protection from
collection, importation, and possession
in Utah. Since 2013, there has been a
zero quota for trapping or harvest of
wolverine in Montana.
Incidental trapping of wolverines has
been documented in the contiguous
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United States (as recently as December
2017), though not all events have
resulted in mortality. In the wolverine
SSA report, we provide a summary of
the number of wolverines that have
been incidentally trapped in Idaho (18
since 1965, including 6 known to be
released alive and 7 known mortalities),
Montana (4 since 2013, 3 mortalities
and 1 released unharmed), and
Wyoming (2 since 1996, 1 mortality and
1 released unharmed) (Service 2018, p.
66). Both Idaho and Montana are
implementing trapper education
programs to minimize nontarget
wolverine captures.
As discussed in the SSA report,
regulated trapping and hunting of
wolverines occurs in parts of Alaska and
Canada, and appears to be sustainable
based on population and density
estimates.
(4) Comment: We received several
public comments identifying potential
threats to wolverines from winter
recreation activities, such as
snowmobiling and back-country skiing.
Our Response: In the SSA report, we
present a summary of winter recreation
studies (Heinemeyer et al. 2015;
Heinemeyer et al. 2017), future
projections of winter recreation activity
in the contiguous United States (White
et al. 2014), and projections of
snowpack relative to changes in the
length of the winter recreation season
(Wobus et al. 2017). We reported results
from Heinemeyer (2016, pers. comm.)
indicating a behavioral response to
recreation activities, but also
maintenance of home ranges within
some areas of relatively high recreation
activity over several years. The study
has not yet been able to determine
whether resident wolverines are
reproductively successful due to the
limited monitoring information
available for reproducing female
wolverines. Nor was the study able to
determine if recreational activities had a
negative impact on wolverine
reproductive success.
We also note here that we received
the final report of this multiyear study
(Heinemeyer et al. 2017) in midDecember 2017 (results of this study
were recently published (Heinemeyer et
al. 2019)), which was after we submitted
the draft SSA report for review to four
peer reviewers and to our State, Federal,
and Tribal partners. Much of the report
presents a modeling exercise to evaluate
wolverine behavior patterns with winter
recreation activities. The study found
that wolverines maintained multiyear
home ranges, and the authors suggest
that wolverines are able to tolerate
winter recreation at some scales
(Heinemeyer et al. 2017, p. iv;
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Heinemeyer et al. 2019, p. 16). The
study described habitat selection as
complex for female wolverines and was
likely driven by a combination of abiotic
(snow, cold) and biotic (predator
avoidance, food availability) factors
(Heinemeyer et al., 2017, p. 36;
Heinemeyer et al. 2019, p. 16). This
study did not assess demographic
effects, fitness effects, or population
level effects of winter recreation on
wolverines (Heinemeyer et al. 2019, p.
17 and 19). As discussed in the
wolverine SSA report, management
measures being implemented in areas
within the wolverine’s current extent of
occurrence include road closures to
minimize disturbance to wildlife on
lands managed by the U.S. Forest
Service and National Park Service
(Service 2018, p. 61 and Appendix F).
In addition, management strategies are
identified in State Wildlife Action Plans
(e.g., Oregon, Montana, Idaho) to
address potential impacts from
recreation to the wolverine. Although
we did not rely on these conservation
measures to support our decision, they
do provide some level of protection to
address potential impacts from
disturbance from winter recreation
activity and mortality from roads.
(5) Comment: We received public
comments claiming that wolverines are
dependent on deep snow for survival
and expressing concern for future
changes in snow pack due to the effects
of climate change.
Our Response: After reviewing studies
not previously considered and the
results of new studies/publications
made available after the 2013 and 2014
proposed rules (e.g., Aronsson 2017,
Aronsson and Persson 2016, Aronsson
et al. 2017, Magoun et al. 2017, Persson
et al. 2017, Stewart et al. 2016, Webb et
al. 2016, see complete list of citations in
the wolverine SSA report), we present
in the SSA report a detailed discussion
of the North American wolverine’s
physiology and other life-history
characteristics (e.g., reproductive
behavior). This summary speaks to
several presumed aspects of the
relationship of denning behavior and
other needs of this species regarding the
presence of persistent spring snow. As
summarized below, we now know that
wolverines can and have denned
outside of heavy snowpack, multiple
factors play a role in den site selection,
females will move dens as young
become mobile, and areas of significant
snowpack will likely persist in the
future in areas where wolverines are
known to den at levels that will
continue to support wolverines.
Denning habitat for the wolverine
varies over its range and is dependent
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on local and regional environmental
conditions (e.g., topographic and other
structural features) and biotic (e.g.,
availability of prey; protection from
predators) factors. Reproductive (natal)
dens are not always excavated in deep
snow, particularly in boreal forest
habitats (Dawson et al. 2010; Novikov
1962; Webb et al. 2016; Jokinen 2019.),
and have been observed in spruce tree
root balls, logging slash piles, and
beaver dens/dams. In the contiguous
United States, dens are found at high
elevations, often in talus slopes, which
provides conditions for protection and
food caching (e.g., restricted access and
cold temperatures). Our review of
studies of wolverine denning activity
found no quantitative data reporting
snow depth at the den site when
wolverines abandon the den. More
importantly, wolverine reproductive
success has not been studied relative to
a number of abiotic and biotic
conditions, including depth and
temporal aspect of spring snow cover.
Wolverines begin shifting den
locations in late April, when young
become more mobile and reliant on
solid food brought to them by the
mother (Aronsson 2017, p. 46; Aubry et
al. 2016, p. 24). The bioclimatic model
presented in Copeland et al. (2010) was
used to test wolverine distribution at a
broad scale based on climate variables,
including spring snow cover, using May
15 conditions. They then tested their
hypothesis by comparing and
correlating the location of wolverine
dens across their circumboreal range,
and telemetry locations from wolverine
studies in North America and
Scandinavia (Copeland et al. 2010, p.
234). Since that publication, wolverines
and wolverine dens have been observed
outside the boundaries defined by the
model presented in Copeland et al.
(2010) (e.g., Webb et al. 2016, Webb
2017 pers. comm., Persson 2017, pers.
comm.). While these observations are
found at higher latitudes in the
circumboreal region, they also indicate
wolverines and wolverine dens are
observed in environments that are not
characterized by several feet of spring
snow on May 15. In sum, Copeland et
al. (2010) provided a fairly accurate
assessment of where wolverine
populations are expected to occur, but
it did not evaluate (model) snow
persistence at the den site scale based
on location and denning period.
In the SSA report, we present an
analysis of 34 wolverine den locations
(years 2002–2015) from studies in the
western contiguous United States
relative to ‘‘melt out’’ dates, which
represents the first day of an 8-day
satellite (Moderate Resolution Imaging
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Spectroradiometer (MODIS)) composite
of when the den switches from ‘‘snow’’
to ‘‘no snow’’ at a 500-by-500-m (1,640by-1,640-ft) spatial resolution. For natal
den locations, the range of the melt-out
dates was from May 25 to June 12,
which is considerably later than the
May 15 date used in the Copeland et al.
(2010) analysis. The estimated melt-out
dates indicate that snow is persistent at
these locations past the time when
young wolverines are generally moving
out of natal dens (i.e., late April).
The Copeland et al. (2010) snow
model was then used by McKelvey et al.
(2011) to model effects of climate
change to wolverine habitat in the
western United States to develop
projections of habitat loss. This
modeling exercise used May 1 snow
presence as a proxy for May 15 snow
disappearance and a spatial resolution
of 36.3 km2 (14 mi2)), which is not
relevant at the at the den site scale. As
described in our Response to Comment
#1, in the SSA report, we presented a
finer scale analysis (0.0625 km2 (0.24
mi2)) for two study areas (Glacier
National Park and Rocky Mountain
National Park) that focused directly on
May 15, in addition to the presence or
absence of snow on May 1 and April 15
in our evaluation of the effects of
climate change to snowpack. These
dates are more relevant to wolverine
life-history needs. We also modeled the
depth of ‘‘significant’’ snow (0.5 m (1.64
ft)) on these dates. We found that large
areas (several hundred km2/mi2 for each
study area) of future snow (greater than
0.5 m (20 in) in depth) are projected to
persist on May 1 at elevations currently
used by wolverines for denning. This is
true, on average, across the range of
climate models used out to
approximately year 2055. We recognize
that wolverines are difficult to study
and evaluation of denning habitat and
behavior is challenging. Additional
research is needed to evaluate other
potential physical and biotic variables
that could be important in defining
wolverine distribution and den
locations. These additional variables
include: prey availability, risk of
predation, den-site scale factors such as
structure/snow conditions, and
temporal use of dens.
(6) Comment: We received public
comments identifying the need for
additional research and
recommendations for conservation
measures for the North American
wolverine, including estimates of
population size and further evaluation
of life-history characteristics, and
recommendations for conservation
measures.
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Our Response: We appreciate the
comments acknowledging the need for
additional studies as well as the
difficulties in studying wolverines given
its occupation of remote habitats in the
contiguous United States. In the
wolverine SSA report, we provide a
summary of the preliminary field and
genetic results from the recent Western
States Wolverine Conservation Project
(WSWCP)–Coordinated Occupancy
Study in four western (contiguous
United) States, as well as results from
several new studies presented in peerreviewed publications and in other
reports from Canada and Scandinavia.
As discussed in the SSA report, the
Western States Wolverine Working
Group is continuing to develop studies
to evaluate wolverine population
distribution and occupancy, and
connectivity across four western States.
(7) Comment: We received additional
comments from the public including the
need for collaboration with local
government and community
stakeholders and use of best available
science in developing the proposed rule.
Our Response: During our preparation
of the wolverine SSA report, we
coordinated extensively with many
wolverine researchers in the United
States (including Alaska), Canada, and
Scandinavia. Those communications are
identified in our References Cited
section of the wolverine SSA report.
Their expertise, insights, and published
or soon-to-be published research papers
were invaluable in ensuring that we
used the best available science in
preparing the new status review. We
also communicated with biologists at
several State and Federal agencies to
ensure that we had incorporated the
most recent wolverine detections in the
western United States. The wolverine
SSA report was sent to four
independent peer reviewers, selected by
an outside contractor, and those nonattributable comments were
incorporated, to the extent possible, in
the final document. We also reviewed
comments received from the public and
previous peer reviewers during our
request for comments for our previous
proposed rule and considered the
information provided (78 FR 7864;
February 4, 2013) during the
preparation of the wolverine SSA
report. As a result, this determination is
based upon the best scientific and
commercial data available to us, as
required by the Act.
(8) Comment: We also received public
comments recommending that the North
American wolverine not be listed as
threatened or endangered under the Act.
One commenter stated that State
wildlife agencies are capable of
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managing the species and are able to
provide protections that ensure
continued population growth towards
population objectives established by
these agencies and that mandates of
various Federal resource management
agencies provide a commitment to
managing wildlife habitat in a way that
benefits all wildlife species, including
wolverines and other forest carnivores.
Our Response: We acknowledge that
some members of the public support our
decision to withdraw our proposed rule
to list the North American wolverine as
threatened under the Act. In the
wolverine SSA report (Service 2018,
Appendix G), we provide a summary of
the regulatory protections provided by
western States and Federal agencies as
well as management measures being
implemented to conserve the wolverine
and its habitat. Legal protections in the
contiguous United States include State
listing in California and Oregon
(threatened), endangered in Colorado, a
candidate species in Washington, nongame species protections in Idaho and
Wyoming, a species of concern and
furbearer with a closed season in
Montana, and protection from
collection, importation, and possession
in Utah. Trapping or hunting of
wolverines is currently prohibited in the
contiguous United States.
(9) Comment: In response to our
request for information in our public
notice, several members of the public
provided specific information related to
personal wolverine sightings of the
North American wolverine in the
contiguous United States (e.g., New
Mexico, Wyoming), and information
regarding past and ongoing research
studies of the species in the western
United States and in Canada.
Our Response: We appreciate the
personal observations provided and
encourage members of the public to
document sightings of the North
American wolverine with photographs
and provide additional details to State
wildlife agencies. Information we
received regarding results from research
studies has been incorporated, as
appropriate, in the final wolverine SSA
report.
(10) Comment: We received
comments from several organizations
that support the listing of the North
American wolverine and designation of
critical habitat. Threats cited include
concerns related to migration, habitat
loss and connectivity related to threats
from effects of climate change, nontarget
trapping pressures, road mortality and
other effects of roads (e.g., noise,
pollution, fragmentation of habitat),
motorized recreation and traffic in
wildlife corridors, timber sales and
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associated roads, and effects of
snowmobile traffic (habitat
fragmentation and pollution, and
change in behavior).
Our Response: As discussed in the
Risk Factors for the North American
Wolverine section below, we identified
several potential stressors that may be
affecting the species and its habitat
currently or in the future, including
impacts associated with climate change
effects. We recognize there is limited
information available for the wolverine,
including population estimates and
abundance trends. Based on the best
available information, demographic
risks to the species from either known
or most likely potential stressors (i.e.,
disturbance due to winter recreational
activities, other human disturbances,
effects of wildland fire, disease,
predation, overutilization, genetic
diversity, small population effects,
climate change, and cumulative effects)
are low based on our evaluation of the
best available information as it applies
to current and potential future
conditions for the wolverine and in the
context of the attributes that affect the
needs of the species (Service 2018, p.
103). Thus, we determined that, based
on the best available information, the
North American wolverine in the
contiguous United States does not meet
the definition of a threatened species or
an endangered species under the Act.
(11) Comment: We received public
comments stating that protection of
North American wolverines in the
contiguous United States is needed
under the Act in order to provide
resources and attention needed for
research and monitoring, to better
understand threats, and sustain
wolverines into the future. The
commenter also stated that federally
sponsored wolverine reintroduction in
Colorado will help increase chances of
long-term species survival.
Our Response: We appreciate the
recognition of the need for continued
resources for research and monitoring.
However, we base our listing decisions
on a determination of whether the
species meets the Act’s definitions of a
threatened species or an endangered
species. Regardless, as summarized in
the SSA report, in 2015, State wildlife
agencies in Idaho, Montana,
Washington, and Wyoming, along with
Federal, tribal, and nongovernmental
organization partners, developed a
collaborative and coordinated
monitoring program to be implemented
in a coordinated fashion across the
species’ range in the western United
States. In 2015, the State of Wyoming
contracted with the Wolverine Initiative
to conduct the Wyoming Wolverine
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Occupancy Pilot Study to address
questions pertaining to the status and
distribution of wolverines throughout
the Greater Yellowstone Ecosystem and
the Bighorn Mountains in the winter of
2015–2016. Expanding on this study,
the Western States Wolverine Working
Group designed and implemented the
WSWCP–Coordinated Occupancy
Survey in the winter of 2016–2017, and
preliminary results are presented in the
SSA report. The Western Association of
Fish and Wildlife Agencies (WAFWA)
Wildlife Chiefs Wolverine
Subcommittee (formally endorsed in
2014) currently provides a forum for
western States to work collaboratively
with each other and with the Service,
Tribes, and other partners, for
conserving wolverines across the
western United States. To date,
approximately $1.5 million of that
funding has been applied towards
conservation and management actions,
including the WSWCP (McDonald 2017,
pers. comm.). This group is also
developing a connectivity study project
to support conservation planning efforts
for the Rocky Mountains and North
Cascades regions.
In addition, multiple western States
have identified the North American
wolverine as a Species of Greatest
Conservation Need in their State
Wildlife Action Plans, and the North
American wolverine is a focal species of
conservation strategies for conservation
targets in a number of ecoregions (e.g.,
Cascades, Sierra Nevada) that support
forested lowlands, subalpine-high
montane conifer forest where
wolverines occur. These State
designations provide information to
assist resource managers with proactive
decision making regarding species
conservation and data collection
priorities. Finally, the Nez Perce Tribe
is currently preparing an Integrated
Resource Management Plan, a Plant and
Wildlife Conservation Strategy, and a
Forest Management plan with the
wolverine defined as a species of
conservation concern in all three draft
plans (Miles 2017, pers. comm.).
In total, these funded and volunteer
collaborative, landscape-level
conservation efforts ensure continued
support for the conservation of the
North American wolverine. Although
we did not rely on these plans to
support our decision, we recognize that
these plans, when implemented, will
likely benefit wolverines and their
habitat.
(12) Comment: We received
comments from several industry groups
supporting our August 13, 2014,
withdrawal (79 FR 47522) of our
February 4, 2013, proposed rule (78 FR
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64623
7864) to list the North American
wolverine as threatened. In general,
their support rests on the following: (1)
The DPS determination presented in our
previous proposed rules (both 2010 and
2013) was flawed; (2) the North
American wolverine does not meet the
definition of a threatened species; (3)
the obligate relationship with denning
and need for snow has not been
adequately addressed (and may be a
habitat preference); and (4) climate
model projections do not support
complete loss of snow. They also urged
us to reaffirm prior findings that winter
recreation (motorized and
nonmotorized) is not a threat to
wolverines.
Our Response: Given that our updated
analysis and new information included
in the wolverine SSA report directly
relates to our previous DPS
determination in the 2013 proposed
rule, we reevaluated wolverines in the
contiguous United States under our DPS
Policy. See the Distinct Population
Segment section below for more
information. We provide our analysis of
the status of wolverines in the
contiguous United States below in the
Determination of Species Status. The
topic of denning behavior is discussed
in the wolverine SSA report (see Use of
Dens and Denning Behavior discussion
in the Reproduction and Growth section
in the wolverine SSA report (Service
2018, pp. 23–28)). For our analysis of
the effects of climate change to
wolverines and denning habitat, see
Climate Change and Potential for
Cumulative Effects below.
(13) Comment: We received a
comment from an industry group stating
that our decision to prepare the
February 4, 2013, proposed rule (78 FR
7864) to list the North American
wolverine as threatened was due to a
‘‘misreading’’ of the Service’s obligation
under our 2011 Settlement Agreement,
and therefore the proposed rule was not
developed from ‘‘an open-ended
scientific inquiry.’’ We received a
comment from an industry group stating
that the Service should not ‘‘revert back
to the 2013 proposed rule’’ and should
conduct a new analysis of potential
impacts to the species, revise the
proposed listing using newly available
information, and reevaluate our
previous DPS determination.
Our Response: The Service properly
prepared its 2013 proposed rule. On
October 18, 2016, we published a
document in the Federal Register (81
FR 71670) announcing that we would
initiate a new status review of the North
American wolverine, to determine
whether this DPS meets the definition of
an endangered or threatened species
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under the Act, or whether the species is
not warranted for listing. The Service
has prepared the wolverine SSA report
that includes discussion and analyses of
the best available scientific information
regarding life history, biology, and
consideration of current and future
vulnerabilities. This information was
used to evaluate the current and future
conditions of the species, and to inform
our current determination.
Comments From Tribes
(14) Comment: We received
comments from one consortium of
Tribal nations stating that, based on the
weight of evidence provided in our
previous rules, the North American
wolverine meets the definition of
endangered or threatened and is
therefore warranted for listing. Specific
threats mentioned in the comment letter
included current population status,
winter recreation activities, and effects
of climate change. The Tribes also
included comments documenting the
cultural value of the wolverine and
connection to cultural practices and
concern for the loss of wolverine
populations in the contiguous United
States. The Tribes encouraged the
Service to use sound and solid science
in the listing determination, and noted
that additional population monitoring
and Tribal climate change modeling
efforts are under way to evaluate the
status of the wolverine.
Our Response: We appreciate the
unique perspective provided by the
Tribal nations regarding the
contribution of the North American
wolverine to the Tribes’ culture and
spirituality. We also appreciate the
commitment of the Tribal nations to
continue ongoing studies of wolverines.
As described in the wolverine SSA
report, we evaluated new information,
as well as information not previously
considered, and contacted several
wolverine researchers (both within and
outside the United States) to provide a
more detailed description of the
wolverine’s life history and ecology,
including a detailed discussion of
wolverine denning habitat and behavior.
We conducted new analyses to develop
a current potential extent of occupancy
using the most recent verified
observations. Current potential extent is
the perimeter of the outermost
geographic limits based on all
(available) occurrence records (that is,
the maximum extent of occurrences) of
a species minus those areas where we
believe the species has been extirpated
(Service 2017). Conservation measures
and regulatory mechanisms relative to
the wolverine were also provided in the
wolverine SSA report. This information
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was used to evaluate the current
(potential stressors) and future
conditions of the species, and inform
our current determination. We
evaluated results from a fine-scale
analysis of the potential effects of
climate change to future snowpack
conditions in two regions of the Rocky
Mountains. This analysis found that
significant areas (several hundred km2/
mi2) will persist on May 1 at elevations
used by wolverines for denning. We
determined that, based on the best
available information, the North
American wolverine in the contiguous
United States does not warrant listing as
threatened or endangered under the Act.
(15) Comment: We received
comments from one Tribe whose
aboriginal territory is occupied by the
North American wolverine. The Tribe
submitted a comment letter in 2013
supporting our proposed listing. The
Tribe stated that the conservation and
restoration of the wolverine and other
species within this homeland is of great
importance to the Tribe’s subsistence,
culture, religion, and economy. The
letter also identified conservation and
management plans currently under
development and highlighted that the
wolverine is designated as a species of
concern in these current draft plans.
Specific comments were provided
relative to threats from climate change
(including relative to demographic
stochasticity), recreation and urban
development, and incidental take.
Included in those comments were
references to other studies under way
(e.g., Adaptation Partners and climate
change vulnerability assessments;
winter recreation study) to evaluate
these potential stressors.
Our Response: We appreciate the
perspective provided regarding the
importance of the wolverine and other
species to the Tribe and its commitment
to current and future conservation and
management actions. We also appreciate
and evaluated the information presented
in the citations that were provided in
the comment letter. As described in the
wolverine SSA report, we evaluated
several new scientific publications and
information not previously considered
in preparing a new status review. This
information was used to evaluate the
current conditions (i.e., potential
stressors, including winter recreation)
and future conditions (e.g., effects of
climate change) of the species. Based on
the best available information, we
determined that the North American
wolverine in the contiguous United
States does not warrant listing as
threatened or endangered under the Act.
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State Agency Comments
We received extensive comments
from several western States, requesting
that we consider previously submitted
comments in response to our previously
proposed listing rule (78 FR 7864;
February 4, 2013) as well as additional
comments submitted in response to our
2016 notice reopening public comment
(81 FR 71670; October 18, 2016). These
comments were grouped together and
summarized as described below:
(16) Comment: We received detailed
comments critical of our reliance on
‘‘unverified’’ climate model projections
in our 2013 proposed rule, the lack of
discussion of assumptions in adopting
the model findings, the lack of
evaluating alternative hypotheses, and
the need to evaluate these effects at the
den-site scale. One State agency
recommended that, given the
disagreements in the scientific
community on the interpretation of
these results, the Service solicit an
independent, scientific review of the
proposed rule.
Our Response: This withdrawal was
based on the scientific analysis using
the structure of the Service’s Species
Status Assessment (SSA) Framework
(https://www.fws.gov/endangered/
improving_ESA/ssa.html). An SSA is a
focused and rigorous assessment of a
species’ ability to maintain selfsustaining populations over time. This
assessment is based on the best
available scientific and commercial
information regarding life history,
biology, and consideration of current
and future vulnerabilities. The result is
a single document (SSA report) that
delivers foundational science for
informing decisions under the Act,
including listing determinations,
consultations, grant allocations,
permitting, and recovery planning.
In preparing the final SSA report for
the North American wolverine
(available at www.regulations.gov, at
Docket No. FWS–R6–ES–2016–0106),
we reviewed available reports and peerreviewed literature, incorporated survey
information for the purpose of preparing
updated maps of the known species’
current and historical occurrences, and
contacted species experts to collect
additional unpublished information. We
evaluated the appropriate analytical
tools to address data gaps and
uncertainties. In some instances, we
used publications and other reports of
the Eurasian subspecies (Gulo gulo gulo)
to fully inform our knowledge of the
North American wolverine (Gulo gulo
luscus).
Before finalizing the SSA report, the
draft wolverine SSA report was
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submitted for peer review to four
independent peer reviewers in
accordance with our July 1, 1994, peer
review policy (59 FR 34270), the
Service’s August 22, 2016, Director’s
Memo on the Peer Review Process, and
the Office of Management and Budget’s
December 16, 2004, Final Information
Quality Bulletin for Peer Review
(revised June 2012). Results of this
structured peer review process can be
found at https://www.fws.gov/mountainprairie/science/peerReview.php. This
draft was also submitted to our Federal,
State, and Tribal partners for scientific
review. In preparing this determination
to withdraw the proposed rule, we
incorporated the results of these reviews
in the final wolverine SSA report, as
appropriate.
As noted in our previous responses to
public comments (see response to
Comments 1 and 5 above), in our
wolverine SSA report, we recognize that
current climate trends and future (2055
and later) climate model projections
indicate warming temperatures for
much of western North America, and
changes to snow pack conditions. Our
review of the literature found that,
overall, higher elevation areas (e.g.,
Rocky Mountains, Sierra Nevada
Mountains) are more resilient to
projected changes in temperature and
precipitation as compared to lower
elevations (Wobus et al. 2017, p. 12). In
general, models indicate higher
elevations will retain more snow cover
than lower elevations, particularly in
early spring (April 30/May 1). We
present in the wolverine SSA report a
summary of new, fine-scale analysis of
future snow persistence in two regions
of the western United States, Glacier
National Park and Rocky Mountain
National Park. Glacier National Park
represents a high-latitude and relatively
low-elevation area currently occupied
by North American wolverines. Rocky
Mountain National Park is a lower
latitude and high-elevation area within
the North American wolverine’s
historical range, which was occupied by
a male wolverine from 2009 to at least
2012.
As described above in Comment 5,
this new analysis built upon previous
model projections presented in
McKelvey et al. (2011), but with
significant differences such as finer
spatial resolution, incorporation of
slope and aspect, snow depth estimates,
additional years of historical data, and
wider temporal analyses of snow
persistence (April–June). Details of this
modeling exercise are presented in Ray
et al. (2017), and summarized in the
wolverine SSA report. That analysis
indicates large areas (several hundred
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km2/mi2 for each study area) of future
snow (greater than 0.5 m (20 in) in
depth) will persist on May 1 at
elevations currently used by wolverines
for denning. This is true, on average,
across the range of climate models used
out to approximately year 2055.
After reviewing studies not previously
considered and new studies/
publications made available after the
2013 and 2014 proposed rules, we
present in the wolverine SSA report, a
detailed discussion of the North
American wolverine’s physiology and
other life-history characteristics (e.g.,
reproductive behavior). The analysis
speaks to several presumed aspects of
the relationship of denning behavior
and other needs of this species
regarding the presence of persistent
spring snow.
Also, see our response to Comment 5
above for a short summary and our SSA
report for more details regarding our
analysis of the effects of climate change
to denning habitat.
(17) Comment: We received
comments critical of our previous
support for findings by Schwartz et al.
2009 regarding effective population size.
Relatedly, several States commented on
recent dispersal/movements of
wolverines into California, Colorado,
and Utah as evidence of population
expansion.
Our Response: See our response to
Comment 2 above for a discussion of
effective population size. Regarding
recent occurrences of wolverines in the
contiguous United States, wolverines
have recently been found in areas where
they were once extirpated in the
contiguous United States. See the
Population Abundance and Density
section below for more information.
(18) Comment: We received
comments from several western States
presenting clarifications or updates to
incidental trapping events and trapping
regulations.
Our Response: In the wolverine SSA
report, we include a summary of
trapping or hunting of wolverines in the
contiguous United States. At the present
time, trapping or hunting of wolverines
is not allowed in any western State
(with the exception of Alaska, which
was not included in the DPS in our
proposed rule). Legal protections for
wolverines are codified in western State
laws and regulations concerning
hunting and trapping. These protections
include: Endangered in Colorado,
threatened in California and Oregon,
candidate species in Washington, nongame species protections in Idaho and
Wyoming, a species of concern and
furbearer with a closed season in
Montana, and protection from
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collection, importation, and possession
in Utah. Since 2013, there has been a
zero quota for trapping or harvest of
wolverine in Montana.
Incidental trapping of wolverines has
been documented in the contiguous
United States (as recently as December
2017), though not all events have
resulted in mortality (see response to
Comment 3 above). Both Idaho and
Montana are implementing trapper
education programs to minimize
nontarget wolverine captures.
(19) Comment: Several States
provided comments in response to our
2013 proposed rule and to our 2016
reopening of the public comment period
indicating their disagreement with our
determination of a DPS for the
contiguous United States. Specifically,
some commenters stated that the criteria
of significance should be reevaluated,
noting that the proposed rule did not
provide any substantive information to
support our conclusion that the loss of
the wolverine in the contiguous United
States would result in a significant gap
in the range of the species; that is, our
previous use of the loss of latitudinal
range does not provide a rational basis
for concluding that the loss of the
wolverine in the contiguous United
States would be significant in relation to
the taxon. Another commenter stated
that the wolverine population in the
contiguous United States is connected
geographically and genetically to the
Canada/Alaska populations and these
northern populations were likely the
source of recolonization during the 20th
century. Further, this commenter stated
there is not a difference in control of
exploitation and conservation status
between the United States and Canada.
Another commenter noted that,
throughout the 2013 proposed rule, the
Service acknowledged that, historically,
the wolverine population in the
contiguous United States was markedly
reduced by systematic predator control
programs and unregulated trapping. Yet,
as the commenter pointed out, areas of
suitable habitat in the North Cascades,
where trapping has been minimal or
nonexistent for decades, and northern
Rockies, were recolonized by animals
from Canada, where relatively liberal
trapping is still allowed. Thus, our
characterization in the 2013 proposed
rule of ‘‘liberal’’ Canadian regulations as
sufficient to ‘‘maintain the robust
conservation status of the Canadian
population,’’ does not comport with our
characterization that the very limited
trapping in the contiguous United States
(Montana only) is insufficient to
maintain the rebounding population
designated as a DPS.
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Our Response: In light of the updated
analysis and new information included
in the wolverine SSA report, we
reevaluated wolverines in the
contiguous United States under our DPS
Policy. We conclude that the population
of wolverines in the contiguous United
States is not discrete in relation to the
remainder of the species in North
America. As a result, the population of
wolverines in the contiguous United
States is not a listable entity under
section 3(16) of the Act. See the Distinct
Population Segment section below for
more information.
(20) Comment: State agencies
provided citations or copies of
publications and reports relevant to
wolverine ecology that were published
after the 2013 proposed rule.
Our Response: We appreciate the
comprehensive lists of published
literature and survey reports provided
by the State agencies. We evaluated this
information during the preparation of
the wolverine SSA report, and have
incorporated this information, as
appropriate, to ensure that the
wolverine SSA report presents the best
available information regarding the
status of the North American wolverine.
(21) Comment: We received
information providing background
information and preliminary results of
ongoing collaborative conservation
efforts being implemented through the
WSWCP–Coordinated Occupancy
Survey.
Our Response: We appreciate the
additional information provided by the
State agencies participating in the
WSWCP–Coordinated Occupancy
Study. In the wolverine SSA report, we
provide a summary of the preliminary
field and genetic results from the recent
WSWCP–Coordinated Occupancy Study
in four western (contiguous United)
States (see wolverine SSA report for
additional details). We also
incorporated technical comments
received from several State agencies
during the review of the draft wolverine
SSA report into the final report. As
discussed in the wolverine SSA report,
the Western States Wolverine Working
Group is continuing to develop studies
to evaluate wolverine population
distribution and occupancy, and
connectivity across four western States.
(22) Comment: Information was
provided by State agencies describing
the legal protections of wolverines in
individual States and conservation
measures being implemented.
Our Response: In the wolverine SSA
report (Service 2018, Appendix G), we
provide a detailed discussion of current
State (and Federal) regulatory
mechanisms and other conservation
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measures that offer protections for the
North American wolverine. In addition
to the WSWCP–Coordinated Occupancy
Study (Service 2018, Appendix B),
several western States have identified
the North American wolverine as a
Species of Greatest Conservation Need
in their State Wildlife Action Plans, and
the North American wolverine is a focal
species of conservation strategies for
conservation targets in a number of
ecoregions (e.g., Cascades, Sierra
Nevada) that support forested lowlands,
subalpine–high-montane conifer forest
where wolverines occur. These State
designations provide information to
assist resource managers with proactive
decision-making regarding species
conservation and data collection
priorities, and support the conservation
of the North American wolverine and its
habitat.
(23) Comment: We received a
comment from one State agency noting
that the State does not recognize the
North American wolverine as a native
species due to lack of evidence that a
population ever existed within New
Mexico (i.e., unverified species); thus,
the State does not recognize the species
in any of its wildlife statutes or
regulations.
Our Response: We appreciate the
clarification and information provided
by the State agency and have considered
this in our analysis to define the current
potential extent of occurrence for the
North American wolverine in the
contiguous United States (see Figures 1
and 2 below) and in our assessment of
population status in the wolverine SSA
report. In their analysis of wolverine
distribution records in the contiguous
United States, Aubry et al. (2001, p.
2,150) identified 1860 as the most recent
verifiable documentation of wolverine
in northern New Mexico. We received
two unverified accounts of wolverine
sightings in New Mexico from the
general public during the most recent
public comment period. We are
unaware of any recent verifiable
individuals or populations of
wolverines in New Mexico.
(24) Comment: In response to our
request for information in our October
18, 2016, Federal Register document (81
FR 71670), we received comments from
the U.S. Forest Service submitting
verifiable and new records of
wolverines from 2000 to 2016. These
records include observations from
camera surveys by both governmental
and nongovernmental organizations,
photos from private citizens, and
locations from a regional study.
Our Response: We appreciate the
information provided and incorporated
these observations and detections in our
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analysis to define the current potential
extent of occurrence for the North
American wolverine in the contiguous
United States (see Figures 1 and 2
below) and in our assessment of
population status in the wolverine SSA
report.
Background
A comprehensive review of the life
history, population trends, and ecology
of the North American wolverine is
presented in the wolverine SSA report
(Service 2018, pp. 3–44). The Service
recognizes the North American
wolverine as the subspecies Gulo gulo
luscus (Service 2018, p. 8). Wolverines
are a medium-sized (about 1 m (3.3 ft)
in length) carnivore, with a large head,
broad forehead, and short neck (Service
2018, p. 4). Wolverines have heavy
musculature and relatively short legs,
and large feet with strong, curved claws
for digging and climbing (Service 2018,
p. 4). Their feet are adapted for travel
through deep snow and, during the
winter, dense, stiff, bristle-type hairs are
found between the toes and around the
foot pad; this characteristic becomes
diminished in the summer (Service
2018, p. 4). The wolverine is the largest
terrestrial member of the Mustelidae
family (weasels, fisher, mink, marten,
and others) and resembles a small bear
with a bushy tail (Service 2018, p. 1).
Wolverines possess a number of
morphological and physiological
adaptations that allow them to travel
long distances and they maintain large
territories in remote areas (Service 2018,
p. 1). They have been described as
curious, intelligent, and playful, but
cautious animals, though their social
behavior and social organization has not
been well-studied (Service 2018, p. 1).
Wolverines have a distribution that
includes the northern portions of
Europe, Asia, and North America. In
North America, they are found in
Alaska, much of Canada, and the
western-northwestern United States.
During the late 1800s and early 1900s,
the wolverine population declined or
was extirpated in much of the
contiguous United States (lower 48
States), which has been largely
attributed to unregulated trapping (Hash
1987, p. 583). Wolverine numbers have
recovered to some extent from this
decline and, in the United States,
wolverines are currently found in parts
of Washington, Oregon, Idaho, Montana,
Wyoming, California (single male), and
Alaska, and as recently as 2010 in
Michigan, 2012 in Colorado, and 2016
in Utah. Known reproducing wolverine
populations are found in Washington,
Idaho, Montana, and Wyoming (Service
2018, p. 1).
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Regulatory and Analytical Framework
Regulatory Framework
Section 4 of the Act (16 U.S.C. 1533)
and its implementing regulations (50
CFR part 424) set forth the procedures
for determining whether a species is an
‘‘endangered species’’ or a ‘‘threatened
species.’’ The Act defines an
endangered species as a species that is
‘‘in danger of extinction throughout all
or a significant portion of its range,’’ and
a threatened species as a species that is
‘‘likely to become an endangered
species within the foreseeable future
throughout all or a significant portion of
its range.’’ The Act requires that we
determine whether any species is an
‘‘endangered species’’ or a ‘‘threatened
species’’ because of any of the following
factors:
(A) The present or threatened
destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial,
recreational, scientific, or educational
purposes;
(C) Disease or predation;
(D) The inadequacy of existing
regulatory mechanisms; or
(E) Other natural or manmade factors
affecting its continued existence.
These factors represent broad
categories of natural or human-caused
actions or conditions that could have an
effect on a species’ continued existence.
In evaluating these actions and
conditions, we look for those that may
have a negative effect on individuals of
the species, as well as other actions or
conditions that may ameliorate any
negative effects or may have positive
effects.
We use the term ‘‘threat’’ to refer in
general to actions or conditions that are
known to or are reasonably likely to
negatively affect individuals of a
species. The term ‘‘threat’’ includes
actions or conditions that have a direct
impact on individuals (direct impacts),
as well as those that affect individuals
through alteration of their habitat or
required resources (stressors). The term
‘‘threat’’ may encompass—either
together or separately—the source of the
action or condition or the action or
condition itself.
However, the mere identification of
any threat(s) does not necessarily mean
that the species meets the statutory
definition of an ‘‘endangered species’’ or
a ‘‘threatened species.’’ In determining
whether a species meets either
definition, we must evaluate all
identified threats by considering the
expected response by the species, and
the effects of the threats—in light of
those actions and conditions that will
ameliorate the threats—on an
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individual, population, and species
level. We evaluate each threat and its
expected effects on the species, then
analyze the cumulative effect of all of
the threats on the species as a whole.
We also consider the cumulative effect
of the threats in light of those actions
and conditions that will have positive
effects on the species, such as any
existing regulatory mechanisms or
conservation efforts. The Secretary
determines whether the species meets
the definition of an ‘‘endangered
species’’ or a ‘‘threatened species’’ only
after conducting this cumulative
analysis and describing the expected
effect on the species now and in the
foreseeable future.
The Act does not define the term
‘‘foreseeable future,’’ which appears in
the statutory definition of ‘‘threatened
species.’’ Our implementing regulations
at 50 CFR 424.11(d) set forth a
framework for evaluating the foreseeable
future on a case-by-case basis. The term
‘‘foreseeable future’’ extends only so far
into the future as the Services can
reasonably determine that both the
future threats and the species’ responses
to those threats are likely. In other
words, the foreseeable future is the
period of time in which we can make
reliable predictions. ‘‘Reliable’’ does not
mean ‘‘certain’’; it means sufficient to
provide a reasonable degree of
confidence in the prediction. Thus, a
prediction is reliable if it is reasonable
to depend on it when making decisions.
It is not always possible or necessary
to define foreseeable future as a
particular number of years. Analysis of
the foreseeable future uses the best
scientific and commercial data available
and should consider the timeframes
applicable to the relevant threats and to
the species’ likely responses to those
threats in view of its life-history
characteristics. Data that are typically
relevant to assessing the species’
biological response include speciesspecific factors such as lifespan,
reproductive rates or productivity,
certain behaviors, and other
demographic factors.
Analytical Framework
The SSA report documents the results
of our comprehensive biological status
review for the species, including an
assessment of the potential threats to the
species. The SSA report does not
represent a decision by the Service on
whether the species should be proposed
for listing as an endangered or
threatened species under the Act. It
does, however, provide the scientific
basis that informs our regulatory
decisions, which involve the further
application of standards within the Act
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and its implementing regulations and
policies. The following sections provide
summaries of the key results and
conclusions from the SSA report; the
full SSA report can be found on the
Mountain-Prairie Region website at
https://www.fws.gov/mountain-prairie/
and at https://www.regulations.gov under
Docket No. FWS–R6–ES–2016–0106.
To assess wolverine viability, we used
the three conservation biology
principles of resiliency, redundancy,
and representation (Shaffer and Stein
2000, pp. 306–310). Briefly, resiliency
supports the ability of the species to
withstand environmental and
demographic stochasticity (for example,
wet or dry, warm or cold years),
redundancy supports the ability of the
species to withstand catastrophic events
(for example, droughts, large pollution
events), and representation supports the
ability of the species to adapt over time
to long-term changes in the environment
(for example, climate changes). In
general, the more resilient and
redundant a species is and the more
representation it has, the more likely it
is to sustain populations over time, even
under changing environmental
conditions. Using these principles, we
identified the species’ ecological
requirements for survival and
reproduction at the individual,
population, and species levels, and
described the beneficial and risk factors
influencing the species’ viability.
The SSA process can be categorized
into three sequential stages. During the
first stage, we evaluated the individual
species’ life-history needs. The next
stage involved an assessment of the
historical and current condition of the
species’ demographics and habitat
characteristics, including an
explanation of how the species arrived
at its current condition. The final stage
of the SSA involved making predictions
about the species’ responses to positive
and negative environmental and
anthropogenic influences. This process
used the best available information to
characterize viability as the ability of a
species to sustain populations in the
wild over time. We use this information
to inform our regulatory decision.
Distinct Population Segment
Pursuant to the Act, we must consider
for listing any species, subspecies, or,
for vertebrates, any distinct population
segment (DPS) of these taxa, if there is
sufficient information to indicate that
such action may be warranted. To
interpret and implement the DPS
provision of the Act and Congressional
guidance, the Service and the National
Marine Fisheries Service published, on
February 7, 1996, an interagency Policy
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Regarding the Recognition of Distinct
Vertebrate Population Segments under
the Act (61 FR 4722; February 7, 1996).
This policy addresses the recognition of
DPSs for potential listing actions. The
policy allows for more refined
application of the Act that better reflects
the biological needs of the taxon being
considered, and avoids the inclusion of
entities that do not require its protective
measures.
Under our DPS policy, three elements
are considered in a decision regarding
the status of a possible DPS as
endangered or threatened under the Act.
These are applied similarly for
additions to the list of endangered and
threatened species, reclassification, and
removal from the list. They are: (1)
Discreteness of the population segment
in relation to the remainder of the taxon;
(2) the biological or ecological
significance of the population segment
to the taxon to which it belongs; and (3)
the population segment’s conservation
status in relation to the Act’s standards
for listing (i.e., whether the population
segment is, when treated as if it were a
species or subspecies, an endangered or
threatened species). Discreteness refers
to the degree of isolation of a population
from other members of the species, and
we evaluate this factor based on specific
criteria. If a population segment is
considered discrete, we must consider
whether the discrete segment is
‘‘significant’’ to the taxon to which it
belongs by using the best available
scientific and commercial information.
When determining if a potential DPS is
significant, our policy directs us to
sparingly list DPSs while encouraging
the conservation of genetic diversity. If
we determine that a population segment
is both discrete and significant, we then
evaluate it for endangered or threatened
species status based on the Act’s
standards.
Both new and updated information
and analyses presented in the wolverine
SSA report, summarized below in
support of our listing determination,
along with public comment, have
prompted us to reevaluate our previous
assessment of the DPS (presented in our
2013 proposed listing rule, which in
turn relied on the DPS analysis
completed in our 2010 12-month
finding) with respect to wolverine in the
contiguous United States. Below we
provide our revised evaluation of
discreteness under the DPS policy of the
segment of the North American
wolverine occurring in the contiguous
United States.
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Distinct Population Segment Analysis
for Wolverine in the Contiguous United
States
Analysis of Discreteness
Under our DPS Policy, a population
segment of a vertebrate species may be
considered discrete if it satisfies either
one of the following conditions: (1) It is
markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors
(quantitative measures of genetic or
morphological discontinuity may
provide evidence of this separation); or
(2) it is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the Act (inadequacy of
existing regulatory mechanisms).
Discreteness Based on Marked
Separation
In our February 4, 2013, proposed
listing rule (78 FR 7864), we did not rely
on marked separation from other
populations to support discreteness of
the contiguous United States wolverine
population. As supported by
information in the SSA report, we
maintain that there are no physical,
physiological, ecological, or behavioral
factors separating wolverines in the
contiguous United States from
wolverines in Canada. We do not
consider wolverines in the contiguous
United States to be genetically isolated
from wolverines in Canada (McKelvey
et al. 2014; Pilgrim and Schwartz 2018).
Therefore, wolverines in the contiguous
United States are not discrete based on
marked separation from other
populations of the same taxon.
Discreteness Based on the International
Border—Legal Status Conveyed by
National, State, and Provincial
Governments; Differences in Control of
Exploitation
Our 2013 proposed rule (78 FR 7864),
which incorporated by reference our
DPS analysis from our 2010 12-month
finding, found there was no significant
difference between the legal status of
wolverines between Canada and the
United States (75 FR 78030; December
14, 2010). In the wolverine SSA report,
we provide an updated assessment of
legal protections and regulatory
mechanisms for wolverine in North
America (Service 2018, pp. 70–71,
Appendix G). Legal protections in the
contiguous United States include State
listing in California and Oregon
(threatened), endangered in Colorado, a
candidate species in Washington, non-
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game species protections in Idaho and
Wyoming, a species of concern and
furbearer with a closed season in
Montana, and protected from collection,
importation, and possession in Utah. In
Canada, provincial designations range
from endangered to threatened in
eastern provinces, and sensitive/special
concern to no ranking in other
provinces (definitions provided by the
Committee on the Status of Endangered
Wildlife in Canada, 2014). As was
determined in our 2013 proposed listing
rule (78 FR 7864), we again find no
significant differences in legal status.
In the 2010 12-month finding (75 FR
78030) and reiterated in our 2013
proposed listing rule (78 FR 7864), we
stated that differences in control of
exploitation exist, but favor the
contiguous United States population.
Trapping or hunting of wolverines is
currently prohibited in the contiguous
United States and regulated as
appropriate in Canada (Service 2018,
pp. 68–69). In the wolverine SSA report,
we included a new analysis of trapping
in southern Canada and trapping effort
along the U.S.–Canada border, which
we found to be limited. Thus, we
conclude that the differences in
exploitation are not significant in light
of section 4(a)(1)(D) of the Act
(inadequacy of existing regulatory
mechanisms).
Discreteness Based on the International
Border—Differences in Management of
Habitat
As we outlined in the proposed 4(d)
rule (78 FR 7888) management activities
(e.g., timber harvest, wildland
firefighting, prescribed fire, and
silviculture) can modify wolverine
habitat, but this generalist species
appears to be little affected by changes
to the vegetative characteristics of its
habitat. In addition, most wolverine
habitat occurs at high elevations in
rugged terrain that is not conducive to
intensive forms of silviculture and
timber harvest. Habitat management is
not a conservation need for wolverine.
Therefore, differences in management of
habitat between the United States and
Canada are not significant in light of
section 4(a)(1)(D) of the Act.
Discreteness Based on the International
Border—Differences in Conservation
Status
In the December 14, 2010, 12-month
finding (75 FR 78030), which is
incorporated and discussed in the
February 4, 2013, proposed listing rule
(78 FR 7864), we found that the
wolverine population in the contiguous
United States met the second DPS
discreteness condition because of
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differences in conservation status as
delimited by the U.S.–Canada
international governmental boundary.
We found that those differences were
substantial and significant in light of
section 4(a)(1)(D) of the Act. We stated
that in the remaining current range in
Canada and Alaska, wolverines exist in
well-distributed, interconnected, large
populations. We added that, conversely,
wolverine populations in the remaining
United States range appear to be at
numbers so low that their continued
existence could be at risk, especially in
light of the threats to the species. In the
2010 finding, we stated that risks come
from three main factors: (1) Small total
population size; (2) effective population
size below that needed to maintain
genetic diversity and demographic
stability; and (3) the fragmented nature
of wolverine habitat in the contiguous
United States that results in smaller,
isolated sky island patches separated by
unsuitable habitat. We stated it was
apparent that maintaining wolverines
within their native range in the
contiguous United States into the future
is likely to require regulatory
mechanisms that are not currently in
place. As a result, we concluded that the
contiguous United States population of
the wolverine meets the discreteness
criterion in our DPS Policy (61 FR 4722,
February 7, 1996). Consequently, we
used the international border between
the United States and Canada to define
the northern boundary of the contiguous
United States wolverine DPS in our
December 14, 2010, 12-month finding
(75 FR 78030) and our February 4, 2013,
proposed listing rule (78 FR 7864).
Below we provide a reevaluation of that
determination supported by information
presented in the wolverine SSA report.
Small Total Population Size—
Wolverine densities vary across North
America and have been described as
naturally low (van Zyll de Jong 1975, p.
434); wolverine populations are
naturally uncommon given the species’
large home range, wide-ranging
movements, and solitary characteristics
(Service 2018, p. 56). There are many
fewer wolverines in the contiguous
United States than there are in Canada
and Alaska (Committee on the Status of
Endangered Wildlife in Canada
(COSEWIC) 2014, p. 36; Inman et al.
2013, p. 282; Service 2018, p. 71), but
this is more a reflection of the amount
of suitable habitat available within the
contiguous United States (both currently
and historically) for a species that needs
large exclusive territories, than it is a
reflection of poor conservation status.
Wolverines in Canada are considered to
occur as a single large group as they are
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easily able to move between areas of
suitable habitat and because wolverine
habitat is relatively contiguous
(Harrower 2017, pers. comm.). However,
wolverines in the contiguous United
States are considered to be a
metapopulation connected with
wolverine populations in Canada
(Inman et al. 2013, p. 277). Wolverines
currently occupy areas in the
contiguous United States where they
were once extirpated and continue to
repopulate the contiguous United States
after decades of unregulated trapping,
hunting, and poisoning (Service 2018, p.
iv). The same holds true for Canada,
where wolverines are being detected in
areas once extirpated (COSEWIC 2014,
p. v).
These movement patterns are
supported by recent genetic information
that indicates wolverines from Canada
have slowly repopulated the contiguous
United States over the past century
since the era of unregulated persecution
(Service 2018, pp. 45–50). This point is
discussed in detail below in the Genetic
Diversity section of this withdrawal. We
stated in the December 14, 2010, finding
that differences in population sizes
between the contiguous United States
and Canada were reflective of a
difference in conservation status (75 FR
78030). However, based on new
information, we now conclude that the
contiguous United States wolverines
represent a peripheral population at the
southern extent of the North American
wolverine range. Thus, we now consider
the small population size of wolverines
in the United States to be a natural
result of habitat fragmentation and not
reflective of a difference in conservation
status (see Habitat Fragmentation below
for more detail). Therefore, any
difference in population size on the
contiguous United States side of the
international border is not a significant
difference in conservation status in light
of section 4(a)(1)(D) of the Act as it
applies to discreteness.
Effective Population Size—Effective
population size (Ne) is defined as ‘‘the
size of an idealized population that
would experience the same amount of
genetic drift and inbreeding as the
population of interest (Service 2018,
Box 2). In popular terms, Ne is the
number of individuals in a population
that contribute offspring to the next
generation’’ (Hoffman et al. 2017, p.
507). Effective population size can be
interpreted differently depending on
how it’s defined and used, and the
concept of effective population size (Ne)
(see review by Wang et al. 2016) and,
relatedly, minimum viable population,
has been a topic of debate, particularly
the 50/500 rule (population size of 50
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for short-term, and 500 for long-term
genetic health). Importantly, the concept
and guidelines for genetically effective
population size were developed for a
single, isolated population (Laikre et al.
2016, p. 280). The term ‘‘effective
population size’’ is not a meaningful
term unless additional context is
provided relative to which concept of
population size is being evaluated
(Ewens 1990, p. 309). Demographic
factors are needed when interpreting
actual population size from an effective
population size; thus, there is no
justification for a fixed, genetically
derived minimum viable population
size value of ‘500’ as each case is unique
and is dependent on such factors as sex
ratio, subpopulations, dispersal, and
immigration (Ewens 1990, pp. 311–313).
As noted above, we do not consider
the wolverine population in the
contiguous United States to be
genetically isolated from wolverines on
the other side of the international border
in Canada. In the wolverine SSA report,
we provide a contextual discussion of
the effective population size concept,
particularly in the context of genetic
studies related to the phylogeographic
history of the North American
wolverine (Service 2018, pp. 45–50),
which was not well described in the
2013 proposed rule. In sum, the spatial
distribution of genetic variability
currently observed in wolverines in
North America appears to be a reflection
of a complex history in which
population abundance has fluctuated
since the time of the last glaciation with
insufficient time passing since human
persecution, since at least the 1700s, to
allow for full recovery of wolverine
densities (Cardinal 2004, pp. 23–24;
Zigouris et al. 2012, p. 1,554). This
history and the fact that wolverines in
the contiguous United States occupy the
southern periphery of its entire North
American range are important
considerations. The wolverine SSA
report also presents information from
genetic and observational studies that
provide support for wolverine
movement across the international
border of the contiguous United States
and Canada (Aubry et al. 2016, pp. 16,
20; Lucid et al. 2016, p. 184; Service
2018, pp. 9–23). Thus, we consider
wolverines that occupy the contiguous
United States to be genetically
continuous with wolverines in adjacent
Canadian provinces.
Wolverines travel (disperse) through
areas outside high-elevation, forested
habitats. For example, tracked
movements of a male wolverine, M56,
from Wyoming into Colorado and its
subsequent discovery in North Dakota,
indicate extensive travel outside of
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modeled primary wolverine habitat (i.e.,
Inman et al. 2013), including through
arid grasslands and shrubland habitats
of the Wyoming Basin ecoregion
(Packila et al. 2017, entire). This
animal’s movement also supports some
level of connectivity (and potential gene
flow) between currently occupied
habitat (Wyoming) and unoccupied
habitat within the wolverine’s historical
range (Colorado) (Packila et al. 2017, p.
404). Similarly wolverines in the North
Cascades region have moved from
Washington and Idaho into British
Columbia, and from Montana to British
Columbia and Alberta (Service 2018, p.
45). Based on genetic analyses, the male
wolverine currently occupying an area
within the Sierra Nevada Mountains of
California also represents evidence of
connectivity between wolverine
populations of the Rocky and Sierra
Nevada Mountain Ranges (Moriarty et
al. 2009, p. 154). Within the
Southwestern Crown of the Continent
(SWCC) in northwestern Montana,
cross-valley movements of wolverines
have been detected, which researchers
believe is an indication of good
connectivity in this region (SWCC
Working Group 2016, pers. comm.).
A preliminary mitochondrial DNA
analysis was prepared for wolverine
samples collected during the winters of
2015–2016 and 2016–2017 as part of the
Western States Wolverine Conservation
Project—Coordinated Occupancy
Survey (Pilgrim and Schwartz 2018,
entire). All 45 wolverines identified
from samples collected in Idaho,
Montana, and Wyoming match
haplotype Wilson-A, which is common
throughout the Rocky Mountains,
Alaska, and Canada, while all 5
wolverines identified from samples
collected in Washington match
haplotype Wilson-C (Pilgrim and
Schwartz 2018, p. 3). Previous analyses
of recent or modern (1989–2012)
samples from the Cascades Range in
northern Washington and southern
British Columbia, as presented in
McKelvey et al. (2014, p. 328), were
characterized as haplotype C, and one
historical (defined in this study as pre1930) sample as haplotype A (McKelvey
et al. 2014, p. 327). Outside of this
region, haplotype C has been found only
in Alberta, Saskatchewan, and Nunavut
provinces (McKelvey et al. 2014, p.
330). Based on mitochondrial DNA,
McKelvey et al. (2014, p. 330)
concluded that modern (defined in their
study as 1989–2012) wolverine
populations in the contiguous United
States are the result of recolonization
(following persecution during a period
of unregulated hunting or trapping and
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poisoning) from the north. The
additional mitochondrial analysis from
samples collected in 2015, 2016, and
2017 provides further support that all
contiguous United States historical (pre1900) and recent wolverine populations
are likely descendants of immigrants
from Canada.
The 2013 proposed rule presented an
effective population size estimate for
wolverines in the contiguous United
States from a publication by Schwartz et
al. (2009), which estimated a summed
effective population size of 35, with
credible limits from 28 to 52 (Schwartz
et al. 2009, p. 3,226). As described in
the wolverine SSA report, the study
used wolverine samples from the main
part of the Rocky Mountain wolverine
populations and did not include
subpopulations from two other
mountain regions in Montana, and
samples were missing from other parts
of the wolverine range in Idaho and
other areas of Montana. Thus, the
analysis missed wolverine
subpopulations and individuals, which
would underestimate the results for this
type of analysis. Furthermore, a small
effective population size would be more
of a concern if the population was in
isolation; however, wolverines in the
contiguous United States are not
genetically or physically isolated from
wolverines in Canada.
To summarize, the currently known
spatial distribution of genetic variability
in wolverines in North America appears
to be a reflection of a complex history
where population abundance has
fluctuated since the time of the last
glaciation and insufficient time has
passed since human persecution for a
full recovery of wolverine densities
(Cardinal 2004, pp. 23–24; Zigouris et
al. 2012, p. 1,554). Zigouris et al. (2012,
p. 1,545) noted that the genetic diversity
reported in Cegelski et al. (2006) and
Kyle and Strobeck (2001, 2002) for the
southwestern edge of the North
American range represented only part of
the diversity in the northern
populations of wolverines. Zigouris et
al. (2012, p. 1,545) posit that the
irregular distribution of wolverines in
the southwestern periphery and the
genetic diversity observed in those
analyses is a result of population
bottlenecks that were caused by range
contractions from a panmictic (random
mating) northern core population
approximately 150 years ago coinciding
with human persecution. Recent
dispersals of wolverines into Colorado
(2009), California (2008), and Utah
(2014) provide evidence for connectivity
and the potential for gene flow between
Northern Rocky Mountain populations
and areas where wolverines were
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extirpated. As noted above, there is also
recent evidence of wolverine movement
across the international border.
Furthermore, our analysis of trapping
levels in the wolverine SSA report does
not support previous assumptions that
trapping in Canada near the border acts
as a barrier to wolverine movement into
the contiguous United States (Service
2018, pp. 68–69). Finally, very few
successful migrants are needed per
generation to maintain at least 95
percent of the genetic variation in the
next 100 generations (approximately
750 years) in the contiguous United
States (Cegelski et al. 2006, p. 209).
We conclude that this level of
migration from the north has already
been occurring following the end of
intense persecution of this species;
wolverines are currently observed in
previously occupied areas within the
contiguous United States. Given the
recent observations of wolverines
moving vast distances over varied
terrain and across the U.S.–Canada
border, our recent assessment of the low
levels of trapping mortality in Canada
near the border, and further
confirmation of Canada as the source of
wolverine genetics present in
contiguous United States wolverines,
we believe that wolverines in the
contiguous United States are not
separated genetically from the larger
population in Canada. Wolverines in the
contiguous United States exhibit genetic
and phenotypic similarities with
wolverines in Canada that implies
connectivity with Canada. As such, we
conclude that it is not biologically
appropriate to consider the low effective
population size of wolverines on the
contiguous United States side of the
border as a difference in conservation
status that is significant in light of
section 4(a)(1)(D) of the Act as it applies
to discreteness. For additional
information related to wolverine genetic
diversity and effective population size,
see Genetic Diversity below and the
wolverine SSA report (Service 2018, pp.
45–50).
Habitat Fragmentation—In our 2010
12-month finding (incorporated into the
2013 proposed listing rule), we stated
that wolverine habitat in the contiguous
United States consists of small, isolated
islands of high-elevation habitat
separated from each other by low
valleys of unsuitable habitat. We also
described that these ‘habitat islands’ are
represented by areas containing spring
snow, citing Copeland et al. (2010). We
concluded that the fragmented nature
and distribution of wolverine habitat in
the contiguous United States results in
a population that is highly vulnerable to
extirpation because of lack of
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connectivity between subpopulations,
and this also makes them more
vulnerable to external threats (75 FR
78030; December 14, 2010).
Our previous analysis of wolverine
habitat fragmentation relied upon the
assumption that wolverines are
constricted to habitats that contain
deep, persistent spring snow cover and,
therefore, are more or less confined to
areas that were defined by the Copeland
et al. (2010) spring snow cover model.
However, wolverines are observed in
and move through areas without snow
cover (e.g., male wolverines dispersing
to California and Colorado), and female
wolverines have successfully denned in
areas outside previously modeled
projections of deep, persistent spring
snow cover (e.g., Webb et al. 2016;
Persson 2017, pers. comm.; Jokinen
2018, pers. comm.).
We now conclude that it is not
accurate to categorize the occupied
habitat of wolverines in the contiguous
United States as ‘habitat islands.’ As
discussed above, wolverine populations
in the contiguous United States
represent the southern periphery of a
much larger range of the North
American wolverine due to naturally
occurring landscape features such as
high elevation and topographic
roughness of mountain regions. Thus,
the distribution of persistent spring
snow cover in mountainous regions
does not represent the only determining
habitat feature for wolverines. The
availability of prey and avoidance of
predators are also important elements of
wolverine habitat (Inman et al. 2012, p.
785; Scrafford et al. 2017, p. 34)). As
described in the SSA report, wolverines
use a unique and productive ecological
niche that allows them to occupy highelevation regions across the
northwestern portion of the contiguous
United States (Service 2018, pp. 27, 38).
Finally, as noted above, wolverine
movement in the contiguous United
States is not constrained by highelevation habitat or snow cover, and
wolverines can easily move and
disperse long distances in the western
United States (e.g., SWCC Working
Group 2016, pers. comm.; Packila et al.
2017, entire). Therefore, habitat
fragmentation in the context of
availability of persistent spring snow
cover or loss of connectivity in the
contiguous United States is not an
appropriate difference in conservation
status in light of section 4(a)(1)(D) of the
Act as it applies to discreteness.
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Discreteness Based on the International
Border—Differences in Regulatory
Mechanisms
Because there aren’t significant
differences in control of exploitation,
legal conservation status, and
management of habitat, nor other threats
to the wolverine requiring regulatory
mechanisms to address them, we
conclude that there are not differences
in regulatory mechanisms between the
United States and Canada that are
significant in light of section 4(a)(1)(D).
Conclusion on Discreteness
Based on our updated analysis
described above and supported by
information in the wolverine SSA
report, the contiguous United States
population of wolverine does not meet
the discreteness criterion in our DPS
Policy (61 FR 4722; February 7, 1996).
As a result, the contiguous United States
population of wolverines does not
qualify as a DPS and is not a listable
entity under section 3(16) of the Act.
After determining that a vertebrate
population is not discrete, we are not
required to complete an analysis to
determine if the population in question
is significant according to our DPS
Policy.
DPS Conclusion
Based on the best available
information, we conclude that the
population of wolverines in the
contiguous United States is not discrete
in relation to the remainder of the
species in North America. As a result,
the population of wolverines in the
contiguous United States is not a
listable entity under section 3(16) of the
Act.
The DPS Policy sets forth a three-step
process for determining whether a
vertebrate population as a separate
entity warrants listing: (1) Determine
whether the population is discrete; (2) if
the population is discrete, determine
whether the population is significant to
the taxon as a whole; and (3) if the
population is both discrete and
significant, then evaluate the
conservation status of the population to
determine whether it is endangered or
threatened. Although we have
determined that wolverines in the
contiguous United States do not qualify
as a DPS and, therefore, are not a
listable entity, we provide below a
status determination of the wolverine
population in the contiguous United
States. The DPS Policy neither requires
nor prohibits completion of a status
determination once we have determined
that a population does not qualify as a
DPS. Nevertheless, in this instance, we
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concluded that completing an
assessment—and detailing the nature,
scope, and likely effect of the threats to
the population and the species—would
provide us and the public with useful
information regarding wolverines
occupying the contiguous United States.
Summary of Biological Status and
Threats
In preparing the SSA report for the
wolverine, we reviewed available
reports and peer-reviewed literature,
incorporated survey information, and
contacted species experts to collect
additional unpublished information for
the North American subspecies (Gulo
gulo luscus), including Canada and
Alaska. We identified uncertainties and
data gaps in our assessment of the
current and future status of the species.
We also evaluated the appropriate
analytical tools to address these gaps
and conducted discussions with species
experts and prepared updated maps of
the known species’ range in North
America. In some instances, we used
publications and other reports
(primarily from Fenno-Scandinavia) of
the Eurasian subspecies (G. g. gulo) in
completing this assessment.
Since the publication of the February
4, 2013, proposed listing rule (78 FR
7864), several new wolverine studies
have been published (e.g., Aronsson
2017, Aronsson and Persson 2016,
Aronsson et al. 2017, Heinemeyer et al.
2019, Jokinen et al, 2019, Magoun et al.
2017, Persson et al. 2017, Stewart et al.
2016, Webb et al. 2016, see additional
list of citations in the wolverine SSA
report), which have added to our
understanding of wolverine biology
while also highlighting new insights
into identifying key species’ needs and
their interactions with both abiotic and
biotic factors. This new information is
particularly relevant for a difficult-tostudy animal like the wolverine.
Using the species, individual, and
population needs identified for the
wolverine and location results from
surveys and studies, we conducted a
geospatial analysis to estimate the
current potential extent of occurrence
for the North American wolverine in
North America including the contiguous
United States (Figure 1; Service 2018).
‘‘Current potential extent’’ represents
the perimeter of the outermost
geographic limits based on all
(available) occurrence records (that is,
the maximum extent of occurrences) of
a species minus those areas where we
believe the species has been extirpated
(Service 2017). We then evaluated this
area and previous estimates of
potentially suitable habitat in the
western-northwestern United States to
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assess the species’ current conditions
within that region. Our future-condition
analysis includes the potential
conditions that the species or its habitat
may face, that is, the most probable
scenario if those conditions are realized
in the future. This most probable
scenario includes consideration of the
sources that have the potential to most
likely impact the species at the
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known to be currently occupied). The
two regions studied include a highlatitude area near tree line within
Glacier National Park, where tree line
occurs at ∼ 1,800 to 2,100 m (5,906 to
6,890 ft) that is currently occupied by
wolverines; and a lower latitude area
within Rocky Mountain National Park,
where tree line occurs at higher
elevation (∼ 3,500 m (11,483 ft)) (Ray et
al. 2017, p. 2). These sites were selected
to bracket the range of latitude and
elevation wolverines currently occupy
in the contiguous United States (Ray et
al. 2017, p. 2).
For the purpose of this assessment,
we generally define viability as
‘‘consisting of self-sustaining
Our analysis of potential future effects
to the North American wolverine and its
habitat that are associated with climate
change (probabilistic estimates for
temperature and precipitation) is
presented in the SSA report and
summarized here. This analysis was
based on downscaled (high resolution
local climate information derived from
global climate models) climate model
projections, including a detailed study
of two regions in the western United
States—Glacier National Park (currently
occupied by reproducing wolverines)
and Rocky Mountain National Park
(occupied by a single male wolverine
from 2009 to at least 2012, but not
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population or rangewide scales in the
future, including potential cumulative
impacts.
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populations that are well distributed
throughout the species’ range,’’ and
where ‘‘[s]elf-sustaining populations are
those that are sufficiently abundant and
have sufficient genetic diversity to
display the array of life history
strategies and forms that will provide
for their persistence and adaptability in
the planning area over time’’
(Committee of Scientists 1999, p. 38).
We use a timeframe of approximately 38
to 50 years for assessing future effects to
wolverine viability. This timeframe
captures consideration of the projected
future conditions related to trapping/
harvesting, climate change, or other
potential cumulative impacts (Service
2018, p. 73). Beyond this range, climate
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modeling uncertainty increases
substantially. We believe this is a
reasonable timeframe to consider as it
includes the potential for observing
these effects over several generations of
the wolverine.
As discussed above in Analytical
Framework, we consider what the
species needs to maintain viability by
characterizing the status of the species
in terms of resiliency, redundancy, and
representation (Wolf et al. 2015, entire).
Resiliency is having sufficiently large
populations for the species to withstand
stochastic events (arising from random
factors). We can measure resiliency
based on metrics of population health;
for example, birth versus death rates
and population size. Resilient
populations are better able to withstand
disturbances such as random
fluctuations in birth rates (demographic
stochasticity), variations in rainfall
(environmental stochasticity), or the
effects of anthropogenic activities.
Redundancy is having a sufficient
number of populations for the species to
withstand catastrophic events (such as a
rare destructive natural event or episode
involving many populations).
Redundancy is about spreading the risk
and can be measured through the
duplication and distribution of
populations across the range of the
species. The greater the number of
populations a species has distributed
over a larger landscape, the better it can
withstand catastrophic events.
Representation is having the breadth
of genetic makeup of the species to
adapt to changing environmental
conditions. Representation can be
measured through the genetic diversity
within and among populations and the
ecological diversity (also called
environmental variation or diversity) of
populations across the species’ range.
The more representation, or diversity, a
species has, the more it is capable of
adapting to changes (natural or human
caused) in its environment. In the
absence of species-specific genetic and
ecological diversity information, we
evaluate representation based on the
extent and variability of habitat
characteristics within the geographical
range.
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Life-History Needs
Wolverines are capable of moving and
dispersing over great distances over
short periods of time. Wolverine
populations are characterized by
naturally low densities in North
America. The species is highly
territorial, with very little overlap
between same-sex adults. Wolverines
occupy a variety of habitats, but
generally select habitat in locations
away from human settlements.
Wolverines consume a variety of food
resources, and seasonal switching of
prey is commonly observed. As with
other Arctic mammals, wolverines have
the ability to adapt to both warm and
cold ambient temperatures and solar
radiation through both physiological
and behavioral responses, such as
vasodilation, increase in skin
temperature, seasonal adjustments in fur
insulation, and micro- and macrohabitat selection.
Wolverine reproduction includes the
following characteristics: polygamous
behavior (i.e., male mates with more
than one female each year), delayed
implantation (up to 6 months), a short
gestation period (30–40 days), denning
behavior, and an extended period of
maternal care. The reproductive
behavior in wolverines is temporally
adapted to take advantage of the
availability of food resources, limited
interspecific competition, and snow
cover in the winter.
Since the publication of the Service’s
2013 proposed rule to list the distinct
population segment of the North
American wolverine in the contiguous
United States (78 FR 7864; February 4,
2013), several new wolverine studies
have been published (e.g., Aronsson
2017, Aronsson and Persson 2016,
Aronsson et al. 2017, Heinemeyer et al.
2019, Jokinen et al, 2019, Magoun et al.
2017, Persson et al. 2017, Stewart et al.
2016, Webb et al. 2016, see additional
list of citations in the wolverine SSA
report). These studies have improved
our understanding of wolverine biology
while also highlighting new insights
into identifying key species’ needs and
their interactions with both abiotic and
biotic factors. Of particular importance
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relative to life history needs and
wolverine reproductive behavior,
wolverine populations and wolverine
dens have been observed outside
previously modeled projections of
spring snow cover.
Overall, the best available information
indicates that within the contiguous
United States the wolverine’s physical
and ecological needs include:
(1) Large territories in relatively
inaccessible landscapes, at high
elevation (1,800 to 3,500 m (5,906 to
11,483 ft));
(2) access to a variety of food
resources, which vary with seasons; and
(3) physical/structural features (e.g.,
talus slopes, rugged terrain) linked to
reproductive behavioral patterns.
Current Condition
Current Potential Extent of Occurrence
As noted above, using the best
available information on current
distribution and recent occurrences, we
created maps to describe an area of
‘‘current potential extent of occurrence’’
(current potential extent) of wolverine
for the western-northwestern contiguous
United States (Service 2018, pp. 12–13,
15). The current potential extent
represents the perimeter of the
outermost geographic limits based on all
(available) occurrence records (that is,
the maximum extent of occurrences) for
the wolverine minus those areas where
we believe the species has been
extirpated (Service 2018, pp. 11–12).
The current potential extent area
identified in Figure 2 encompasses
approximately 280,316 km2 (69,267,592
acres (ac)) (Service 2018, p. 12). We also
prepared a current potential extent map
for all of North America, including
Canada and Alaska, for a total estimated
current potential extent of 8,114,878
km2 (2,005,230,024 ac) (Service 2018, p.
12 and Figure 1 of this document). The
current potential extent area in the
contiguous United States represents
approximately 3.5 percent of the total
current potential extent of wolverines in
North America (Service 2018, p. 13 and
Figure 2 of this document).
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Population Abundance and Density
Areas in the western contiguous
United States have been previously
identified as suitable for wolverine
survival (long-term survival; used by
resident adults) or primary habitat,
reproduction (used by reproductive
females), and dispersal (female and
male) of wolverines (see methodology in
Inman et al. 2013, pp. 279–280). From
these results, the researchers estimated
potential and current distribution and
abundance of wolverines in the western
contiguous United States. They
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estimated current population size of
wolverines to be 318 individuals (range
249–626) located within the Northern
Continental Divide (Montana) and areas
within the following ecoregions:
Salmon-Selway (Idaho, portion of
eastern Oregon), Central Linkage
(primarily Idaho, Montana), Greater
Yellowstone (Montana, Idaho,
Wyoming), and Northern Cascades
(Washington) (Inman et al. 2013, p.
282). Potential wolverine population
capacity based on habitat modeling was
estimated to be 644 individuals (range:
506–1,881) (Inman et al. 2013, p. 282);
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however, we do not have information
indicating wolverine abundance in the
contiguous United States.
In the wolverine SSA report, we
provide a discussion of recent studies of
wolverine detections and observations
in the western United States (Service
2018, pp. 51–56); however, no
comprehensive surveys have been
conducted across the entire area defined
as the species’ maximum extent of
occurrence (Service 2018, p. 14; Figure
2) or current potential extent of
occurrence (Figure 2 above) in the
contiguous United States. Below we
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provide a summary of recent wolverine
observations and detections in the
western United States.
A recent study (2007–2015) has
demonstrated that the Cascades region
of Washington and Canada supports a
resident wolverine population (Aubry et
al. 2016, p. 40). For the first time in
recent history, a breeding female
wolverine was detected south of I–90 in
the south Cascades of Washington, as
well as her potential mate, indicating
wolverines may be extending their
current range in that area (Flatt 2018, p.
1). Wolverines have been detected in the
Eagle Cap Wilderness Area in the
Wallowa Mountains of northeastern
Oregon in 2011–2012, 2016, and 2017
(Magoun et al. 2013, p. 17; Magoun
2017, pers. comm.). In California, a
single male wolverine occurs in the
Truckee area as of March 2017
(Shufelberger 2017, pers. comm.).
Since 2010, survey and monitoring
efforts in the Idaho Panhandle and
adjoining areas of Washington,
Montana, and British Columbia, Canada,
have detected five individual male
wolverines (Service 2018, p. 52). One
male was also detected in British
Columbia, north of Canadian Highway 3
(Lucid et al. 2016, p. 184), which some
consider to be a barrier to wildlife
passage (IDFG 2017, pers. comm.). This
male was most recently detected in
Idaho, on March 6, 2013 (Lucid et al.
2016, p. 175). One likely wolverine den
was located in the Saint Joe Mountains
in Idaho (Lucid et al. 2017, p. 12).
Results from a pilot study to evaluate
wolverine occupancy in Wyoming
indicated at least three individual
wolverines (at five stations) with at least
one individual in the Gros Ventre and
Wind River mountain ranges, and at
least two individuals in the Southern
Absaroka mountain range (Inman et al.
2015, p. 9). Occupancy modeling
estimated a probability of occupancy for
sampled sites of 62.9 percent (Inman et
al. 2015, p. 8).
Building on the results of the
Wyoming pilot study, the Western
Association of Fish and Wildlife
Agencies (WAFWA), in coordination
with Tribal partners, formed a multiState, multi-agency working group
(Western States Wolverine Working
Group) to design and implement the
Western States Wolverine Conservation
Project (WSWCP)–Coordinated
Occupancy Survey. The primary
objectives of the WSWCP include: (1)
Implement a monitoring program to
define a baseline wolverine distribution
and genetic characteristics of the
metapopulation across Montana, Idaho,
Wyoming, and Washington; (2) model
and maintain the connectivity of the
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wolverine metapopulation in the
western United States; and (3) develop
policies to address socio-political needs
to assist wolverine population
expansion as a conservation tool,
including translocation of wolverines
(IDFG 2016, pers. comm.; Montana FWP
2016, pers. comm.; WGFD 2016, pers.
comm.).
The Wyoming Game and Fish
Department began implementation of
the survey in Wyoming in the Greater
Yellowstone Ecosystem region and the
Bighorn Mountains in the winter of
2015–2016 (WGFD 2016, pers. comm.).
That initial survey detected at least
three unique wolverines in the Wind
River and southern Absaroka Mountain
Ranges (WGFD 2016, pers. comm.).
The monitoring effort was expanded
in the winter of 2016–2017 in four
States (Washington, Idaho, Montana,
and Wyoming), and our review of the
results indicate that wolverines were
detected in all four States (Service 2018,
p. 53). From this study, a total of 43
unique individuals were identified, 20
males and 23 females (Pilgrim et al.
2018, no page number).
We also received additional wolverine
observations from State and Federal
agencies in northwestern Wyoming. A
wolverine was detected by camera in
northern Grand Teton National Park,
and a member of the public reported
wolverine tracks in southwestern Grand
Teton National Park while skiing, which
was confirmed by a Forest Service
biologist (Service 2018, p. 53). Both of
these observations occurred in March
2017. South of this area in the Wyoming
Range (about 4 miles east of Alpine,
Wyoming), a wolverine was detected by
camera in May 2017 (Service 2018, p.
53).
Wolverine densities vary across North
America and have been described as
naturally low and wolverine
populations as naturally uncommon
given the species’ large home range,
wide-ranging movements, and solitary
characteristics (Service 2018, p. 56). In
the contiguous United States, density
estimates (number of wolverines per
1,000 km2 (386 mi2)) ranged from 3.5 for
the Greater Yellowstone region (2001–
2008) (areas above 2,150 m (7,054 ft)
(latitude-adjusted elevation), 4.5 for
central Idaho (1992–1995), to 15.4 for
northwestern Montana (1972–1977)
(Service 2018, p. 57).
We note here that in our 2013
proposed listing rule for the wolverine
(78 FR 7864), we discussed the
occurrences of two dispersing
individuals in California and Colorado
(the Colorado wolverine was later killed
in North Dakota). We know of one male
wolverine in California that has
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consistently occupied an area much
farther north in the Sierra Nevada
Mountains, and we have no evidence of
any other wolverines currently in the
State. We have no recent records of
wolverines in Arizona. Aubry et al.
(2007, p. 2,150) identified the year 1860
as the most recent verifiable
documentation of wolverines in
northern New Mexico. We know of no
wolverines currently occupying
Colorado. As presented in Aubrey et al.
(2007, p. 2,151; Figure 1), prior to 1900,
the most recent verifiable record for
wolverine in New Mexico was 1860 and
1887 for Nebraska; no records were
found for Arizona, Texas, Oklahoma.
This was also true for most midwestern
and mid-Atlantic States (Aubrey et al.,
2007, p. 2,152, Figure 1). Additionally,
historical range maps shown in Seton
(1909, p. 947; Map 51), Aubrey et al.
(2007, p. 2,152; Figure 1), and the
assessment and status review for the
wolverine in Canada (COSEWIC 2014,
p. 12; Figure 3) do not extend the
distribution of wolverines into these
regions.
Our updated analysis of wolverine
occurrence in the contiguous Unted
States is based on a more scientifically
robust and spatially explicit assessment
of the current areas occupied by
wolverines in the contiguous United
States, which was prepared based on
verifiable wolverine records and
comments received by reviewers of a
draft of the wolverine SSA report (see
the Historical Range and Distribution
section of the wolverine SSA report for
more on the information used to assess
the maximum extent of occurrences
(‘historical range’) and current extent of
occurrence (Service 2018, pp. 9–16;
Figures 2–4)). Using the current
potential extent of occurrence, as
presented in Figures 1 and 2 above,
provides a more accurate reflection of
the areas currently occupied by
wolverines in the contiguous United
States supported by the best available
information.
Alaska and Canada
In the wolverine SSA report, we
provide a summary of population
abundance in Alaska and Canada where
wolverines are more abundant than in
the contiguous United States (Service
2018, pp. 57–60). Much of what we
know about wolverine occurrences and
abundance has been gathered from
trapping records (see summary in
Service 2018, pp. 53–56).
In Alaska and Yukon, density
estimates presented by Inman et al.
(2012, p. 789) range from 3 to about 14
wolverines per 1,000 km2 (386 mi2),
using a number of methods. For
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example, Royle et al. (2011, p. 609)
estimated wolverine densities for
southeastern Alaska (Tongass National
Forest; 2008) from 8.2 to 9.7 per 1,000
km2 (386 mi2) (using mark-recapture),
where the higher estimate incorporates
a positive, trap-specific behavioral
response. Density of wolverines were
recently reported as an estimated 5–10
wolverines per 1,000 km2 (386 mi2)
(based on snow tracking) for
southcentral Alaska, and approximately
10 per 1,000 km2 (386 mi2) (based on
DNA mark-recapture methods) for
southeastern Alaska (Golden 2017, pers.
comm.). A wolverine occupancy study
in 2015 within an area of central Alaska
reported a density estimate of 9.48
wolverines per 1,000 km2 (386 mi2)
(Alaska Department of Fish and Game
(ADF&G) 2015, p. 7).
Wolverine density estimates for
Canada vary across regions, from 5 to 10
per 1,000 km2 (386 mi2) in northern
mountain and boreal regions to 1 to 4
per 1,000 km2 (386 mi2) in southern
boreal areas (COSEWIC 2014, p. 27).
More recently, Clevenger et al. (2017,
entire) presented a density estimate
(using spatial capture/recapture models)
for the Kootenay region of British
Columbia of 0.78 wolverines per 1,000
km2 (386 mi2), for 3 study years (2014–
2016), which they reported as lower
than expected (Clevenger et al. 2017, p.
6). Researchers in Canada are currently
conducting a landscape level analysis to
estimate the size and sustainable harvest
for wolverine populations within British
Columbia (Weir 2017, pers. comm.).
According to the most recent
COSEWIC Assessment and Status
Report on the Wolverine, Gulo gulo in
Canada (COSEWIC 2014, entire),
Canada’s western subpopulation has
been estimated at 15,688 to 23,830
adults, which we recognize is an
estimate based on several assumptions,
such as consistent trapping effort and
uniform densities across the species’
range (COSEWIC 2014, p. 36). In Alaska,
estimates of populations are not
available and are best evaluated based
on density with recent density estimates
ranging from 5 to 10 wolverines per
1,000 km2 (386 mi2) for Alaska (Parr
2017, pers. comm.). In Alaska, which,
like Canada, allows regulated hunting
and trapping of wolverines, an average
of 590 wolverines have been taken each
year over the past 6 years (Service 2018,
p. 68). The consistent harvest levels in
these regions suggest relatively stable
wolverine populations in Alaska that
more likely than not number in the
thousands of individuals in order to
sustain such level of harvest.
We do not have reliable current
population estimates for wolverines in
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the contiguous United States. As
discussed above, the only estimate
available is from 2013, when
researchers, using spatial modeling
methods, estimated the then-current
population size of wolverines to be 318
(range: 249–626) (Inman et al. 2013, p.
282). Potential wolverine population
capacity in the contiguous United States
based on habitat modeling was
estimated to be 644 individuals (range:
506–1,881) (Inman et al. 2013, p. 282).
However, these capacity estimates did
not consider spatial characteristics
related to behavior, such as territoriality
(home range), of wolverine populations.
Given all the assumptions, differing
methods of estimation, limitations, and
uncertainties of the available estimates
of North American wolverines (as
discussed in the wolverine SSA report
(Service 2018, pp. 50–56)), we believe
caution should be used relative to
comparing the number of wolverines in
the contiguous United States to the
remainder of the taxon. However, even
assuming the high population estimate
from 2013 for the contiguous United
States (n=626) and the low estimate of
wolverines in western Canada from
2014 (15,688 adults), the contiguous
United States conservatively contains
approximately 4 percent of the total
wolverines within these two regions.
This estimate does not account for
wolverines in Alaska, for which we
have no population estimate, but, based
on a rough estimate of land area for the
State occupied by wolverines and
estimated wolverine densities of
between 5 to 10 animals per 1000 km2
(386 mi2) (Parr 2017, pers. comm.), it is
reasonable to assume there are
thousands of wolverines in the State.
The actual percentage of wolverines in
the contiguous United States compared
to the overall taxon (Canada and Alaska
included) is still significantly less than
4 percent of the overall North American
wolverine population. Wolverine
densities vary across North America and
have been described as being naturally
low, due in large part to the species
having large home ranges, wide-ranging
movements, and solitary characteristics
(Service 2018, p. 56). It is important to
understand that the amount of suitable
habitat in the contiguous United States
identified both in historical and current
distribution maps (see, for example,
1909, p. 947; Map 51), Aubrey et al.
(2007, p. 2,152) does not support the
larger numbers of wolverines and higher
densities found in Canada and Alaska
(see Figure 3 in the wolverine SSA
report (Service 2018, p. 15)).
Summary of Factors Affecting the
North American Wolverine
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As mentioned above in Regulatory
Framework, a species may be
determined to be an endangered or
threatened species due to one or more
of the five factors described in section
4(a)(1) of the Act: (A) The present or
threatened destruction, modification, or
curtailment of its habitat or range; (B)
overutilization for commercial,
recreational, scientific, or educational
purposes; (C) disease or predation; (D)
the inadequacy of existing regulatory
mechanisms; or (E) other natural or
manmade factors affecting its continued
existence. Listing actions may be
warranted based on any of the above
threat factors, singly or in combination.
Potential stressors evaluated for
wolverine in the contiguous United
States include effects from roads
(Factors A and E); disturbance due to
winter recreational activity (Factors A
and E); other human disturbance
(Factors A and E); effects from wildland
fire (Factor A); disease (Factor C);
predation (Factor C); overutilization
(trapping) (Factor B); genetic diversity
(Factor E); small-population effects
(Factor E); and climate change (Factors
A and E). A summary of the potential
stressors affecting wolverine in the
contiguous United States is presented
below; for a full description of our
evaluation of the effects of these
stressors, refer to the wolverine SSA
report (Service 2018, pp. 57–101).
Effects from Roads: Wolverines are
associated with habitat found in highelevation areas, but are known to
disperse over great distances. Major
highways can present mortality risks to
dispersing individuals and affect
immigration to open territories, but
roads do not represent absolute barriers
to wolverine movements. Wolverines
den during winter months in locations
that are often inaccessible or restricted
to motorized vehicles, though secondary
roads and trails are used for winter
recreational activity. Although we
recognize there are likely additional
events that have not been reported, we
estimated the total number of wolverine
mortalities due to roads from 1972 to
2016 (44 years) in North America was
20, at least 11 of which are from Canada
(Service 2018, p. 60). In the SSA report,
we calculated a low proportion of major
highways in both modeled primary
habitat and a low mean density of roads
at high elevations where wolverines
have been observed, with the exception
of the southern Rocky Mountains
(Service 2018, p. 60). We therefore
determine that the effects from roads
present a low stressor to wolverines at
the individual and population level in
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most of its current area of occupancy
within the contiguous United States.
Disturbance due to Winter
Recreational Activity: Wolverine
behavior patterns, such as denning,
rearing of young, movement and
dispersal, and foraging/scavenging, may
be affected by recreational activities
(COSEWIC 2014, p. 42), although
several wolverines have been captured
for research on or near ski areas (e.g.,
Teton Mountains) (Montana FWP 2017,
pers. comm.). In Norway, one study
found, at the home-range scale, a
minimal threshold distance of
approximately 1.5 km (0.93 mi) for
wolverine den sites from private roads
and/or recreational cabins (May et al.
2012, p. 201). Another study found that
in an area of active recreation (Columbia
Mountains, Canada), female wolverines
were negatively associated with
helicopter and backcountry skiing in
their winter models (Krebs et al. 2007,
pp. 2,187–2,188). In summer months,
Copeland et al. (2007, p. 2,210) reported
that wolverines in their study area of
central Idaho were not uncommonly
found near maintained trails and active
campgrounds, which suggests some
level of tolerance to human presence/
recreational activity.
The Wolverine–Winter Recreation
Study represents an ongoing project to
evaluate the potential effects of
backcountry winter recreation (e.g.,
backcountry skiers, heli-skiers, catskiers, snowmobilers) on wolverines in
central Idaho and areas in the western
Yellowstone region (Island Park area
and Teton Mountains) (Heinemeyer
2016, pers. comm.; Heinemeyer 2019,
entire; Heinemeyer and Squires 2015, p.
3). Early analysis of the data suggested
that wolverines demonstrate a
behavioral response to recreation
activities, such as increased movement
rates and a reduction in resting periods
in areas of high-recreation activity,
especially high-recreation days
(Saturday and Sunday) (Heinemeyer
and Squires 2013, pp. 5, 7–8). However,
this research also found that wolverines
maintained their home ranges within
areas with relatively high winterrecreation activity over several years of
monitoring, including some areas found
to contain the highest recreational
activities (Heinemeyer 2016, pers.
comm.). The study has not been able to
determine whether these resident
wolverines are reproductively
successful due to the limited monitoring
information available for reproductive
females (Heinemeyer 2016, pers.
comm.).
A final Winter Recreation Study
report found that wolverines maintained
multi-year home ranges in areas that
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support relatively intensive winter
recreation, suggesting that wolverines
are able to tolerate winter recreation at
some scales (Heinemeyer et al. 2017, p.
iv; Heinemeyer et al. 2019, p. 16).
Wolverines responded negatively to
increasing intensity of winter recreation,
with off-road and dispersed recreation
having a greater effect than recreation
that was concentrated on access routes
(Heinemeyer et al. 2017, p. 34;
Heinemeyer et al. 2019, p. 13).
Wolverine avoidance of roads and
groomed areas used by winter
recreationists was found to be less than
estimated for dispersed recreation,
suggesting that wolverines may be less
sensitive to predictable winterrecreational use patterns (Heinemeyer et
al. 2017, p. 40; Heinemeyer et al. 2019,
p. 15). Habitat selection in females
evaluated in the multi-year study was
complex, and likely driven by a
combination of abiotic (snow, cold) and
biotic factors (predator avoidance, food
availability) (Heinemeyer et al. 2017, p.
36; Heinemeyer et al. 2019, p. 16). This
study did not assess demographic
effects, fitness effects, or population
level effects of winter recreation on
wolverines (Heinemeyer et al. 2019, p.
17 and 19).
Conservation measures currently
being implemented that address the
effects of roads in the Teton Mountains
include winter closures in certain areas
(generally from November 1 through
May 1), including road closures in the
Bridger-Teton and Caribou-Targhee
National Forests and in Grand Teton
National Park (Service 2018, p. 67,
Appendix F). These closures are being
implemented to help minimize
disturbance to wildlife (e.g., migration
pathways). State Wildlife Action Plans
prepared for individual western States
identify recreation management
strategies within wolverine habitats. For
example, in Oregon, the Oregon
Department of Fish and Wildlife
Conservation Strategy identifies
management of winter-recreation use as
a conservation action to avoid impacts
to wolverines (ODFW 2016). In
Montana’s State Wildlife Action Plan,
conservation actions for the wolverine
are identified to address potential
impacts from recreation, such as
consideration of seasonal closures
during denning season (Montana FWP
2015, p. 63). The Idaho Department of
Fish and Game Management Plan for the
Conservation of Wolverines in Idaho
also includes conservation strategies
related to developing a better
understanding of the relationships
between wolverine behavior and winter
recreation activities (IDFG 2014, p. 35),
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and the State continues to support the
Wolverine-Winter Recreation Study.
Appendix G in the SSA report provides
additional details on individual State
conservation strategies. Although we do
not rely on these conservation measures
to support our decision on listing status,
they do provide some protections to
address potential impacts to wolverine
from disturbance from winter
recreational activity and mortality from
roads.
Based on the studies summarized
above, wolverine behavior (movement)
is potentially affected by winter
recreational activity. However,
wolverines can maintain residency in
high winter-recreational use areas
(Heinemeyer et al. 2017, p. iv;
Heinemeyer et al. 2019, p. 16). Based on
the best available scientific and
commercial information, the effect of
winter recreational activity represents a
low stressor to wolverines in the
contiguous United States at the
individual and population level.
Other Human Disturbance:
Infrastructure, such as pipelines, active
logging or clearcuts, seismic lines, and
activities associated with mining (e.g.,
producing mines, mines under
development, mineral exploration
areas), may also affect individual
wolverine behavior (e.g., avoidance) or
loss or modification of wolverine
habitat. In the SSA report, we
summarize a recently published study
of habitat selection of wolverines in
response to human disturbance in
western Canadian forested habitat
(Service 2018, p. 62). That study found
that wolverines avoided interior areas of
some logged areas, but also found that
wolverines were attracted to all-season
road sections with borrow pits
(Scrafford et al. 2017, pp. 32–34). The
authors concluded that wolverine
selection patterns relative to industrial
activity and infrastructure in their study
area represented a balance between
exposure to predators and foraging
opportunities (Scrafford et al. 2017, p.
32). Based on the best available
scientific and commercial information,
we find that these human disturbance
effects are likely to be small or narrow
in scope and scale for wolverines in the
contiguous United States.
Effects from Wildland Fire: Wildland
fire can produce both direct and indirect
effects to wildlife. Direct effects include
injury and mortality as well as escape or
emigration movement away from fires
(Lyon et al. 2000, pp. 17–21). We are
unaware of any studies evaluating direct
effects of wildland fire to wolverines.
Wildland fire is likely to temporarily
displace wolverines, which could affect
home range dynamics. Given that
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wolverines can travel long distances in
a short period of time, individuals
would be expected to move away from
fire and smoke (Luensmann 2008, p.
14). In addition, because young
wolverines are born in underground or
otherwise sheltered dens during winter
months and in locations where wildland
fire risk is low due to snow cover or
increased moisture (Luensmann 2008, p.
14), the potential effects of fire at that
critical life stage is very low
(Luensmann 2008, p. 14). Indirect
effects of wildland fire can include
habitat-related effects or effects to prey
and competitors/predators; however, we
are unaware of empirical studies
evaluating these potential effects as they
relate to wolverines.
Given the diversity of habitats
occupied by wolverines, their
opportunistic foraging habitats and
seasonal switching of food sources, their
occupancy of high elevations, and
extensive mobility, wildland fire
represents a limited indirect and direct
stressor, in scope and scale, to
wolverine habitat and its prey in the
contiguous United States range (Service
2018, pp. 63–64) such that it would not
be expected to have population or
species-level impacts.
Disease: We are unaware of
comprehensive surveys evaluating the
prevalence of diseases in wolverines in
the contiguous United States. Other
than a parasitic pneumonia mortality
event and a single rabies case, we are
not aware of any other studies
documenting impacts of disease to
wolverines in North America (Service
2018, p. 65). At this time, based on the
best available scientific and commercial
information, we do not find that disease
is a population- or species-level stressor
to the wolverine in the contiguous
United States (Service 2018, pp. 64–65).
Predation: A number of potential
natural predators have been identified
for wolverines within its North
American range, including intraspecific
predation (Service 2018, p. 65).
However, we have no information that
suggests predation represents a
significant stressor to the wolverine at
the population level. At the individual
level, we recognize that wolverines
likely avoid areas of potential predation
risk from wolves and other potential
predators (Service 2018, p. 65). Thus,
indirect effects of predators may result
in predator avoidance behavior of
individual wolverines through habitat
selection. However, the best scientific
and commercial information available
indicates that predation is not a stressor
for the wolverine (Service 2018, p. 65).
Overutilization for Commercial,
Recreational, Scientific, or Educational
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Purposes: During the late 1800s and
early 1900s, the wolverine population
declined or was extirpated in much of
the contiguous United States, which has
been attributed in large part to
unregulated persecution (Service 2018,
p. 1). Similar range reductions and
extirpations of some wolverine
populations were observed in parts of
Canada during this time period (van
Zyll de Jong 1975, entire; COSEWIC
2014, p. iv). However, after unregulated
harvest of wolverines ceased, the
numbers of wolverines in Canada and
the contiguous United States began to
recover from this decline (e.g., Aubry et
al., 2007, p. 2,151; Aubry et al., 2012,
entire; Aubry et al. 2016, pp. 14–15;
Magoun et al. 2013, p. 27).
In Montana, wolverines were a legally
harvested furbearer up until 2012
(Service 2018, p. 65). There is, however,
no evidence to suggest that the harvest
of wolverines in Montana at historical
rates (about 10 animals per year) was
detrimental to wolverine populations
(Service 2018, pap. 65–66 and
Appendix G). Furthermore, States
within the wolverine range in the
contiguous United States have adopted
protective regulations to prevent
unauthorized take and are
implementing other measures to limit
incidental mortality of wolverines
(Service 2018, p. 66). There is currently
no allowable trapping or harvesting of
wolverines in the contiguous United
States, although incidental trapping,
shooting, and poisoning mortalities
have been documented (Service 2018,
pp. 65–69).
In Alaska, wolverine trapping and
hunting is controlled by seasons and bag
limits, with about 550 animals
harvested each year (ADF&G 2017a).
This level of harvest has been fairly
consistent since 2010 (Service 2018,
Table 7).
Trapping and harvesting of
wolverines occurs over much of the
range in Canada (COSEWIC 2014, pp.
10, 29–35). Specifically, wolverines are
harvested in the northern and western
territories—Manitoba, Saskatchewan,
Alberta, British Columbia, Yukon,
Northwest Territories, and Nunavut
(COSEWIC 2014, p. 43). The population
of wolverines in British Columbia is
estimated to be 2,700–4,760 and 1,500–
2,000 animals in Alberta (COSEWIC
2014, p. 36). In the wolverine SSA
report, we evaluated trapping of
wolverines in British Columbia and
Alberta regions of southern Canada in
an effort to document potential impacts
to dispersing wolverines along the U.S.–
Canada border (Service 2018, pp. 68–
69). This type of analysis was not
conducted for the 2013 proposed listing
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rule (78 FR 7864; February 4, 2013) or
for our 2014 withdrawal (79 FR 47522;
August 13, 2014). The results of our
spatial analysis for British Columbia
indicates a total of 77 wolverines were
trapped in wildlife management units
within 110 km (68.35 mi) of the U.S.–
Canada border in the period 2007–2015,
or an average of 8.5 animals per year
(Service 2018, pp. 68–69). We used this
distance since it is similar to both the
average maximum distance per
dispersal movement of 102 km (63 mi)
for male wolverines in the Greater
Yellowstone region of Montana (Inman
et al. 2012, p. 784), and a reported 100km (62-mi) dispersal distance for a
juvenile male for Ontario, Canada
(COSEWIC 2014, p. 24, citing
unpublished data from Dawson et al.
2013). For Alberta, we identified a total
of 15 wolverines harvested by trappers
and data presented in other studies
within 110 km (68.35 mi) of the U.S.—
Canada border in the period 1989–2014
(average of less than 1.0 animal per
year) (Service 2018, p. 68).
Based on this new analysis, legal
trapping effort along the U.S.–Canada
border does not represent a barrier to
wolverine movement and dispersal
along the international border. As
discussed below and in the DPS
analysis above, results from genetic
analyses provide further evidence of
movement and dispersal of wolverines
across the international border (see
Genetic Diversity below).
In summary, overutilization does not
currently represent a stressor to the
wolverine in the contiguous United
States at the individual, population, or
species level. Wolverine populations in
the contiguous United States are
currently protected under several State
laws and regulations. Regulated hunting
and trapping activities for wolverines
are currently suspended or closed
entirely for animals that occupy western
States of the contiguous United States,
though occasional incidental trapping
can occur. Current trapping in Alaska
and Canada appears to be sustainable
and wolverine populations along the
Alaska—Canada border are continuous
with the Yukon region of Canada, which
suggests a rescue effect (animals from a
higher population density area moving
to areas of lower population density,
preventing local extirpation) for
Canadian populations along this
international boundary (COSEWIC 2014,
p. 37). Trapping or harvesting of
wolverines along the contiguous U.S.–
Canada border does not represent a
barrier or stressor to wolverines
migrating into the contiguous United
States at the individual or population
level.
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Genetic Diversity: The geographical
genetic structure of wolverines is
believed to be largely structured around
the strong female philopatry
characteristic of this species (Rico et al.
2015, p. 2) and the species’ polygamous
behavior. Results from Scandinavia
indicate that wolverine population
distributions are primarily limited by
dispersal of the more philopatric sex
(females) (Aronsson 2017, p. 13). The
extensive and often asymmetrical
movement of male wolverines from core
populations to the periphery of their
range can result in the addition of
nuclear genetic material to these edges
(Zigouris et al. 2012, p. 1,553). Thus, the
dispersal pattern for male wolverines
may help explain why allelic richness
(i.e., nuclear DNA, which is inherited
from both parents) can be similar across
regions, but haplotype richness
(mitochondrial DNA, which is
maternally inherited) is lower at the
periphery of the species’ range (Zigouris
et al. 2012, p. 1,553).
Studies evaluating the genetic
structure of wolverines, primarily
within its core range in North America,
were presented in Chappell et al. (2004)
and Kyle and Strobeck (2001, 2002).
Using microsatellite markers, Kyle and
Strobeck (2002) and Zigouris et al.
(2012) found greater genetic structure of
wolverines toward the eastern and
southern peripheries of their North
American distribution, likely due to a
west-to-east recolonization during the
Holocene (Zigouris et al. 2013, p. 9).
Similarly, based on an evaluation of
mitochondrial DNA, which is used
primarily for an evaluation of
phylogenetic structure and
phylogeography, McKelvey et al. (2014,
p. 330) concluded that modern
wolverine populations in the contiguous
United States are the result of
recolonization (following persecution
during a period of unregulated hunting
or trapping and poisoning) from the
north.
Genetic diversity and population
genetic structure of a larger sample size
of wolverines were examined by
Cegelski et al. (2006, entire) for the
southern extent of their North American
range using both microsatellite markers
and mitochondrial DNA. They
concluded that the wolverine
populations in the contiguous United
States were not sources for dispersing
individuals into Canada (Cegelski et al.
2006, p. 208). They found that there was
significant differentiation between most
of the populations in Canada and the
United States (Cegelski et al. 2006, p.
208). However, they cautioned that their
statistical analysis may not have been
able to detect ‘‘effective migrants’’ and
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that sample size can affect the detection
of dispersers (Cegelski et al. 2006, p.
208). They concluded that some
migration of wolverines was occurring
between the Rocky Mountain Front
region (northwestern Montana) and
Canada as well as among wolverine
populations in the United States, with
the exception of Idaho (Cegelski et al.
2006, p. 208).
This study also conducted model
simulations of the number of effective
wolverine breeders necessary to
maintain genetic variation
(heterozygosity) in their sampled
population of the contiguous United
States in the absence of gene flow
(Cegelksi et al. 2006, p. 201). They
indicated that two effective migrants
from either Canada or Wyoming into the
Rocky Mountain Front population
would be needed (per generation, 7.5
years) to maintain the levels of genetic
diversity in that population, and one
effective migrant was needed to
maintain levels of diversity in the
Gallatin, Crazybelt, or Idaho
populations (Cegelski et al. 2006, p.
209). They also found that to maintain
at least 95 percent of the genetic
variation in the next 100 generations
(we estimate this to be approximately
750 years, based on generation time) 200
to 300 wolverine breeding pairs were
needed in the Wyoming and Rocky
Mountain Front populations,
respectively, and 200 breeding pairs
were needed in the Gallatin, Crazybelts,
and Idaho wolverine populations
(Cegelski et al., 2006, pp. 208–209). The
authors concluded that migration is
essential for maintaining diversity in
wolverine populations in the contiguous
United States since effective population
size may never be reached due to the
naturally low population densities of
wolverines (Cegelski et al. 2006, p. 209).
More recently, an analysis of
mitochondrial DNA was prepared for
wolverine samples collected during the
winters of 2015–2016 and 2016–2017 as
part of the Western States Wolverine
Conservation Project–Coordinated
Occupancy provides further support
that all contiguous United States
historical (pre-1900) and recent
wolverine populations are likely
descendants of immigrants from Canada
and suggest continued connectivity
between the contiguous United States
and Canadian wolverine populations
(Pilgrim and Schwartz 2018, entire).
Effective population size (Ne) is
defined as ‘‘the size of an idealized
population that would experience the
same amount of genetic drift and
inbreeding as the population of interest.
In popular terms, Ne is the number of
individuals in a population that
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contribute offspring to the next
generation’’ (Hoffman et al. 2017, p.
507; see also Service 2018, Box 2). It
represents a metric for quantifying rates
of inbreeding and genetic drift and is
often used in conservation management
to set genetic viability targets (Olsson et
al. 2017, p. 1). It is not the same as the
more commonly used metric, census
population size (N), but is often
assumed to represent the genetically
effective population size.
In his review of the minimum viable
population size concept, Ewens (1990,
entire) emphasized that the term
‘‘effective population size’’ is not a
meaningful term unless additional
context is provided relative to which
concept of population size is being
evaluated (Ewens 1990, p. 309). He
introduced the concept of mutation
effective population size, defined as the
size of population defined by its
capacity to maintain genetic variation
(Ewens 1990, p. 307), which is different
than actual population size (Ewens
1990, p. 309). Demographic factors such
as sex ratio, subpopulations, dispersal,
and immigration are needed when
interpreting actual population size from
an effective population size; thus, there
is no justification for a fixed, genetically
derived minimum viable population
size value of ‘500’ as each case is unique
(Ewens 1990, p. 310). A review of the
minimum viable population concept by
Flather et al. (2011, entire) also found
that any ‘‘rule of thumb’’ used for
minimum viable population will likely
be a poor estimate for that population
(Flather et al. 2011, pp. 311, 313).
Minimum viable population estimates
therefore vary considerably both within
and among species and are sensitive to
the timeframe in which data are
collected (Flather et al. 2011, p. 314).
An effective population size analysis
for wolverines in the contiguous United
States was presented in Schwartz et al.
(2009, p. 3,225) using wolverine
samples from the main part of the Rocky
Mountains populations (e.g., central and
eastern Idaho, Montana, northwestern
Wyoming). Subpopulations from the
Crazy and Belt Mountains in Montana
were excluded from this analysis based
on suggestion by Cegelski et al. (2003)
that they represented separate groups
(Schwartz et al. 2009, p. 3,225). The
summed effective population size was
estimated at 35, with credible limits
from 28–52, and the summed values for
the three timeframes was reported as
follows: Ne 1989–1994 = 33, credible
limits 27–43; Ne 1995–2000 = 35,
credible limits 28–57; Ne 2001–2006 =
38, credible limits 33–59 (Schwartz et
al. 2009, p. 3,226). Thus, the two later
time-frames evaluated indicate an
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(increasing) effective population size
with credible limits above 50.
Of direct relevance to potential gene
flow and genetic structure at the
landscape level, wolverines travel
(disperse) through areas outside highelevation, forested habitats. For
example, tracked dispersal movements
of a male wolverine, M56, from
Wyoming into Colorado and its
subsequent discovery in North Dakota,
indicate extensive travel outside of
modeled primary wolverine habitat (i.e.,
Inman et al. 2013), including through
arid grasslands and shrubland habitats
of the Wyoming Basin ecoregion
(Packila et al. 2017, entire). This
animal’s movement also supports some
level of connectivity (and gene flow)
between currently occupied habitat
(Wyoming) and unoccupied habitat
within the wolverine’s historic range
(Colorado) (Packila et al. 2017, p. 404).
Similarly, wolverines have recently
moved from Washington and Idaho into
British Columbia, and earlier from
Montana to British Columbia and
Alberta (Service 2018, p. 45). Based on
genetic analyses, the male wolverine
currently occupying an area within the
Sierra Nevada Mountains of California
also represents evidence of connectivity
between wolverine populations of the
Rocky and Sierra Nevada Mountain
Ranges (Moriarty et al. 2009, p. 154).
Wolverines have been detected making
cross-valley movements in the
Southwestern Crown of the Continent
(SWCC) in northwestern Montana,
which researchers believe is an
indication of good connectivity in this
region (SWCC Working Group 2016,
pers. comm.).
It can be difficult to make inferences
about the relationship between
population size and point estimates of
genetic diversity without continued
genetic monitoring and an
understanding of the demographic
history of a species’ population
(Hoffman et al. 2017, p. 507), including
factors that have historically influenced
and continue to influence movement
patterns and connectivity. Additionally,
the extensive dispersal movements of
both male and female wolverines can
produce gene flow among diverged
populations, making it difficult to
distinguish, without additional
sampling and analysis, between longdistance dispersal and fragmentation
based on the patchy distribution of
some haplotypes (Zigouris et al. 2013, p.
10). Genetic diversity can be a reflection
of favorable adaptations (natural
selection) and is necessary for species to
locally adapt to environmental stressors
or to facilitate range shifts (Zigouris et
al. 2012, p. 1,544). Genetic
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distinctiveness in peripheral
populations may therefore play a role in
both maintaining and generating
biological diversity for a species
(Zigouris et al. 2012, p. 1,544; citing
results presented in Channell and
Lomolino 2000, p. 84). Relatedly,
genetic variation that is adaptive is a
better predictor of the long-term success
of populations as compared to overall
genetic variation (Hoffman et al. 2017,
p. 510). The challenge is to be able to
determine whether genetic variation is
adaptive and is a reflection of remnants
of high genetic diversity from ancestral
populations, or whether that variation is
a reflection of accumulated deleterious,
nonadaptive genes due to genetic drift
in small populations (Hoffman et al.
2017, p. 509).
In summary, the currently known
spatial distribution of genetic variability
in wolverines in North America appears
to be a reflection of a complex history
where population abundance has
fluctuated since the time of the last
glaciation and insufficient time has
passed since human persecution for a
full recovery of wolverine densities
(Cardinal 2004, pp. 23–24; Zigouris et
al. 2012, p. 1,554). Zigouris et al. (2012,
p. 1,545) noted that the genetic diversity
reported in Cegelski et al. (2006) and
Kyle and Strobeck (2001, 2002) for the
southwestern edge of the North
American range represented only part of
the diversity in the northern
populations of wolverines. Zigouris et
al. (2012, p. 1,545) posit that the
irregular distribution of wolverines in
the southwestern periphery and the
genetic diversity observed in those
analyses is a result of population
bottlenecks that were caused by range
contractions from a panmictic (random
mating) northern core population
approximately 150 years ago coinciding
with human persecution. As described
here, recent dispersals of wolverines
into Colorado, California, and Utah
provide evidence for connectivity and
the potential for gene flow between
Northern Rocky Mountain populations
and areas where wolverines were
extirpated.
As noted above in this section (and in
the Distinct Population Segment
section), there is recent evidence of
wolverines traveling across the
international border. Furthermore, our
analysis of trapping levels in the
wolverine SSA report (summarized in
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes above) does not support
previous assumptions that trapping in
Canada near the border acts as a barrier
to wolverine movement into the
contiguous United States. Cegelski et al.
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(2006, p. 209) determined that very few
successful migrants are needed per
generation to maintain at least 95
percent of the genetic variation in the
next 100 generations (approximately
750 years) in the contiguous United
States (Cegelski et al. 2006, p. 209). We
have no reason to believe that this level
of migration from the north has not
already been occurring following the
end of intense persecution of this
species to repopulate previously
occupied areas within the contiguous
United States. This repopulation has
occurred without human-assisted
introductions and with unregulated
trapping from about the 1930s to 1970
in Montana. Given the recent
observations of dispersing wolverines
moving vast distances over varied
terrain and movement of wolverines
across the U.S.–Canada border, our
recent assessment of the low levels of
trapping mortality in Canada near the
border, and further confirmation of
Canada as the source of wolverine
genetics present in contiguous United
States wolverines, we conclude that
wolverines in the contiguous United
States are not separated genetically from
the larger population in Canada.
Furthermore, even if they were
separated genetically, the multiple
generations it would take for genetic
isolation to potentially result in
significantly lower genetic diversity and
for the deleterious effects of decreased
genetic diversity to then manifest into
negative population-level effects is
likely beyond the foreseeable future
used for this determination (38 to 50
years, see Future Condition section
below). As such, we conclude that loss
of genetic diversity is not a stressor for
wolverines in the contiguous United
States now or within the foreseeable
future.
Small Population Effects: As
described above in Population
Abundance and Density, the number of
wolverines in the contiguous United
States is relatively small compared to
the remainder of the range in Canada
and Alaska, in large part due to limited
suitable habitat and previous
persecution and unregulated trapping
pressures. As described above in
Genetic Diversity, we now consider
wolverines in the contiguous United
States to be genetically connected to
wolverines in Canada (McKelvey et al.
2014; Pilgrim and Schwartz 2018) and
wolverines in the contiguous United
States are not separated from the larger
North American wolverine population
to the North (Canada and Alaska). In
previous proposed rules and findings,
we have discussed small population
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size as a vulnerability that places
wolverines in the contiguous United
States at risk of extirpation. However,
those assertions were predicated on a
belief that wolverines in the contiguous
United States were effectively isolated
regionally within the United States and
isolated from Canada, thereby
increasing the risk of deleterious genetic
effects (countered above in Genetic
Diversity) and susceptibility to
stochastic events and limited rescue
effect (migrants) from Canada. With
further genetic evidence of the
recolonization of once-extirpated areas
of the contiguous United States by
wolverines from Canada postunregulated trapping over the last
approximately 100 years, history has
demonstrated the resiliency of the North
American wolverine population to
recover from extreme persecution and
unprecedented direct mortality. We do
not currently foresee any stochastic or
catastrophic events that could result in
a similar population-level effect on
wolverines in the contiguous United
States. It is no longer accurate to
consider contiguous United States
wolverines in isolation from the rest of
North American wolverines; rather, it is
more accurate to consider the
contiguous United States wolverines a
portion of a much larger and proven
resilient North American wolverine
population. We conclude that small
population effects are not a stressor for
wolverines in the contiguous United
States now or within the foreseeable
future.
Climate Change: In the SSA report,
included in our discussion of future
conditions, we provide a summary of
current trends related to observed
climate change effects, such as
increased temperatures and changes in
precipitation patterns, in areas that
encompass the current potential extent
of occurrence for the wolverine. We are
not aware of any adverse effects of these
observed changes to the wolverine in
the contiguous United States. The
potential effects of future climate
change (projections) are fully
considered in our future condition
analysis in the wolverine SSA report
(Service 2018, pp. 73–99). We
summarize the results of that analysis in
the Future Condition section of this
document below.
Summary of Current Condition
Wolverine populations in much of
North America are still recovering from
large losses of individuals from
intensive hunting and unregulated
persecution pressures in the late 1880s
into the mid-20th century (Service 2018,
p. 104). The distribution of wolverines
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within suitable habitat provides a more
appropriate method for estimating
population status than using abundance
of animals, although there is limited
rangewide survey information. Based on
the best available information,
wolverines continue to be detected
within suitable habitat within the
western-northwestern contiguous
United States including Washington,
Oregon, Idaho, Montana, and Wyoming
(Service 2018, p. 71). Studies are
currently under way to provide a better
assessment of the species’ current
distribution and genetic characteristics
of these populations. The best available
information does not indicate the
portion of the North American
wolverine population in the contiguous
United States is currently negatively
impacted by lower genetic diversity,
and there is no evidence that wolverine
numbers in the contiguous United
States are declining.
We prepared a map of the current
potential extent of occurrence to
illustrate the species’ current
distribution in the contiguous United
States (Figure 2). We estimate this area
represents approximately 3.5 percent of
the wolverine’s current potential extent
in North America (Service 2018, p. 71).
We determined that 72 percent of our
current potential extent of the wolverine
in the contiguous United States is found
on lands owned or managed by the
Federal Government (Service 2018, p.
72 and Appendix D). We also evaluated
previously modeled wolverine primary
habitat in the contiguous United States
(Inman et al. 2013, entire) and estimated
that 96 percent of this area is owned or
managed by Federal agencies and 41
percent of this area is located in
designated wilderness areas (Service
2018, p. 72). In our SSA report, we
provide a detailed summary of
regulatory mechanisms and
conservation measures affecting
wolverines related to State and Federal
land management in the contiguous
United States (Service 2018, Appendix
G).
We evaluated several potential
stressors that may be affecting wolverine
populations or its habitat, including
effects from roads, disturbance due to
winter recreation and other activities,
effects from wildland fire, disease and
predation, overutilization for (primarily)
commercial purposes, genetic diversity,
and small-population effects.
We determined that the effects of
roads (evaluated by number of miles,
density, and location) and disturbance
represent low-level stressors to the
wolverine in the contiguous United
States. Wildland fire was determined to
be a short-term stressor to wolverine
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habitat and its prey. Disease and
predation, genetic diversity, and small
population size are not considered
stressors to the wolverine.
Trapping or hunting of wolverines is
currently prohibited in the contiguous
United States. Incidental trapping of
wolverines is infrequent in the
contiguous United States and, in Idaho
and Montana, education programs are
being implemented to reduce this
stressor. Wolverines are harvested in
several Canadian provinces and near the
U.S.–Canada border with management
and monitoring oversight based on
spatial and temporal elements. We
evaluated historical trapping
information to assess potential impacts
to dispersing wolverines into the United
States. Based on the best available
commercial and scientific information,
overutilization does not represent a
stressor to the wolverine in the
contiguous United States.
We also determined that the
wolverines in the contiguous United
States are connected to and an extension
of the Canadian population that is not
genetically isolated nor considered a
small population that may be more
vulnerable to stressors.
Future Condition
The foreseeable future timeframe
evaluated in our SSA analysis is
approximately 38 to 50 years, which
captures consideration of the projected
future conditions related to trapping/
harvesting, climate change, or other
potential cumulative impacts (Service
2018, p. 73). We use a timeframe of
approximately 38 to 50 years because,
beyond this range, climate modeling
uncertainty increases substantially. We
believe this is a reasonable timeframe to
consider as it includes the potential for
observing these effects over several
generations of the wolverine.
Evaluations of future conditions for
species have an inherent level of
uncertainty relative to demographic
risks, particularly those related to
climate change projections. After
considering the current conditions for
the wolverine and its habitat, we
determine that climate change effects
(i.e., significantly elevated temperatures
resulting in decline in snowpack) that
may modify suitable habitat, including
reproductive denning habitat, could also
change the scope of the wildland fire
stressor and is the most likely future
scenario to potentially have an effect on
wolverines at the population level in the
contiguous United States. Based on our
review of the best available information,
we determined that there were no other
plausible future scenarios that were
likely to have population-level impacts
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to wolverine in the contiguous United
States (Service 2018, p. 73). As
described in detail in the wolverine SSA
report (Service 2018, pp. 57–72), the
effects of disease, predation,
overutilization (trapping), genetic
diversity, small-population effects, and
effects of wildland fire are expected to
continue to be at low levels in the future
but are not expected to result in
population-level effects to wolverine.
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Climate Change Effects
In the wolverine SSA report, we
considered climate changes that may
affect environmental conditions upon
which the wolverine relies. As defined
by the Intergovernmental Panel on
Climate Change (IPCC), the term
‘‘climate’’ refers to the mean and
variability of different types of weather
conditions over time (IPCC 2013, p.
1,450). Thus, the term ‘‘climate change’’
refers to a change in the mean or the
variability of relevant properties, which
persists for an extended period,
typically decades or longer, due to
natural conditions (e.g., solar cycles) or
human-caused changes in the
composition of atmosphere or in land
use (IPCC 2013a, p. 1,450).
Multiple lines of evidence, not just
projections derived from quantitative
models, should be examined when
conducting climate vulnerability
assessments (Michalak et al. 2017,
entire). Thus, we evaluated projected
effects from climate change in the
western United States relative to both
abiotic (e.g., temperature, precipitation,
snow cover) and biotic (e.g., phenology,
behavior) factors. Refer to the wolverine
SSA report for a complete discussion of
our analysis of the effects of climate
change to wolverine in the contiguous
United States (Service 2018, pp. 73–99).
We summarize the results of that
analysis below.
Summary of Future Condition
Abiotic Factors: Observed trends and
future climate model projections
indicate warming temperatures for
much of the western United States,
including areas within the current
potential extent of the wolverine
(Service 2018, pp. 75–81). The degree of
future warming varies by region and is
dependent upon the future emission
scenario used during the modeling
process. Future precipitation trends are
less certain for many regions, in part,
due to naturally high inter-annual
variability; some regions are projected to
experience greater winter precipitation
(Service 2018, p. 81). Wolverines have
been found to have a wide range in their
physiological critical temperature
depending on season and undergo
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seasonal changes in fur insulation to
adapt to warmer temperatures in
summer (Service 2018, p. 81).
Wolverines also exhibit changes in
behavior, such as moving to higher
elevations in summer months (Service
2018, p. 81). Wolverines continue to
occupy areas that have exhibited
increases in temperature (e.g.,
California, parts of Montana and
Washington) due to effects of climate
change; however, no empirical studies
have evaluated these physiological and
behavioral adaptations, including
sublethal effects, relative to warming
temperatures (Service 2018, p. 81).
Biotic Factors: In addition to
evaluating changes in abiotic factors,
biotic interactions should be considered
in evaluating species’ response to
climate change (reviewed by Post 2013).
Although abiotic changes drive
ecological processes, the alterations in
biotic interactions (e.g., competition
among conspecifics, interactions with
competitors, resources, and predators)
represent the ecological responses that
result from those changes (Post 2013, p.
1). Changes in certain abiotic factors,
such as snow and ice cover, should also
be considered in an ecological context
since they represent habitat for many
species (Post 2013, p. 11).
The results presented in the
wolverine SSA report indicate biotic
effects resulting from climate change,
varying from phenological changes to
shifts in vegetation and vegetation
succession (Service 2018, pp. 81–82).
We are unaware of studies that have
directly evaluated these types of effects
to the North American wolverine or its
habitat. Given the relatively large area
and varied habitats occupied by
wolverines in the contiguous United
States, the projected shifts in vegetation
are likely to be relatively narrow in
scope and scale relative to potential
effects to wolverines. Furthermore, we
have no information to suggest that
wolverines selectively use any specific
vegetation type, and some projected
changes in vegetation may be
advantageous for wolverine prey
(Service 2018, p. 82).
Climate Change and Potential for
Cumulative Effects
Threats can work in concert with one
another to cumulatively create
conditions that may impact the
wolverine or its habitat beyond the
scope of each individual threat (Service
2018, p. 82). Given an expected increase
in temperature in the western United
States, the best available information
indicates that, if there are any
cumulative impacts in the future, the
most likely population-level effects on
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wolverine in the contiguous United
States could be: (1) Changes in
snowpack from the combination of
increased temperature and changes in
precipitation patterns, or (2) changes in
snowpack and increase in wildland fire
potential (Service 2018, p. 83).
Snowpack/Snow Cover: The effects of
climate change on snow persistence has
been suggested as an important negative
impact on wolverine habitat and
populations by the mid-21st century
and was the primary basis of our 2013
proposed rule to list the North
American wolverine in the contiguous
United States (78 FR 7864; February 4,
2013). In light of the court decision
remanding our consideration of our
withdrawal of the 2013 proposed rule
relative to climate change effects to
wolverine, the Service pursued a refined
methodology to provide insights into
the potential impacts of climate change
on snow persistence (Service 2018, p.
85; Ray et al. 2017, entire).
The Service engaged the National
Oceanic and Atmospheric
Administration and University of
Colorado in Boulder, Colorado, to
evaluate and model fine-scale
persistence of snow in occupied and
potential wolverine habitat in the
contiguous United States. The primary
objective of this study was to refine
spatial and temporal scale of snow
modeling efforts and improve the
scientific understanding of the extent of
spring snow retention currently and into
the future under a changing climate
(Ray et al. 2017, p. 9). The objectives of
the study included (Ray et al. 2017, p.
10):
• Use of fine-scale models to analyze
the topographic effects of snow,
including slope and aspect (compass
direction that slope faces).
• Use of a range of plausible future
climate change scenarios to assess snow
persistence.
• Analysis of extremes and year-toyear variability by selecting
representative wet, dry, and near normal
years (using observed conditions) and
then modeling changes for those base
years under several future climate
scenarios.
• Assessment of changes in snow
persistence by elevation.
The study was designed to parallel as
much as possible and thereby refine the
previous assessment of snow cover
persistence in the western United States
presented in McKelvey et al. (2011).
However, an exact replication of the
McKelvey et al. (2011) study was not
possible given the time, funding, and
computational constraints needed to
develop a fine-scale assessment. The
current study was limited to two study
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areas (approximately 1,500 to 3,000 km 2
(579 to 1,158 mi 2) each) in the northern
and southern Rocky Mountains (see
Service 2018, Appendix H). These two
National Parks bound the Northern and
Southern part of the wolverine historic
range, and were selected because they
encompass the latitudinal and
elevational range of wolverines within
the contiguous United States. Glacier
National Park is representative of a
high-latitude and relatively lowelevation area currently occupied by
wolverines. The Rocky Mountain
National Park region is a lower latitude
and higher elevation area within the
wolverine’s historical range, which was
recently occupied by a wolverine from
2009 to at least 2012. See the wolverine
SSA report for a summary of the
methods used in Ray et al. (2017)
(Service 2018, pp. 86–87).
Comparison with McKelvey et al.
(2011): Although the methods used in
this study have similarities with those
presented in McKelvey et al. (2011),
there are several key differences.
• Ray et al. (2017) used a finer spatial
resolution model than McKelvey et al.
(2011) (0.0625 km2 vs. 37 km2) (see
Service 2018, Appendix I for a
comparison figure) that also
incorporated slope and aspect.
• The grid cells represented in
McKelvey et al. (2011) were assumed to
be flat (i.e., north-facing slopes treated
as identical to south-facing slopes).
• McKelvey et al. (2011) focused on
May 1 snow depth as a proxy for May
15 snow disappearance, while Ray et al.
(2017) focused directly on May 15 snow
disappearance and produced results for
the presence or absence of deeper snow
(nominally greater than or equal to 0.5
m (20 in) depth) on May 1 and April 15.
Ray et al. (2017) originally focused on
May 15 to compare to the McKelvey et
al. (2011) study, and June 1 to bracket
the snowmelt season. However, the
April 15 and April 30 dates were added
to the evaluation of snow-covered areas
to align with temporal reproductive
patterns of the wolverine (see Use of
Dens and Denning Behavior discussion
in the Reproduction and Growth section
of the wolverine SSA report (Service
2018, pp. 23–28)).
• Because of the increased resolution
of this study, Ray et al. (2017) were able
to consider whether any areas of snow
with depth greater than 0.5 m (20 in)
will persist in these areas in the future
at time periods encompassing the end of
the wolverine denning period.
Additional comparisons are outlined in
the wolverine SSA report (Service 2018,
p. 88; Table 8) and our rationale
supporting the use of snow depth
greater than 0.5 m (20 in) is documented
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in the wolverine SSA report (Service
2018, p. 87) and in Ray et al. (2017;
Table 5–2).
Interpretation of results and
additional analysis relative to wolverine
den site scale: Recent studies of
wolverine populations and distribution
in Sweden have observed wolverine
populations and reproductive den sites
outside areas modeled with persistent
spring snow cover (Aronsson and
Persson 2016, p. 266; Persson 2017,
pers. comm.). Another recent study,
from Canada, concluded wolverines are
adaptable and do not require large areas
of deep spring snowpack for successful
reproduction, and may select small
areas covered with deep snow at a finer
scale than can be detected using satellite
imagery (Webb et al. 2016, p. 1,468).
Jokinen et al (2019) reported seven
wolverine den sites in hollow mounds
(caused by the uplifted root masses from
fallen Black Spruce trees) in the boreal
forest of Alberta. These areas were
largely devoid of spring snow cover
(mean distance from dens to nearest
spring snow cover was 15.2 km) and the
authors stated wolverines appear to be
using ‘‘locally-available denning
structures in the lowland boreal forest,
despite a lack of deep snow, persistent
spring snow cover, or large boulders
documented in other studies.’’
Regardless as to whether or not
wolverines are obligated to den in areas
of deep snow, the Service was interested
in exploring the question, ‘‘If snow
cover is required for wolverine denning,
will there be a sufficient amount of
significant snow cover in the future in
areas wolverines have historically used
for denning in the contiguous United
States?’’ The Service integrated future
Distributed Hydrology Soil Vegetation
Model projections (2000–2013 averages)
of snow-covered area (greater than 0.5 m
(20 in) depth) on May 1 for Glacier
National Park and Rocky Mountain
National Park with new information
obtained from a spatial analysis of
documented den sites in the contiguous
United States. This analysis indicated
31 of 34 documented den sites in the
contiguous United States were located
in areas with slopes less than 25
degrees. Avalanche risk increases
significantly in areas with slope greater
than 25 degrees (Scott 2017, pers.
comm.) and thus wolverines maybe
avoiding these areas for denning due to
this risk (Service 2018, p. 91).
The Service calculated areal estimates
for future snow covered area in both
study sites and limited these estimates
to elevation bands wolverines have used
historically for denning and for areas
with slopes less than 25 degrees. This
approach resulted in providing the most
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conservative estimates of future snow
covered area in the areas wolverines are
most likely to use for denning.
Using the projections prepared by Ray
et al. (2017), the wolverine SSA report
presents the spatial distribution of
significant snow-covered area with
slopes less than 25 degrees and within
the elevation bands expected to be used
by wolverines for denning for three
future climate scenarios in each study
area (Service 2018, pp. 92–98). The
three scenarios for Glacier National Park
and Rocky Mountain National Park were
chosen to span the range of Global
Climate Model uncertainty regarding
temperature and precipitation, and by
extension significant snow-covered area
(Service 2018, p. 93). A detailed
description of methods describing the
process of Global Climate Model
selection can be found in Ray et al.
(2017, pp. 35–38). We found that large
portions of the study areas meet all
three criteria—greater than 0.5 m (20 in)
snow depth on May 1, at elevation
1,514–2,252 m (4,967–7,389 ft) for
Glacier National Park or 2,700 to 3,600
m (8,858 to 11,811 ft) for Rocky
Mountain National Park, and with
slopes less than 25 degrees—across both
study sites in the future (See map
legends in Figures 10–15 in the SSA
report, (Service 2018, pp. 94–98)).
We also determined that large tracts
(several hundred km2/mi2) of significant
snow (greater than 0.5 m (20 in) in
depth) are projected in close proximity
to documented historical den sites
across all three climate scenarios
(Service 2018, pp. 94–95). This analysis
is limited to Glacier National Park
because this is the only area where new
snow-covered area projections and
historical den locations were both
available. Wolverines would not have to
travel far, or at all, relative to either
distance or elevation to reach areas with
significant snow-covered area for
denning in the future (Service 2018, pp.
94–95).
Based on the best available
information, we have no reason to
believe wolverines are confined to
previously modeled spring snow
covered areas. Furthermore, there is no
quantitative data documenting spring
snow patch size or depth to the denning
needs of wolverines. Even if wolverines
must have spring snow for denning,
which we do not believe to be true, the
wolverines in the lower 48 will likely
have access to areas with significant
spring snow cover in the future. Based
on the new information presented above
and in the wolverine SSA report, we do
not believe wolverines need spring
snow cover for denning. Nevertheless,
new information suggests that spring
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snow cover will not be a limiting
resource for wolverines in the
contiguous United States in the future.
Therefore, based on the best available
information, we do not consider the
effects of changes in snowpack from the
combination of increased temperature
and changes in precipitation patterns to
be a threat to the wolverine.
Wildland Fire
The wolverine SSA report includes a
discussion of available information on
the relationship of predicted future
climate conditions on wildland fire
projections in the western United States
(Service 2018, pp. 99–100). In summary,
based on these projections, wildland fire
risk is likely to increase in the western
United States, with future patterns and
trends of wildland fire dependent on
several factors (e.g., degree of warming
and drought conditions, fuel and soil
moisture, wildland fire management
practices, elevation) and geographic
region (Service 2018, p. 100). However,
given the diversity of habitats occupied
by wolverines, their occupancy of high
elevations, extensive mobility, and the
positive effect wildland fire may have
on wolverine prey species, wildland fire
represents a limited stressor, in scope
and scale, to wolverine habitat and its
prey in the contiguous United States
range (Service 2018, pp. 63–64).
To summarize, based on the best
available information, the cumulative
effects of wildland fire and climate
change (e.g., snowpack) will continue to
represent a low impact to the wolverine
and its habitat into the mid-21st
century, based on climate change
projections (Service 2018, p. 100).
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Other Cumulative Effects
Finally, we note here that the effects
of climate change on snowpack are
projected to negatively affect the season
lengths for winter recreational activities,
such as skiing and snowmobiling,
shortening the winter recreation season
(Service 2018, pp. 100–101). A shorter
winter recreation season would likely
decrease the amount of winter
recreation related disturbance occurring
in wolverine habitat and fewer effects to
wolverines. Alternatively, even though
winter recreation seasons will be
shorter, we could see more winter
human activity at higher elevations due
to snow loss at lower elevations.
However, even at current levels, we do
not consider winter recreational
activities to be a threat to wolverine in
the contiguous United States. For
further discussion of winter recreational
activities see the wolverine SSA report
(Service 2018, pp. 100–101).
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Summary of Future Conditions
Climate change model projections for
the range of the wolverine within the
contiguous United States indicate
increases in temperature by the mid21st century as compared to early to
mid-20th century values (Service 2018,
p. 101). The degree of future warming
varies by region; area specific
discussions are included in the SSA
report (Service 2018, pp. 73–80).
Precipitation patterns into the future are
less clear as the climate models show
significant disagreement in their many
regional projections. Although drought
conditions in the western United States
are not unusual, drought duration and
intensity have the potential to be
exacerbated by projected temperature
increases. Projected temperature and
precipitation changes will affect future
snow cover and the persistence of snow
on the landscape.
Snow cover is projected to decline in
response to warming temperatures and
changing precipitation patterns, but this
varies by elevation, topography, and by
geographic region (Service 2018, p. 101).
Simulations of natural snow
accumulation at winter recreation
locations have found that, overall,
higher elevation areas (e.g., Rocky
Mountains, Sierra Nevada Mountains)
are more resilient to projected changes
in temperature and precipitation as
compared to lower elevations (Wobus et
al. 2017, p. 12). In general, models
indicate higher elevations will retain
more snow cover than lower elevations,
particularly in early spring (April 30/
May 1) (Service 2018, p. 101). In the
wolverine SSA report, we present
results from several recent climate
models projecting snowpack declines in
the western United States (Service 2018,
pp. 83–100). More specifically, we
reviewed a new analysis (Ray et al.
2017, entire) that modeled future snow
persistence for Glacier and Rocky
Mountain National Parks (areas that
encompass the latitudinal and
elevational range of the wolverine in the
contiguous United States) at high spatial
resolution and at the den-site scale
(Service 2018, pp. 85–98). The results
indicate large areas (several hundred
km2/mi2 for each site) of future snow
(greater than 0.5 m (20 in) in depth) will
persist on May 1 (end of the denning
season) at elevations currently used by
wolverines for denning (Service 2018,
pp. 93–98). This is true, on average,
across the range of climate models used
out to approximately year 2055.
Within their North American range,
wolverines are found in a variety of
habitats within primarily high-elevation
areas of the western-northwestern
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United States, and exhibit wide-ranging
movements (Service 2018, p. 102).
Wolverines select den sites for differing
characteristics depending on location,
and natal den locations are generally
associated with snow cover; however,
many natal dens have been observed
outside of the circumpolar boundary of
the snow model presented in Copeland
et al. (2010) (Service 2018, p. 103),
particularly in Scandinavia. In addition,
reproductive success of wolverines has
not been evaluated relative to the depth
and persistence of snow cover at the den
site scale, or in combination with these
or other important key life-history
characteristics, including avoidance
and/or protection from predators, prey
availability, availability of food-caching
habitat.
We also considered temperature and
precipitation projections from climate
change models in conjunction with
wildland fire risk. This risk is likely to
increase across the western United
States, but patterns and trends are
dependent on several factors (e.g.,
degree of warming and drought
conditions, fuel and soil moisture) and
geographic region (Service 2018, p. 102)
and wildland fire represents a limited
stressor, in scope and scale, to
wolverine habitat and prey as described
above in Effects from Wildland Fire.
Overall Assessment
The wolverine’s current potential
extent of occurrence includes the
western-northwestern United States (see
Figure 2), large areas of Canada, and
Alaska (Service 2018, p. 16). The
wolverine is found in a variety of
habitats in North America, but generally
occurs in high-elevation, relatively
inaccessible locations (Service 2018, p.
102). In the contiguous United States,
potentially suitable habitat (i.e., primary
habitat), as determined by the physical
and ecological features and the
ecological needs of the wolverine, is
estimated at 164,125 km2 (63,369 mi2)
(Inman et al. 2013, p. 281). Based on our
review of available relevant literature,
we identified the physical and
ecological needs of the species as
follows: large territories in relatively
inaccessible landscapes, at high
elevation (1,800 to 3,500 m (5,906 to
11,483 ft)) within the contiguous United
States; access to a variety of food
resources, that varies with seasons; and
reproductive behavior linked to both
temporal and physical features (Service
2018, p. 104). These needs are currently
met for wolverines in the contiguous
United States and are expected to be
met in the future (i.e., in 38–50 years)
(Service 2018, p. 104).
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We recognize there is limited
information available for the wolverine,
including population estimates and
abundance trends. In the contiguous
United States, the structure of the
wolverine population is represented as
a metapopulation, although its genetic
structure relative to its entire North
American range has not been
comprehensively evaluated (Service
2018, p. 102). Wolverine populations in
Alaska are considered to be continuous
with populations in the Yukon and
British Columbia provinces of Canada
based on genetic studies (COSEWIC
2014, p. 37). Similarly, studies of
wolverines in the North Cascades region
have documented recent movement of
wolverines from Washington into
British Columbia (Aubry et al. 2016, pp.
16, 20) and from Idaho (Lucid et al.
2016, p. 184) to British Columbia, and
earlier from Montana to British
Columbia and Alberta (e.g., Newby and
Wright 1955, p. 252).
We present in our SSA report a
detailed discussion of wolverine
reproductive behavior. Based on the
best available information, wolverines
select den sites for different
characteristics depending on location.
Dens located under snow cover may be
related to wolverine distribution based
on other life-history traits, including
morphological, demographic, and
behavioral adaptations that allow them
to successfully compete for food
resources (Inman 2013, pers. comm.).
Structure (e.g., uprooted trees, boulders
and talus fields) appears to be an
important requirement for natal den
sites. However, reproductive success of
wolverines has not been evaluated
relative to the depth and persistence of
snow cover, or in combination with
these or other important characteristics,
including prey availability and predator
avoidance. Recent studies of wolverine
populations and distribution in Sweden
have observed wolverine populations
and reproductive den sites outside areas
modeled with persistent spring snow
cover (Aronsson and Persson 2016;
Persson 2017, pers. comm.). Another
recent study concluded that wolverines
are adaptable and do not require large
areas of deep spring snowpack for
successful reproduction, and may select
small areas covered with deep snow at
a finer scale than can be detected using
satellite imagery (Webb et al. 2016, p.
1,468). Most recently, wolverine dens
have been documented in boreal
Alberta, Canada, several kilometers
away from spring snow cover, in hollow
mounds caused by fallen spruce trees
(Jokinen et al, 2019). We would not
expect fallen trees, and the potentially
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suitable denning sites created by them,
to be a limiting resource in wolverine
habitat.
We identified several potential
stressors that may be affecting the
species and its habitat currently or in
the future, including impacts associated
with climate change effects. Based on
the best available information,
demographic risks to the species from
either known or most likely potential
stressors (i.e., disturbance due to winter
recreational activities, other human
disturbances, effects of wildland fire,
disease, predation, overutilization,
genetic diversity, small-population
effects, climate change, and cumulative
effects) are low based on our evaluation
of the best available information as it
applies to current and potential future
conditions for the wolverine and in the
context of the attributes that affect the
needs of the species (Service 2018, p.
103).
Climate change model projections for
the range of the wolverine within the
contiguous United States indicate
increases in temperature by the mid21st century as compared to early to
mid-20th century values (Service 2018,
p. 103). Our evaluation of climate
change indicates that snow cover is
projected to decline in response to
warming temperatures and changing
precipitation patterns, but this varies by
elevation, topography, and by
geographic region (Service 2018, p. 103).
In general, models indicate higher
elevations will retain more snow cover
than lower elevations, particularly in
early spring (April 30/May 1) (Service
2018, p. 103). Although the persistence
of spring snow has not yet been
determined to be critical to wolverine
survival in North America, our review
of projected snow persistence (to
approximately 2055) within the
Northern and Southern Rocky
Mountains, indicates several hundred
km2/mi2 of deep snow will persist on
May 1 at elevations used by the
wolverine for denning (Service 2018, p.
103).
Legal protections of the wolverine in
the contiguous United States include
State listing in California and Oregon (as
threatened); Colorado (as endangered);
candidate species status in Washington;
protected as a non-game species in
Idaho and Wyoming; a species of
concern and furbearer with a closed
season in Montana; and protected from
collection, importation, and possession
in Utah (Service 2018, p. 107). Trapping
or hunting of wolverines is currently
prohibited in the contiguous United
States. Trapping effort along the U.S.–
Canada border does not represent a
barrier to wolverine movement and
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dispersal along the international border
(Service 2018, p. 103).
Management actions for conservation
of the wolverine and its habitat are
included within State Wildlife Action
Plans, the Management Plan for the
Conservation of Wolverines in Idaho
(IDFG 2014), and USDA Forest Service
Land and Resource Management Plans
(Service 2018, Appendix G). Various
provisions of these plans include, but
are not limited to, winter road closures,
fire management, and land acquisition
or conservation easements. These
management measures, currently and in
the future, will alleviate effects
associated with potential impacts
related to stressors. However, we do not
rely on the management measures and
conservation efforts contained in these
plans to support our listing decision. In
addition, the WAFWA Wildlife Chiefs
Wolverine Subcommittee is providing a
forum for western States to work
collaboratively with each other and with
the Service and other partners for
conserving wolverines found in the
western-northwestern United States,
and, to date, approximately $1.5 million
of funding has been applied towards
conservation and management actions
for the wolverine (e.g., Western States
Wolverine Conservation Project)
(McDonald 2017, pers. comm.).
Determination of Species Status
Section 4 of the Act (16 U.S.C. 1533)
and its implementing regulations (50
CFR part 424) set forth the procedures
for determining whether a species meets
the definition of ‘‘endangered species’’
or ‘‘threatened species.’’ The Act defines
an ‘‘endangered species’’ as a species
that is ‘‘in danger of extinction
throughout all or a significant portion of
its range,’’ and a ‘‘threatened species’’ as
a species that is ‘‘likely to become an
endangered species within the
foreseeable future throughout all or a
significant portion of its range.’’ The Act
requires that we determine whether a
species meets the definition of
‘‘endangered species’’ or ‘‘threatened
species’’ because of any of the following
factors:
(A) The present or threatened
destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial,
recreational, scientific, or educational
purposes;
(C) Disease or predation;
(D) The inadequacy of existing
regulatory mechanisms; or
(E) Other natural or manmade factors
affecting its continued existence.
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Determination of Status Throughout All
of Its Range in the Contiguous United
States
Since the publication of the February
4, 2013, proposed rule (78 FR 7864) and
reinstatement of that proposed rule on
October 18, 2016 (81 FR 71670), we
prepared a comprehensive assessment
of the current and future status of
wolverines in the contiguous United
States as presented in the wolverine
SSA report (Service 2018, entire). New
information from recent surveys and a
reevaluation of the species’ current
range, new genetic information, new
studies of wolverine reproductive
behavior and denning habitat, and
results from detailed modeling of future
spring snow persistence are included in
the wolverine SSA report and contribute
to our current understanding of the
species. The wolverine SSA report also
provides a comprehensive summary of
wolverine life history and ecology,
including an assessment of wolverine
physiology, and an analysis of new
information on wolverine trapping
pressure in Canada near the United
States-Canada border, as well as
analyses of new information relevant to
other potential threats to the species.
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to North American
wolverines in the contiguous United
States including effects from roads
(Factors A and E); disturbance due to
winter recreational activity (Factors A
and E); other human disturbance
(Factors A and E); effects from wildland
fire (Factor A); disease (Factor C);
predation (Factor C); overutilization
(trapping) (Factor B); genetic diversity
(Factor E); small-population effects
(Factor E); and climate change (Factors
A and E). We also assessed the adequacy
of existing regulatory mechanisms
(Factor D).
Consideration of Cumulative Effects—
Threats can work in concert with one
another to cumulatively create
conditions that may impact the
wolverine or its habitat beyond the
scope of each individual threat. See the
Climate Change and Potential for
Cumulative Effects section above and
the wolverine SSA report for an indepth analysis of cumulative effects
(Service 2018, pp. 82–101). We note that
by using the SSA framework to guide
our analysis of the scientific information
documented in the SSA report, we have
not only analyzed individual effects on
the species, but we have also analyzed
their potential cumulative effects. We
incorporate the cumulative effects into
our SSA analysis when we characterize
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the current and future condition of the
species. Our assessment of the current
and future conditions encompasses and
incorporates the threats individually
and cumulatively. Because the SSA
framework considers not just the
presence of the factors, but to what
degree they collectively influence risk to
the entire species, our assessment
integrates the cumulative effects of the
factors.
Our future-condition analysis in the
wolverine SSA report includes the
potential conditions that the species or
its habitat may face, that is, the most
probable scenario if those conditions are
realized in the future. This most
probable scenario includes
consideration of the sources that have
the potential to most likely impact the
species at the population or rangewide
scales in the future, including potential
cumulative impacts. Given an expected
increase in temperature in the western
United States, the best available
information indicates that, if there are
any cumulative impacts in the future,
the most likely to have population-level
effects on wolverine in the contiguous
United States could be: (1) Changes in
snowpack from the combination of
increased temperature and changes in
precipitation patterns, or (2) changes in
snowpack and increase in wildland fire
potential (Service 2018, p. 83). The best
available information does not indicate
that the effects of trapping and mortality
from roads will act cumulatively with
effects of climate change, and those
stressors are expected to remain lowlevel impacts into the future. We
provide a detailed analysis of climate
change and the potential for cumulative
effects in the wolverine SSA report
(Service 2018, pp. 82–102). Based on the
best available information, the
cumulative effects of wildland fire and
climate change (e.g., snowpack) will
continue to represent a low impact to
the wolverine and its habitat into the
mid-21st century, based on climate
change projections.
Resilience, Representation, and
Redundancy—In order to characterize a
species’ viability and demographic
risks, we consider the concepts of
resilience, representation, and
redundancy. We also consider known
and potential stressors that may
negatively impact the physical and
biological features that the species
needs for survival and reproduction.
Stressors are expressed as risks to its
demographic features such as
abundance, population and spatial
structure, and genetic or ecological
diversity. We consider the level of
impact a stressor may have on a species
along with the consideration of
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demographic factors (e.g., whether a
species has stable, increasing, or
decreasing trends in abundance,
population growth rates, diversity of
populations, and loss or degradation of
habitat).
Wolverine populations in much of
North America are still recovering from
large losses of individuals from
unregulated hunting and persecution
pressures in the late 1880s into the mid20th century (Service 2018, p. 104).
Surveys conducted in the winter of
2015–2016 and 2016–2017 continue to
document its presence within portions
of its historical range in the western
contiguous United States
(representation) (Service 2018, p. 104).
Redundancy, the ability to rebound
after stochastic perturbation, can be
characterized by the distribution and
connectivity of populations. In
considering wolverine in the contiguous
United States, individuals are found in
alpine, boreal, and subalpine habitats,
with breeding populations in four
western States. Additionally, wolverines
in the contiguous United States are
connected to wolverine populations in
Canada along the U.S.–Canada border,
which contributes to current and future
redundancy (Service 2018, p. 104).
Resiliency, the ability to withstand
stochastic events, can be characterized
by numbers of individuals and
abundance trends. As indicated above,
actual current population size, growth
rate, and current population trends are
unknown for wolverines in the
contiguous United States due to the lack
of abundance information. However,
according to recent estimates, Canada’s
western subpopulation (which is
connected to wolverines in the
contiguous United States) has been
estimated at 15,688 to 23,830 adult
wolverines, with expansion of
wolverines into historically occupied
areas in both Canada and the contiguous
United States with movement across
both international borders (Service
2018, pp. 54, 105). The 2014 Committee
on the Status of Endangered Wildlife in
Canada report concluded that a climatedriven decline in wolverine populations
in North America is not evident at this
time in much of its range (COSEWIC
2014, p. 22). Wolverine populations in
Canada are considered stable (Service
2018, p. 105). We also note that density
estimates indicate no declining trend in
wolverine populations in Alaska
(Service 2018, p. 105). We recognize
that there is limited information on
populations (representation) or genetic
diversity (resiliency and representation)
for the wolverine in the contiguous
United States, and no comprehensive
studies to indicate what a viable (or
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minimal) wolverine population size
should be across its North American
range. However, the best available
information does not indicate either
increasing or declining numbers of the
wolverine in North America, including
the contiguous United States. Further, at
this time, the best available information
does not indicate that the species’
abundance is significantly impacted by
the stressors evaluated (singly or
cumulatively), and this situation is
unlikely to change in the future,
supporting current and future
resiliency.
As discussed in the wolverine SSA
report, both direct and cumulative
effects of climate change (e.g., higher
temperatures, loss of snow cover,
wildland fire) may affect the resilience
of the wolverine in the future by
creating an environment that is less
favorable to its physiological and
ecological needs (Service 2018, p. 105).
We are unaware of studies of the
wolverine that have formally evaluated
the species’ responses (e.g.,
reproductive success or survival) to
warming temperatures or other climate
change effects.
As described in the wolverine SSA
report, the best available information
indicates confirmed observations of
wolverines denning in areas with
patchy snow cover in Alaska, Canada,
and Scandinavia (Service 2018, p. 105).
Further, using fine-scale snow
modeling, we estimated that large areas
of spring snow (May 1) will remain
within Glacier National Park, where
wolverines are known to den (Service
2018, p. 105). Given their high rate of
movement, large dispersal distances,
including travel through areas not
covered with snow, and other lifehistory traits (e.g., behavioral plasticity)
observed in wolverines, we do not
predict a significant loss of individual
and population resiliency to the species
in the future (i.e., 38–50 years) within
its North America range, including the
contiguous United States (Service 2018,
p. 105).
Currently, we are unaware of any
documented specific risks for the
wolverine related to a substantial
change or loss of diversity in life-history
traits, population demographics,
morphology, behavior, or genetic
characteristics that can be used to
characterize species representation (the
ability to adapt to change). Rates of
dispersal or gene flow are not known to
have changed, and recent evidence
supports continued connectivity with
contiguous United States wolverines
and wolverines in Canada. Additionally,
there is no currently available
information to indicate that the current
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abundance of the wolverine across its
current potential extent in the
contiguous United States is at a level
that is causing inbreeding depression or
that loss of genetic variation is affecting
representation or that would affect
representation in the future (Service
2018, p. 105). Nor is there any
information to indicate that this species
is unable to adapt or adjust to changing
conditions (e.g., potential reduction in
snow cover). We do not expect a
reduction in representation of the
wolverines in the contiguous United
States in the future. We have
determined that the needs of the species
are provided within the contiguous
United States currently and into the
future. Thus, after assessing the best
available information, we conclude that
the North American wolverine in the
contiguous United States is not in
danger of extinction throughout all of its
range (endangered) nor is it likely to
become so in the foreseeable future
(threatened).
Because we determined that the North
American wolverine in the contiguous
United States is not in danger of
extinction or likely to become so in the
foreseeable future throughout all of its
range, we will consider whether there
are any significant portions of its range
in which it is in danger of extinction or
likely to become so in the foreseeable
future.
Determination of Status Throughout a
Significant Portion of Its Range
Under the Act and our implementing
regulations, a species may warrant
listing if it is in danger of extinction or
likely to become so in the foreseeable
future throughout all or a significant
portion of its range. Having determined
that the wolverine is not in danger of
extinction or likely to become so in the
foreseeable future throughout all of its
range, we now consider whether it may
be in danger of extinction or likely to
become so in the foreseeable future in
a significant portion of its range—that
is, whether there is any portion of the
species’ range for which it is true that
both (1) the portion is significant; and,
(2) the species is in danger of extinction
now or likely to become so in the
foreseeable future in that portion.
Depending on the case, it might be more
efficient for us to address the
‘‘significance’’ question or the ‘‘status’’
question first. We can choose to address
either question first. Regardless of
which question we address first, if we
reach a negative answer with respect to
the first question that we address, we do
not need to evaluate the other question
for that portion of the species’ range.
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64647
In undertaking this analysis for the
North American wolverine in the
contiguous United States, we choose to
address the status question first—we
consider information pertaining to the
geographic distribution of both the
species and the threats that the species
faces to identify any portions of the
range where the species is endangered
or threatened.
For the North American wolverine in
the contiguous United States, we
considered whether the threats are
geographically concentrated in any
portion of the species’ range at a
biologically meaningful scale. We
examined the following threats: effects
from roads, disturbance due to winter
recreational activity, other human
disturbance, wildland fire, disease,
predation, overutilization (trapping),
genetic diversity, small-population
effects, climate change, and cumulative
impacts of these potential threats
(Service 2018, entire). All of these
potential stressors are relatively evenly
distributed geographically throughout
the range of the wolverine in the
contiguous United States. We found no
concentration of threats in any portion
of the wolverine’s range at a biologically
meaningful scale. Therefore, no portion
of the species’ range can provide a basis
for determining that the species is in
danger of extinction now or likely to
become so in the foreseeable future in
a significant portion of its range, and we
find the species is not in danger of
extinction now or likely to become so in
the foreseeable future in any significant
portion of its range. This is consistent
with the courts’ holdings in Desert
Survivors v. Department of the Interior,
No. 16-cv-01165–JCS, 2018 WL 4053447
(N.D. Cal. Aug. 24, 2018), and Center for
Biological Diversity v. Jewell, 248 F.
Supp. 3d, 946, 959 (D. Ariz. 2017).
Determination of Status
We have reviewed the best available
scientific and commercial information
regarding the past, present, and future
threats to the North American wolverine
in the contiguous United States and we
have determined that, if it were to be a
listable entity, it does not meet the
definition of an endangered species or a
threatened species in accordance with
sections 3(6) and 3(20) of the Act.
Furthermore, we have determined that
the population of wolverines in the
contiguous United States is not a DPS.
As a consequence of these
determinations, we are withdrawing our
proposed rule to list the distinct
population segment of the North
American wolverine occurring in the
contiguous United States as a threatened
species.
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References Cited
A complete list of references cited in
this document and the wolverine SSA
report are available on the internet at
https://www.regulations.gov at Docket
No. FWS–R6–ES–2016–0106 and upon
request from the Montana Ecological
Services Office (see FOR FURTHER
INFORMATION CONTACT).
Authors
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The primary authors of this document
are the staff members of the Montana
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Ecological Services Office and the
Mountain-Prairie Regional Office.
document on August 10, 2020, for
publication.
Signing Authority
Authority
The Director, U.S. Fish and Wildlife
Service, approved this document and
authorized the undersigned to sign and
submit the document to the Office of the
Federal Register for publication
electronically as an official document of
the U.S. Fish and Wildlife Service.
Aurelia Skipwith, Director, U.S. Fish
and Wildlife Service, approved this
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
PO 00000
Madonna Baucum,
Regulations and Policy Chief, Division of
Policy, Economics, Risk Management, and
Analytics of the Joint Administrative
Operations, U.S. Fish and Wildlife Service.
[FR Doc. 2020–19538 Filed 10–9–20; 8:45 am]
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Agencies
[Federal Register Volume 85, Number 198 (Tuesday, October 13, 2020)]
[Proposed Rules]
[Pages 64618-64648]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2020-19538]
[[Page 64617]]
Vol. 85
Tuesday,
No. 198
October 13, 2020
Part II
Department of the Interior
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Fish and Wildlife Service
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50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Withdrawal of the
Proposed Rule for the North American Wolverine; Proposed Rule
Federal Register / Vol. 85 , No. 198 / Tuesday, October 13, 2020 /
Proposed Rules
[[Page 64618]]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R6-ES-2016-0106; FF09E21000 FXES11110900000 201]
RIN 1018-BB78
Endangered and Threatened Wildlife and Plants; Withdrawal of the
Proposed Rule for the North American Wolverine
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Proposed rule, withdrawal.
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SUMMARY: We, the U.S. Fish and Wildlife Service, withdraw the proposed
rule to list the distinct population segment (DPS) of the North
American wolverine (Gulo gulo luscus) occurring in the contiguous
United States as a threatened species under the Endangered Species Act
of 1973, as amended (Act). This withdrawal is based on our conclusion
that the factors affecting the species as identified in the proposed
rule are not as significant as believed at the time of the proposed
rule. We base this conclusion on our analysis of current and future
threat factors. We also find that North American wolverines occurring
in the contiguous United States do not qualify as a DPS. Therefore, we
are withdrawing our proposal to list the wolverine within the
contiguous United States as a threatened species.
DATES: The proposed rule that published February 4, 2013 (78 FR 7864),
to list the distinct population segment of the North American wolverine
occurring in the contiguous United States as a threatened species is
withdrawn on October 13, 2020.
ADDRESSES: The withdrawal of our proposed rule, comments, and
supplementary documents are available on the internet at https://www.regulations.gov at Docket Nos. FWS-R6-ES-2012-0107 and FWS-R6-ES-
2016-0106.
FOR FURTHER INFORMATION CONTACT: Jodi Bush, Field Supervisor, U.S. Fish
and Wildlife Service, Montana Ecological Services Office, (see
ADDRESSES). Persons who use a telecommunications device for the deaf
may call the Federal Relay Service at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Executive Summary
Why we need to publish a rule. Under the Act, if we determine that
a species may be an endangered or threatened species throughout all or
a significant portion of its range, we are required to promptly publish
a proposal in the Federal Register and make a determination on our
proposal within 1 year. To the maximum extent prudent and determinable,
we must designate critical habitat for any species that we determine to
be an endangered or threatened species under the Act. Listing a species
as an endangered or threatened species and designation of critical
habitat can only be completed by issuing a rule.
What this document does. We withdraw the proposed rule to list the
DPS of the North American wolverine occurring in the contiguous United
States as a threatened species under the Act.
The basis for our action. Under the Act, we may determine that a
species is an endangered or threatened species because of any of five
factors: (A) The present or threatened destruction, modification, or
curtailment of its habitat or range; (B) overutilization for
commercial, recreational, scientific, or educational purposes; (C)
disease or predation; (D) the inadequacy of existing regulatory
mechanisms; or (E) other natural or manmade factors affecting its
continued existence. We have determined that the factors affecting the
species as identified in the proposed rule (loss of habitat due to
climate change) are not as significant as believed at the time of the
proposed rule. We also find that North American wolverines occurring in
the contiguous United States do not qualify as a DPS.
Peer review. In accordance with our July 1, 1994, peer review
policy (59 FR 34270; July 1, 1994), the Service's August 22, 2016,
Director's Memo on the Peer Review Process, and the Office of
Management and Budget's December 16, 2004, Final Information Quality
Bulletin for Peer Review (revised June 2012), we sought the expert
opinions of four appropriate specialists regarding the species status
assessment report. We received responses from four specialists, which
informed this proposed rule. The purpose of peer review is to ensure
that our listing determinations are based on scientifically sound data,
assumptions, and analyses. The peer reviewers have expertise in the
biology, habitat, and threats to the species. Results of this
structured peer review process can be found at https://www.fws.gov/mountain-prairie/science/peerReview.php. A draft analysis was also
submitted to our Federal, State, and Tribal partners for scientific
review. In preparing this withdrawal, we incorporated the results of
these reviews in the final SSA report, as appropriate.
During the reopening of the public comment periods for the proposed
listing rule, we requested any new information and announced that we
initiated a new and comprehensive status review of the North American
wolverine to determine whether the species meets the definition of an
endangered or threatened species under the Act, or whether the species
is not warranted for listing. The wolverine SSA report provides the
scientific basis for the decision to withdraw the proposed listing rule
for the DPS of wolverine occurring in the contiguous United States.
Both new and updated information and analyses presented in the
wolverine SSA report, summarized below in support of our listing
determination, along with public comment, have also prompted us to
reevaluate our previous assessment of the DPS with respect to wolverine
in the contiguous United States.
Supporting Documents
A team prepared a Species Status Assessment (SSA) for the North
American Wolverine (Gulo gulo luscus) (Service, 2018) (hereafter
referred to as the wolverine SSA report). The SSA team was composed of
U.S. Fish and Wildlife Service biologists, who consulted with other
species experts. The wolverine SSA report represents a compilation of
the best scientific and commercial data available concerning the status
of the species, including the impacts of past, present, and future
factors (both negative and beneficial) affecting the wolverine. The
wolverine SSA report underwent independent peer review by scientists
with experience with mesocarnivores and their conservation and
management, genetics, population modeling, and climate change. The
wolverine SSA report and other materials relating to this proposal can
be found on the Mountain-Prairie Region website at https://www.fws.gov/mountain-prairie/ and at https://www.regulations.gov under Docket No.
FWS-R6-ES-2016-0106.
Previous Federal Actions
Please refer to the proposed listing rule for the wolverine (78 FR
7864; February 4, 2013) for a detailed description of previous Federal
actions concerning the wolverine prior to 2013. On February 4, 2013, we
published a proposed rule to list the DPS of wolverine occurring in the
contiguous United States as threatened, under the Act, with a proposed
rule under section 4(d) of the Act that outlines the prohibitions
necessary and advisable for the conservation of the wolverine (78 FR
[[Page 64619]]
7864). We also published a proposed rule on February 4, 2013, to
establish a nonessential experimental population (NEP) area for the
North American wolverine in the Southern Rocky Mountains of Colorado,
northern New Mexico, and southern Wyoming (78 FR 7890). On October 31,
2013, we reopened the comment period on the proposed listing rule for
an additional 30 days (78 FR 65248).
Following publication of the 2013 proposed rules, there was
scientific disagreement and debate about the interpretation of the
habitat requirements for wolverines and the available climate change
information used to determine the extent of threats to the DPS. Based
on this substantial disagreement regarding the sufficiency or accuracy
of the available data relevant to the proposed listing, on February 5,
2014 (79 FR 6874), we announced a 6-month extension of the final
determination of whether to list the wolverine DPS as a threatened
species. We also reopened the comment period on the proposed rule to
list the contiguous United States DPS of the North American wolverine
for 90 days.
On August 13, 2014, we withdrew the proposed rule to list the DPS
of the North American wolverine as a threatened species under the Act
(79 FR 47522). This withdrawal was based on our conclusion that the
factors affecting the DPS as identified in the proposed rule were not
as significant as believed at the time of the proposed rule's
publication in 2013. As a result, we also withdrew our associated
proposed rule under section 4(d) of the Act contained in the proposed
listing rule and withdrew the proposed NEP designation under section
10(j) of the Act for the southern Rocky Mountains.
In October 2014, three complaints were filed in the District Court
for the District of Montana by Defenders of Wildlife, WildEarth
Guardians, Center for Biological Diversity, and other organizations
challenging the withdrawal of the proposal to list the North American
wolverine DPS. Numerous parties intervened in the litigation. These
three cases were consolidated, and on April 4, 2016, the court issued a
decision. The court granted plaintiff's motion for summary judgment
with respect to the Service's determination regarding (1) the threat
posed to the wolverine by the effects of climate change at the
reproductive denning scale, (2) the threat posed to the wolverine by
small population size and lack of genetic diversity, and (3) the
application of the significant portion of its range policy to the
wolverine. As a result of the court order, the August 13, 2014,
withdrawal (79 FR 47522) was vacated and remanded to the Service for
further consideration consistent with the order. As documented in the
SSA report, the Service conducted additional analyses and reviewed new
literature regarding climate change effects at the denning scale (see
pages 73-99 of the SSA report) and included additional life-history
information relevant to this potential stressor (see pages 25-39). With
regard to population size, we also provide in the SSA report an
analysis of information on wolverine population abundance and
distribution (to date) and have included a discussion of population
structure (genetics, effective population size) in the context of the
species' known genetic variability (see pages 44-50). Finally, in this
withdrawal, we have provided an updated significant portion of its
range analysis (see discussion below).
In effect, the court's action returned the process to the proposed
rule stage, and the status of the wolverine under the Act reverted to
that of a proposed species for the purposes of consultation under
section 7 of the Act. On October 18, 2016, we published a notice (81 FR
71670) reopening the comment period on the February 4, 2013, proposed
rule (78 FR 7864) to list the DPS of wolverine occurring in the
contiguous United States as threatened, under the Act. We also
requested new information and announced that we initiated a new and
comprehensive status review of the North American wolverine, to
determine whether the species meets the definition of an endangered or
threatened species under the Act, or whether the species is not
warranted for listing. The wolverine SSA report provides the scientific
basis for the decision to withdraw the proposed listing rule for the
DPS of wolverine occurring in the contiguous United States. Both new
and updated information and analyses presented in the wolverine SSA
report, summarized below in support of our listing determination, along
with public comment, have prompted us to reevaluate our previous
assessment of the DPS (presented in our 2013 proposed listing rule,
which in turn relied on the DPS analysis completed in our 2010 12-month
finding) with respect to wolverine in the contiguous United States.
Summary of Comments and Recommendations
As stated above in the Previous Federal Actions section, on October
18, 2016 (81 FR 71670), we opened a public comment period on our
February 4, 2013, proposed rule (78 FR 7864) to list the DPS of
wolverine occurring in the contiguous United States as threatened. We
also contacted appropriate Federal and State agencies, scientific
experts and organizations, Tribes, and other interested parties and
invited them to comment on the proposed rule. Many of the comments we
received from State agencies during our notice for reopening the
comment period (81 FR 71670) were similar to those received for the
previously proposed rule (78 FR 7864). All substantive information
provided during comment periods has either been incorporated directly
into this final determination or is addressed below.
Public Comments
(1) Comment: We received several public comments claiming that the
North American wolverine faces increasing threats from the effects of
climate change, particularly habitat loss due to declining snow pack.
Our Response: As discussed in the wolverine SSA report, we
recognize that current climate trends and future (2055 and later)
climate model projections indicate warming temperatures for much of
western North America and changes to snow pack conditions. Our review
of the literature found that, overall, higher elevation areas (e.g.,
Rocky Mountains, Sierra Nevada Mountains) are more resilient to
projected changes in temperature and precipitation as compared to lower
elevations (Wobus et al. 2017, p. 12). In general, models indicate
higher elevations, where documented historical wolverine denning has
occurred, will retain more snow cover than lower elevations,
particularly in early spring (April 30/May 1). We present in the
wolverine SSA report a summary of new, fine-scale analysis of future
snow persistence in two regions of the western United States, Glacier
National Park and Rocky Mountain National Park. The two regions studied
include a high-latitude area near tree line within Glacier National
Park, where tree line occurs at (~1,800 to 2,100 meters (m) (5,906 to
6,890 feet (ft))) that is currently occupied by wolverines; and a lower
latitude area within Rocky Mountain National Park (occupied by a single
male wolverine from 2009 to at least 2012, but not known to be
currently occupied), where tree line occurs at higher elevation (~3,500
m (11,483 ft)) (Ray et al. 2017, p. 2). These sites were chosen to
bracket the range of latitude and elevation wolverines currently occupy
in the contiguous United States (Ray et al. 2017, p. 2). This effort
built upon previous model projections presented in McKelvey et al.
(2011), but with significant differences such as finer
[[Page 64620]]
spatial resolution, incorporation of slope and aspect, snow depth
estimates, additional years of historical data, and wider temporal
analyses of snow persistence (April-June). Details of this modeling
exercise are presented in Ray et al. (2017), and summarized in the SSA
report. That analysis indicates significant areas (several hundred
square kilometers (km\2\)/square miles (mi\2\) for each study area) of
future snow (greater than 0.5 m (20 inches (in) in depth) will persist
on May 1 at elevations currently used by wolverines for denning. This
is true, on average, across the range of climate models used out to
approximately year 2055.
(2) Comment: We received several public comments during our request
for information claiming that low population size (and small effective
population size) warrant listing of the North American wolverine as
threatened or endangered.
Our Response: Small populations in and of themselves do not
constitute a threat such that a species would be endangered or
threatened. When evaluating species status, we take into consideration
the species' life history, population dynamics, and other impacts to
populations and species to determine if small population dynamics
increases the species' vulnerability to extinction such that listing as
threatened or endangered is warranted. Wolverines are difficult animals
to survey, and populations occur in naturally low densities across
their North American range due, in large part, to their need for large,
exclusive territories. At the present time, there is no reliable
estimate of the number of wolverines that currently occupy or
previously occupied the contiguous United States, nor are there
reliable quantitative estimates of wolverine population trends in the
contiguous United States. The often-cited population estimate of 318
wolverines (range: 249-626) in the contiguous United States is derived
from a habitat modeling exercise presented in Inman et al. (2013). That
publication also provided a model estimate of potential wolverine
capacity of 644 (range: 249-626). However, both of these estimates did
not consider important spatial considerations related to wolverine
behavior, such as territoriality, relative to wolverine populations.
Despite the paucity of information regarding wolverine populations, the
SSA analysis is a thorough examination of all of the available
population information.
As discussed in the wolverine SSA report, preliminary field results
from a recent (2016-2017) occupancy study in four western States
(Idaho, Montana, Washington, and Wyoming) and from a pilot occupancy
study in Wyoming (2015-2016) indicate detections of wolverines in areas
where they would be expected to be found, but also no detections in
areas where they are known to occur (e.g., areas within Glacier
National Park) (see Service 2018, Appendix B for a descriptive map). To
date, this study reports a total of 86 photographic detections through
camera-trapping and 157 wolverine hair samples collected for genetic
analysis. It has not yet been determined from the camera-trap images or
hair samples how many of the detections are unique individuals.
Preliminary analysis of the study results indicates an average
estimated probability of occupancy of 0.42 suggesting that wolverines
used nearly half of all sites during the study period (Montana FWP,
pers. comm., 2017); however, the study did not encompass all potential
wolverine habitat in the western United States (Service 2018, Appendix
B). For example, wolverines have also been recently detected in
northeastern Oregon (as of 2017) and in parts of Grand Teton National
Park (two records during the winter of 2017), which were not included
in the surveyed study cells. Our SSA report presents a visual summary
of these recent detections (Service 2018, Figure 3). Although the sum
of these reports cannot confirm previous estimates of population size
or verify population trends, they offer recent evidence that wolverines
continue to be observed across a large area of the western United
States.
The 2013 proposed rule presented an effective population size
estimate from a publication by Schwartz et al. (2009), which estimated
a summed effective population size of 35, with credible limits from 28
to 52 (Schwartz et al. 2009, p. 3,226). As described in the SSA report,
the study used wolverine samples from the main part of the Rocky
Mountain wolverine populations and did not include subpopulations from
two other mountain regions in Montana, and samples were missing from
other parts of the wolverine range in Idaho and other areas of Montana.
Thus, the analysis missed both wolverine subpopulations and
individuals, which would underestimate the results for this type of
analysis.
In the wolverine SSA report, we provide a contextual discussion of
the effective population size concept, particularly in the context of
genetic studies related to the phylogeographic history of the North
American wolverine that were not well described in the 2013 proposed
rule. In sum, the spatial distribution of genetic variability currently
observed in wolverines in North America appears to be a reflection of a
complex history in which population abundance has fluctuated since the
time of the last glaciation with insufficient time passing since human
persecution since at least the 1700s to allow for full recovery of
wolverine densities (Cardinal 2004, pp. 23-24; Zigouris et al. 2012, p.
1,554). This history and the fact that wolverines in the contiguous
United States occupy the southern periphery of the species' entire
North American range are important considerations in estimating and
interpreting current wolverine distribution and abundance. The
wolverine SSA report also presents information from genetic and
observational studies that provide support for wolverine movement
across the international border of the contiguous United States and
Canada. In the 2013 proposed rule, we stated there is an apparent lack
of connectivity between wolverine populations in Canada and the United
States based on genetic data (78 FR 7864; February 4, 2013). We now
consider wolverines that occupy the contiguous United States to be
genetically continuous with wolverines in adjacent Canadian provinces.
A small effective population size would be more of a concern if the
population was in isolation; however, wolverines in the contiguous
United States are not genetically isolated from wolverines in Canada.
For more information, see the Small Total Population Size and Effective
Population Size sections under Distinct Population Segment below.
(3) Comment: We received several public comments during our request
for information claiming that North American wolverine face threats
from indiscriminant trapping in the contiguous United States, or are
threatened by incidental trapping.
Our Response: At the present time, trapping or hunting of wolverine
is not allowed in any State within the range of the wolverine (with the
exception of Alaska). Legal protections for wolverines are codified in
western State laws and regulations and include: Endangered in Colorado,
threatened in California and Oregon, candidate species in Washington,
non-game species protections in Idaho and Wyoming, a species of concern
and a furbearer with a closed season in Montana, and protection from
collection, importation, and possession in Utah. Since 2013, there has
been a zero quota for trapping or harvest of wolverine in Montana.
Incidental trapping of wolverines has been documented in the
contiguous
[[Page 64621]]
United States (as recently as December 2017), though not all events
have resulted in mortality. In the wolverine SSA report, we provide a
summary of the number of wolverines that have been incidentally trapped
in Idaho (18 since 1965, including 6 known to be released alive and 7
known mortalities), Montana (4 since 2013, 3 mortalities and 1 released
unharmed), and Wyoming (2 since 1996, 1 mortality and 1 released
unharmed) (Service 2018, p. 66). Both Idaho and Montana are
implementing trapper education programs to minimize nontarget wolverine
captures.
As discussed in the SSA report, regulated trapping and hunting of
wolverines occurs in parts of Alaska and Canada, and appears to be
sustainable based on population and density estimates.
(4) Comment: We received several public comments identifying
potential threats to wolverines from winter recreation activities, such
as snowmobiling and back-country skiing.
Our Response: In the SSA report, we present a summary of winter
recreation studies (Heinemeyer et al. 2015; Heinemeyer et al. 2017),
future projections of winter recreation activity in the contiguous
United States (White et al. 2014), and projections of snowpack relative
to changes in the length of the winter recreation season (Wobus et al.
2017). We reported results from Heinemeyer (2016, pers. comm.)
indicating a behavioral response to recreation activities, but also
maintenance of home ranges within some areas of relatively high
recreation activity over several years. The study has not yet been able
to determine whether resident wolverines are reproductively successful
due to the limited monitoring information available for reproducing
female wolverines. Nor was the study able to determine if recreational
activities had a negative impact on wolverine reproductive success.
We also note here that we received the final report of this
multiyear study (Heinemeyer et al. 2017) in mid-December 2017 (results
of this study were recently published (Heinemeyer et al. 2019)), which
was after we submitted the draft SSA report for review to four peer
reviewers and to our State, Federal, and Tribal partners. Much of the
report presents a modeling exercise to evaluate wolverine behavior
patterns with winter recreation activities. The study found that
wolverines maintained multiyear home ranges, and the authors suggest
that wolverines are able to tolerate winter recreation at some scales
(Heinemeyer et al. 2017, p. iv; Heinemeyer et al. 2019, p. 16). The
study described habitat selection as complex for female wolverines and
was likely driven by a combination of abiotic (snow, cold) and biotic
(predator avoidance, food availability) factors (Heinemeyer et al.,
2017, p. 36; Heinemeyer et al. 2019, p. 16). This study did not assess
demographic effects, fitness effects, or population level effects of
winter recreation on wolverines (Heinemeyer et al. 2019, p. 17 and 19).
As discussed in the wolverine SSA report, management measures being
implemented in areas within the wolverine's current extent of
occurrence include road closures to minimize disturbance to wildlife on
lands managed by the U.S. Forest Service and National Park Service
(Service 2018, p. 61 and Appendix F). In addition, management
strategies are identified in State Wildlife Action Plans (e.g., Oregon,
Montana, Idaho) to address potential impacts from recreation to the
wolverine. Although we did not rely on these conservation measures to
support our decision, they do provide some level of protection to
address potential impacts from disturbance from winter recreation
activity and mortality from roads.
(5) Comment: We received public comments claiming that wolverines
are dependent on deep snow for survival and expressing concern for
future changes in snow pack due to the effects of climate change.
Our Response: After reviewing studies not previously considered and
the results of new studies/publications made available after the 2013
and 2014 proposed rules (e.g., Aronsson 2017, Aronsson and Persson
2016, Aronsson et al. 2017, Magoun et al. 2017, Persson et al. 2017,
Stewart et al. 2016, Webb et al. 2016, see complete list of citations
in the wolverine SSA report), we present in the SSA report a detailed
discussion of the North American wolverine's physiology and other life-
history characteristics (e.g., reproductive behavior). This summary
speaks to several presumed aspects of the relationship of denning
behavior and other needs of this species regarding the presence of
persistent spring snow. As summarized below, we now know that
wolverines can and have denned outside of heavy snowpack, multiple
factors play a role in den site selection, females will move dens as
young become mobile, and areas of significant snowpack will likely
persist in the future in areas where wolverines are known to den at
levels that will continue to support wolverines.
Denning habitat for the wolverine varies over its range and is
dependent on local and regional environmental conditions (e.g.,
topographic and other structural features) and biotic (e.g.,
availability of prey; protection from predators) factors. Reproductive
(natal) dens are not always excavated in deep snow, particularly in
boreal forest habitats (Dawson et al. 2010; Novikov 1962; Webb et al.
2016; Jokinen 2019.), and have been observed in spruce tree root balls,
logging slash piles, and beaver dens/dams. In the contiguous United
States, dens are found at high elevations, often in talus slopes, which
provides conditions for protection and food caching (e.g., restricted
access and cold temperatures). Our review of studies of wolverine
denning activity found no quantitative data reporting snow depth at the
den site when wolverines abandon the den. More importantly, wolverine
reproductive success has not been studied relative to a number of
abiotic and biotic conditions, including depth and temporal aspect of
spring snow cover.
Wolverines begin shifting den locations in late April, when young
become more mobile and reliant on solid food brought to them by the
mother (Aronsson 2017, p. 46; Aubry et al. 2016, p. 24). The
bioclimatic model presented in Copeland et al. (2010) was used to test
wolverine distribution at a broad scale based on climate variables,
including spring snow cover, using May 15 conditions. They then tested
their hypothesis by comparing and correlating the location of wolverine
dens across their circumboreal range, and telemetry locations from
wolverine studies in North America and Scandinavia (Copeland et al.
2010, p. 234). Since that publication, wolverines and wolverine dens
have been observed outside the boundaries defined by the model
presented in Copeland et al. (2010) (e.g., Webb et al. 2016, Webb 2017
pers. comm., Persson 2017, pers. comm.). While these observations are
found at higher latitudes in the circumboreal region, they also
indicate wolverines and wolverine dens are observed in environments
that are not characterized by several feet of spring snow on May 15. In
sum, Copeland et al. (2010) provided a fairly accurate assessment of
where wolverine populations are expected to occur, but it did not
evaluate (model) snow persistence at the den site scale based on
location and denning period.
In the SSA report, we present an analysis of 34 wolverine den
locations (years 2002-2015) from studies in the western contiguous
United States relative to ``melt out'' dates, which represents the
first day of an 8-day satellite (Moderate Resolution Imaging
[[Page 64622]]
Spectroradiometer (MODIS)) composite of when the den switches from
``snow'' to ``no snow'' at a 500-by-500-m (1,640-by-1,640-ft) spatial
resolution. For natal den locations, the range of the melt-out dates
was from May 25 to June 12, which is considerably later than the May 15
date used in the Copeland et al. (2010) analysis. The estimated melt-
out dates indicate that snow is persistent at these locations past the
time when young wolverines are generally moving out of natal dens
(i.e., late April).
The Copeland et al. (2010) snow model was then used by McKelvey et
al. (2011) to model effects of climate change to wolverine habitat in
the western United States to develop projections of habitat loss. This
modeling exercise used May 1 snow presence as a proxy for May 15 snow
disappearance and a spatial resolution of 36.3 km\2\ (14 mi\2\)), which
is not relevant at the at the den site scale. As described in our
Response to Comment #1, in the SSA report, we presented a finer scale
analysis (0.0625 km\2\ (0.24 mi\2\)) for two study areas (Glacier
National Park and Rocky Mountain National Park) that focused directly
on May 15, in addition to the presence or absence of snow on May 1 and
April 15 in our evaluation of the effects of climate change to
snowpack. These dates are more relevant to wolverine life-history
needs. We also modeled the depth of ``significant'' snow (0.5 m (1.64
ft)) on these dates. We found that large areas (several hundred km\2\/
mi\2\ for each study area) of future snow (greater than 0.5 m (20 in)
in depth) are projected to persist on May 1 at elevations currently
used by wolverines for denning. This is true, on average, across the
range of climate models used out to approximately year 2055. We
recognize that wolverines are difficult to study and evaluation of
denning habitat and behavior is challenging. Additional research is
needed to evaluate other potential physical and biotic variables that
could be important in defining wolverine distribution and den
locations. These additional variables include: prey availability, risk
of predation, den-site scale factors such as structure/snow conditions,
and temporal use of dens.
(6) Comment: We received public comments identifying the need for
additional research and recommendations for conservation measures for
the North American wolverine, including estimates of population size
and further evaluation of life-history characteristics, and
recommendations for conservation measures.
Our Response: We appreciate the comments acknowledging the need for
additional studies as well as the difficulties in studying wolverines
given its occupation of remote habitats in the contiguous United
States. In the wolverine SSA report, we provide a summary of the
preliminary field and genetic results from the recent Western States
Wolverine Conservation Project (WSWCP)-Coordinated Occupancy Study in
four western (contiguous United) States, as well as results from
several new studies presented in peer-reviewed publications and in
other reports from Canada and Scandinavia. As discussed in the SSA
report, the Western States Wolverine Working Group is continuing to
develop studies to evaluate wolverine population distribution and
occupancy, and connectivity across four western States.
(7) Comment: We received additional comments from the public
including the need for collaboration with local government and
community stakeholders and use of best available science in developing
the proposed rule.
Our Response: During our preparation of the wolverine SSA report,
we coordinated extensively with many wolverine researchers in the
United States (including Alaska), Canada, and Scandinavia. Those
communications are identified in our References Cited section of the
wolverine SSA report. Their expertise, insights, and published or soon-
to-be published research papers were invaluable in ensuring that we
used the best available science in preparing the new status review. We
also communicated with biologists at several State and Federal agencies
to ensure that we had incorporated the most recent wolverine detections
in the western United States. The wolverine SSA report was sent to four
independent peer reviewers, selected by an outside contractor, and
those non-attributable comments were incorporated, to the extent
possible, in the final document. We also reviewed comments received
from the public and previous peer reviewers during our request for
comments for our previous proposed rule and considered the information
provided (78 FR 7864; February 4, 2013) during the preparation of the
wolverine SSA report. As a result, this determination is based upon the
best scientific and commercial data available to us, as required by the
Act.
(8) Comment: We also received public comments recommending that the
North American wolverine not be listed as threatened or endangered
under the Act. One commenter stated that State wildlife agencies are
capable of managing the species and are able to provide protections
that ensure continued population growth towards population objectives
established by these agencies and that mandates of various Federal
resource management agencies provide a commitment to managing wildlife
habitat in a way that benefits all wildlife species, including
wolverines and other forest carnivores.
Our Response: We acknowledge that some members of the public
support our decision to withdraw our proposed rule to list the North
American wolverine as threatened under the Act. In the wolverine SSA
report (Service 2018, Appendix G), we provide a summary of the
regulatory protections provided by western States and Federal agencies
as well as management measures being implemented to conserve the
wolverine and its habitat. Legal protections in the contiguous United
States include State listing in California and Oregon (threatened),
endangered in Colorado, a candidate species in Washington, non-game
species protections in Idaho and Wyoming, a species of concern and
furbearer with a closed season in Montana, and protection from
collection, importation, and possession in Utah. Trapping or hunting of
wolverines is currently prohibited in the contiguous United States.
(9) Comment: In response to our request for information in our
public notice, several members of the public provided specific
information related to personal wolverine sightings of the North
American wolverine in the contiguous United States (e.g., New Mexico,
Wyoming), and information regarding past and ongoing research studies
of the species in the western United States and in Canada.
Our Response: We appreciate the personal observations provided and
encourage members of the public to document sightings of the North
American wolverine with photographs and provide additional details to
State wildlife agencies. Information we received regarding results from
research studies has been incorporated, as appropriate, in the final
wolverine SSA report.
(10) Comment: We received comments from several organizations that
support the listing of the North American wolverine and designation of
critical habitat. Threats cited include concerns related to migration,
habitat loss and connectivity related to threats from effects of
climate change, nontarget trapping pressures, road mortality and other
effects of roads (e.g., noise, pollution, fragmentation of habitat),
motorized recreation and traffic in wildlife corridors, timber sales
and
[[Page 64623]]
associated roads, and effects of snowmobile traffic (habitat
fragmentation and pollution, and change in behavior).
Our Response: As discussed in the Risk Factors for the North
American Wolverine section below, we identified several potential
stressors that may be affecting the species and its habitat currently
or in the future, including impacts associated with climate change
effects. We recognize there is limited information available for the
wolverine, including population estimates and abundance trends. Based
on the best available information, demographic risks to the species
from either known or most likely potential stressors (i.e., disturbance
due to winter recreational activities, other human disturbances,
effects of wildland fire, disease, predation, overutilization, genetic
diversity, small population effects, climate change, and cumulative
effects) are low based on our evaluation of the best available
information as it applies to current and potential future conditions
for the wolverine and in the context of the attributes that affect the
needs of the species (Service 2018, p. 103). Thus, we determined that,
based on the best available information, the North American wolverine
in the contiguous United States does not meet the definition of a
threatened species or an endangered species under the Act.
(11) Comment: We received public comments stating that protection
of North American wolverines in the contiguous United States is needed
under the Act in order to provide resources and attention needed for
research and monitoring, to better understand threats, and sustain
wolverines into the future. The commenter also stated that federally
sponsored wolverine reintroduction in Colorado will help increase
chances of long-term species survival.
Our Response: We appreciate the recognition of the need for
continued resources for research and monitoring. However, we base our
listing decisions on a determination of whether the species meets the
Act's definitions of a threatened species or an endangered species.
Regardless, as summarized in the SSA report, in 2015, State wildlife
agencies in Idaho, Montana, Washington, and Wyoming, along with
Federal, tribal, and nongovernmental organization partners, developed a
collaborative and coordinated monitoring program to be implemented in a
coordinated fashion across the species' range in the western United
States. In 2015, the State of Wyoming contracted with the Wolverine
Initiative to conduct the Wyoming Wolverine Occupancy Pilot Study to
address questions pertaining to the status and distribution of
wolverines throughout the Greater Yellowstone Ecosystem and the Bighorn
Mountains in the winter of 2015-2016. Expanding on this study, the
Western States Wolverine Working Group designed and implemented the
WSWCP-Coordinated Occupancy Survey in the winter of 2016-2017, and
preliminary results are presented in the SSA report. The Western
Association of Fish and Wildlife Agencies (WAFWA) Wildlife Chiefs
Wolverine Subcommittee (formally endorsed in 2014) currently provides a
forum for western States to work collaboratively with each other and
with the Service, Tribes, and other partners, for conserving wolverines
across the western United States. To date, approximately $1.5 million
of that funding has been applied towards conservation and management
actions, including the WSWCP (McDonald 2017, pers. comm.). This group
is also developing a connectivity study project to support conservation
planning efforts for the Rocky Mountains and North Cascades regions.
In addition, multiple western States have identified the North
American wolverine as a Species of Greatest Conservation Need in their
State Wildlife Action Plans, and the North American wolverine is a
focal species of conservation strategies for conservation targets in a
number of ecoregions (e.g., Cascades, Sierra Nevada) that support
forested lowlands, subalpine-high montane conifer forest where
wolverines occur. These State designations provide information to
assist resource managers with proactive decision making regarding
species conservation and data collection priorities. Finally, the Nez
Perce Tribe is currently preparing an Integrated Resource Management
Plan, a Plant and Wildlife Conservation Strategy, and a Forest
Management plan with the wolverine defined as a species of conservation
concern in all three draft plans (Miles 2017, pers. comm.).
In total, these funded and volunteer collaborative, landscape-level
conservation efforts ensure continued support for the conservation of
the North American wolverine. Although we did not rely on these plans
to support our decision, we recognize that these plans, when
implemented, will likely benefit wolverines and their habitat.
(12) Comment: We received comments from several industry groups
supporting our August 13, 2014, withdrawal (79 FR 47522) of our
February 4, 2013, proposed rule (78 FR 7864) to list the North American
wolverine as threatened. In general, their support rests on the
following: (1) The DPS determination presented in our previous proposed
rules (both 2010 and 2013) was flawed; (2) the North American wolverine
does not meet the definition of a threatened species; (3) the obligate
relationship with denning and need for snow has not been adequately
addressed (and may be a habitat preference); and (4) climate model
projections do not support complete loss of snow. They also urged us to
reaffirm prior findings that winter recreation (motorized and
nonmotorized) is not a threat to wolverines.
Our Response: Given that our updated analysis and new information
included in the wolverine SSA report directly relates to our previous
DPS determination in the 2013 proposed rule, we reevaluated wolverines
in the contiguous United States under our DPS Policy. See the Distinct
Population Segment section below for more information. We provide our
analysis of the status of wolverines in the contiguous United States
below in the Determination of Species Status. The topic of denning
behavior is discussed in the wolverine SSA report (see Use of Dens and
Denning Behavior discussion in the Reproduction and Growth section in
the wolverine SSA report (Service 2018, pp. 23-28)). For our analysis
of the effects of climate change to wolverines and denning habitat, see
Climate Change and Potential for Cumulative Effects below.
(13) Comment: We received a comment from an industry group stating
that our decision to prepare the February 4, 2013, proposed rule (78 FR
7864) to list the North American wolverine as threatened was due to a
``misreading'' of the Service's obligation under our 2011 Settlement
Agreement, and therefore the proposed rule was not developed from ``an
open-ended scientific inquiry.'' We received a comment from an industry
group stating that the Service should not ``revert back to the 2013
proposed rule'' and should conduct a new analysis of potential impacts
to the species, revise the proposed listing using newly available
information, and reevaluate our previous DPS determination.
Our Response: The Service properly prepared its 2013 proposed rule.
On October 18, 2016, we published a document in the Federal Register
(81 FR 71670) announcing that we would initiate a new status review of
the North American wolverine, to determine whether this DPS meets the
definition of an endangered or threatened species
[[Page 64624]]
under the Act, or whether the species is not warranted for listing. The
Service has prepared the wolverine SSA report that includes discussion
and analyses of the best available scientific information regarding
life history, biology, and consideration of current and future
vulnerabilities. This information was used to evaluate the current and
future conditions of the species, and to inform our current
determination.
Comments From Tribes
(14) Comment: We received comments from one consortium of Tribal
nations stating that, based on the weight of evidence provided in our
previous rules, the North American wolverine meets the definition of
endangered or threatened and is therefore warranted for listing.
Specific threats mentioned in the comment letter included current
population status, winter recreation activities, and effects of climate
change. The Tribes also included comments documenting the cultural
value of the wolverine and connection to cultural practices and concern
for the loss of wolverine populations in the contiguous United States.
The Tribes encouraged the Service to use sound and solid science in the
listing determination, and noted that additional population monitoring
and Tribal climate change modeling efforts are under way to evaluate
the status of the wolverine.
Our Response: We appreciate the unique perspective provided by the
Tribal nations regarding the contribution of the North American
wolverine to the Tribes' culture and spirituality. We also appreciate
the commitment of the Tribal nations to continue ongoing studies of
wolverines.
As described in the wolverine SSA report, we evaluated new
information, as well as information not previously considered, and
contacted several wolverine researchers (both within and outside the
United States) to provide a more detailed description of the
wolverine's life history and ecology, including a detailed discussion
of wolverine denning habitat and behavior. We conducted new analyses to
develop a current potential extent of occupancy using the most recent
verified observations. Current potential extent is the perimeter of the
outermost geographic limits based on all (available) occurrence records
(that is, the maximum extent of occurrences) of a species minus those
areas where we believe the species has been extirpated (Service 2017).
Conservation measures and regulatory mechanisms relative to the
wolverine were also provided in the wolverine SSA report. This
information was used to evaluate the current (potential stressors) and
future conditions of the species, and inform our current determination.
We evaluated results from a fine-scale analysis of the potential
effects of climate change to future snowpack conditions in two regions
of the Rocky Mountains. This analysis found that significant areas
(several hundred km\2\/mi\2\) will persist on May 1 at elevations used
by wolverines for denning. We determined that, based on the best
available information, the North American wolverine in the contiguous
United States does not warrant listing as threatened or endangered
under the Act.
(15) Comment: We received comments from one Tribe whose aboriginal
territory is occupied by the North American wolverine. The Tribe
submitted a comment letter in 2013 supporting our proposed listing. The
Tribe stated that the conservation and restoration of the wolverine and
other species within this homeland is of great importance to the
Tribe's subsistence, culture, religion, and economy. The letter also
identified conservation and management plans currently under
development and highlighted that the wolverine is designated as a
species of concern in these current draft plans. Specific comments were
provided relative to threats from climate change (including relative to
demographic stochasticity), recreation and urban development, and
incidental take. Included in those comments were references to other
studies under way (e.g., Adaptation Partners and climate change
vulnerability assessments; winter recreation study) to evaluate these
potential stressors.
Our Response: We appreciate the perspective provided regarding the
importance of the wolverine and other species to the Tribe and its
commitment to current and future conservation and management actions.
We also appreciate and evaluated the information presented in the
citations that were provided in the comment letter. As described in the
wolverine SSA report, we evaluated several new scientific publications
and information not previously considered in preparing a new status
review. This information was used to evaluate the current conditions
(i.e., potential stressors, including winter recreation) and future
conditions (e.g., effects of climate change) of the species. Based on
the best available information, we determined that the North American
wolverine in the contiguous United States does not warrant listing as
threatened or endangered under the Act.
State Agency Comments
We received extensive comments from several western States,
requesting that we consider previously submitted comments in response
to our previously proposed listing rule (78 FR 7864; February 4, 2013)
as well as additional comments submitted in response to our 2016 notice
reopening public comment (81 FR 71670; October 18, 2016). These
comments were grouped together and summarized as described below:
(16) Comment: We received detailed comments critical of our
reliance on ``unverified'' climate model projections in our 2013
proposed rule, the lack of discussion of assumptions in adopting the
model findings, the lack of evaluating alternative hypotheses, and the
need to evaluate these effects at the den-site scale. One State agency
recommended that, given the disagreements in the scientific community
on the interpretation of these results, the Service solicit an
independent, scientific review of the proposed rule.
Our Response: This withdrawal was based on the scientific analysis
using the structure of the Service's Species Status Assessment (SSA)
Framework (https://www.fws.gov/endangered/improving_ESA/ssa.html). An
SSA is a focused and rigorous assessment of a species' ability to
maintain self-sustaining populations over time. This assessment is
based on the best available scientific and commercial information
regarding life history, biology, and consideration of current and
future vulnerabilities. The result is a single document (SSA report)
that delivers foundational science for informing decisions under the
Act, including listing determinations, consultations, grant
allocations, permitting, and recovery planning.
In preparing the final SSA report for the North American wolverine
(available at www.regulations.gov, at Docket No. FWS-R6-ES-2016-0106),
we reviewed available reports and peer-reviewed literature,
incorporated survey information for the purpose of preparing updated
maps of the known species' current and historical occurrences, and
contacted species experts to collect additional unpublished
information. We evaluated the appropriate analytical tools to address
data gaps and uncertainties. In some instances, we used publications
and other reports of the Eurasian subspecies (Gulo gulo gulo) to fully
inform our knowledge of the North American wolverine (Gulo gulo
luscus).
Before finalizing the SSA report, the draft wolverine SSA report
was
[[Page 64625]]
submitted for peer review to four independent peer reviewers in
accordance with our July 1, 1994, peer review policy (59 FR 34270), the
Service's August 22, 2016, Director's Memo on the Peer Review Process,
and the Office of Management and Budget's December 16, 2004, Final
Information Quality Bulletin for Peer Review (revised June 2012).
Results of this structured peer review process can be found at https://www.fws.gov/mountain-prairie/science/peerReview.php. This draft was
also submitted to our Federal, State, and Tribal partners for
scientific review. In preparing this determination to withdraw the
proposed rule, we incorporated the results of these reviews in the
final wolverine SSA report, as appropriate.
As noted in our previous responses to public comments (see response
to Comments 1 and 5 above), in our wolverine SSA report, we recognize
that current climate trends and future (2055 and later) climate model
projections indicate warming temperatures for much of western North
America, and changes to snow pack conditions. Our review of the
literature found that, overall, higher elevation areas (e.g., Rocky
Mountains, Sierra Nevada Mountains) are more resilient to projected
changes in temperature and precipitation as compared to lower
elevations (Wobus et al. 2017, p. 12). In general, models indicate
higher elevations will retain more snow cover than lower elevations,
particularly in early spring (April 30/May 1). We present in the
wolverine SSA report a summary of new, fine-scale analysis of future
snow persistence in two regions of the western United States, Glacier
National Park and Rocky Mountain National Park. Glacier National Park
represents a high-latitude and relatively low-elevation area currently
occupied by North American wolverines. Rocky Mountain National Park is
a lower latitude and high-elevation area within the North American
wolverine's historical range, which was occupied by a male wolverine
from 2009 to at least 2012.
As described above in Comment 5, this new analysis built upon
previous model projections presented in McKelvey et al. (2011), but
with significant differences such as finer spatial resolution,
incorporation of slope and aspect, snow depth estimates, additional
years of historical data, and wider temporal analyses of snow
persistence (April-June). Details of this modeling exercise are
presented in Ray et al. (2017), and summarized in the wolverine SSA
report. That analysis indicates large areas (several hundred km\2\/
mi\2\ for each study area) of future snow (greater than 0.5 m (20 in)
in depth) will persist on May 1 at elevations currently used by
wolverines for denning. This is true, on average, across the range of
climate models used out to approximately year 2055.
After reviewing studies not previously considered and new studies/
publications made available after the 2013 and 2014 proposed rules, we
present in the wolverine SSA report, a detailed discussion of the North
American wolverine's physiology and other life-history characteristics
(e.g., reproductive behavior). The analysis speaks to several presumed
aspects of the relationship of denning behavior and other needs of this
species regarding the presence of persistent spring snow.
Also, see our response to Comment 5 above for a short summary and
our SSA report for more details regarding our analysis of the effects
of climate change to denning habitat.
(17) Comment: We received comments critical of our previous support
for findings by Schwartz et al. 2009 regarding effective population
size. Relatedly, several States commented on recent dispersal/movements
of wolverines into California, Colorado, and Utah as evidence of
population expansion.
Our Response: See our response to Comment 2 above for a discussion
of effective population size. Regarding recent occurrences of
wolverines in the contiguous United States, wolverines have recently
been found in areas where they were once extirpated in the contiguous
United States. See the Population Abundance and Density section below
for more information.
(18) Comment: We received comments from several western States
presenting clarifications or updates to incidental trapping events and
trapping regulations.
Our Response: In the wolverine SSA report, we include a summary of
trapping or hunting of wolverines in the contiguous United States. At
the present time, trapping or hunting of wolverines is not allowed in
any western State (with the exception of Alaska, which was not included
in the DPS in our proposed rule). Legal protections for wolverines are
codified in western State laws and regulations concerning hunting and
trapping. These protections include: Endangered in Colorado, threatened
in California and Oregon, candidate species in Washington, non-game
species protections in Idaho and Wyoming, a species of concern and
furbearer with a closed season in Montana, and protection from
collection, importation, and possession in Utah. Since 2013, there has
been a zero quota for trapping or harvest of wolverine in Montana.
Incidental trapping of wolverines has been documented in the
contiguous United States (as recently as December 2017), though not all
events have resulted in mortality (see response to Comment 3 above).
Both Idaho and Montana are implementing trapper education programs to
minimize nontarget wolverine captures.
(19) Comment: Several States provided comments in response to our
2013 proposed rule and to our 2016 reopening of the public comment
period indicating their disagreement with our determination of a DPS
for the contiguous United States. Specifically, some commenters stated
that the criteria of significance should be reevaluated, noting that
the proposed rule did not provide any substantive information to
support our conclusion that the loss of the wolverine in the contiguous
United States would result in a significant gap in the range of the
species; that is, our previous use of the loss of latitudinal range
does not provide a rational basis for concluding that the loss of the
wolverine in the contiguous United States would be significant in
relation to the taxon. Another commenter stated that the wolverine
population in the contiguous United States is connected geographically
and genetically to the Canada/Alaska populations and these northern
populations were likely the source of recolonization during the 20th
century. Further, this commenter stated there is not a difference in
control of exploitation and conservation status between the United
States and Canada.
Another commenter noted that, throughout the 2013 proposed rule,
the Service acknowledged that, historically, the wolverine population
in the contiguous United States was markedly reduced by systematic
predator control programs and unregulated trapping. Yet, as the
commenter pointed out, areas of suitable habitat in the North Cascades,
where trapping has been minimal or nonexistent for decades, and
northern Rockies, were recolonized by animals from Canada, where
relatively liberal trapping is still allowed. Thus, our
characterization in the 2013 proposed rule of ``liberal'' Canadian
regulations as sufficient to ``maintain the robust conservation status
of the Canadian population,'' does not comport with our
characterization that the very limited trapping in the contiguous
United States (Montana only) is insufficient to maintain the rebounding
population designated as a DPS.
[[Page 64626]]
Our Response: In light of the updated analysis and new information
included in the wolverine SSA report, we reevaluated wolverines in the
contiguous United States under our DPS Policy. We conclude that the
population of wolverines in the contiguous United States is not
discrete in relation to the remainder of the species in North America.
As a result, the population of wolverines in the contiguous United
States is not a listable entity under section 3(16) of the Act. See the
Distinct Population Segment section below for more information.
(20) Comment: State agencies provided citations or copies of
publications and reports relevant to wolverine ecology that were
published after the 2013 proposed rule.
Our Response: We appreciate the comprehensive lists of published
literature and survey reports provided by the State agencies. We
evaluated this information during the preparation of the wolverine SSA
report, and have incorporated this information, as appropriate, to
ensure that the wolverine SSA report presents the best available
information regarding the status of the North American wolverine.
(21) Comment: We received information providing background
information and preliminary results of ongoing collaborative
conservation efforts being implemented through the WSWCP-Coordinated
Occupancy Survey.
Our Response: We appreciate the additional information provided by
the State agencies participating in the WSWCP-Coordinated Occupancy
Study. In the wolverine SSA report, we provide a summary of the
preliminary field and genetic results from the recent WSWCP-Coordinated
Occupancy Study in four western (contiguous United) States (see
wolverine SSA report for additional details). We also incorporated
technical comments received from several State agencies during the
review of the draft wolverine SSA report into the final report. As
discussed in the wolverine SSA report, the Western States Wolverine
Working Group is continuing to develop studies to evaluate wolverine
population distribution and occupancy, and connectivity across four
western States.
(22) Comment: Information was provided by State agencies describing
the legal protections of wolverines in individual States and
conservation measures being implemented.
Our Response: In the wolverine SSA report (Service 2018, Appendix
G), we provide a detailed discussion of current State (and Federal)
regulatory mechanisms and other conservation measures that offer
protections for the North American wolverine. In addition to the WSWCP-
Coordinated Occupancy Study (Service 2018, Appendix B), several western
States have identified the North American wolverine as a Species of
Greatest Conservation Need in their State Wildlife Action Plans, and
the North American wolverine is a focal species of conservation
strategies for conservation targets in a number of ecoregions (e.g.,
Cascades, Sierra Nevada) that support forested lowlands, subalpine-
high-montane conifer forest where wolverines occur. These State
designations provide information to assist resource managers with
proactive decision-making regarding species conservation and data
collection priorities, and support the conservation of the North
American wolverine and its habitat.
(23) Comment: We received a comment from one State agency noting
that the State does not recognize the North American wolverine as a
native species due to lack of evidence that a population ever existed
within New Mexico (i.e., unverified species); thus, the State does not
recognize the species in any of its wildlife statutes or regulations.
Our Response: We appreciate the clarification and information
provided by the State agency and have considered this in our analysis
to define the current potential extent of occurrence for the North
American wolverine in the contiguous United States (see Figures 1 and 2
below) and in our assessment of population status in the wolverine SSA
report. In their analysis of wolverine distribution records in the
contiguous United States, Aubry et al. (2001, p. 2,150) identified 1860
as the most recent verifiable documentation of wolverine in northern
New Mexico. We received two unverified accounts of wolverine sightings
in New Mexico from the general public during the most recent public
comment period. We are unaware of any recent verifiable individuals or
populations of wolverines in New Mexico.
(24) Comment: In response to our request for information in our
October 18, 2016, Federal Register document (81 FR 71670), we received
comments from the U.S. Forest Service submitting verifiable and new
records of wolverines from 2000 to 2016. These records include
observations from camera surveys by both governmental and
nongovernmental organizations, photos from private citizens, and
locations from a regional study.
Our Response: We appreciate the information provided and
incorporated these observations and detections in our analysis to
define the current potential extent of occurrence for the North
American wolverine in the contiguous United States (see Figures 1 and 2
below) and in our assessment of population status in the wolverine SSA
report.
Background
A comprehensive review of the life history, population trends, and
ecology of the North American wolverine is presented in the wolverine
SSA report (Service 2018, pp. 3-44). The Service recognizes the North
American wolverine as the subspecies Gulo gulo luscus (Service 2018, p.
8). Wolverines are a medium-sized (about 1 m (3.3 ft) in length)
carnivore, with a large head, broad forehead, and short neck (Service
2018, p. 4). Wolverines have heavy musculature and relatively short
legs, and large feet with strong, curved claws for digging and climbing
(Service 2018, p. 4). Their feet are adapted for travel through deep
snow and, during the winter, dense, stiff, bristle-type hairs are found
between the toes and around the foot pad; this characteristic becomes
diminished in the summer (Service 2018, p. 4). The wolverine is the
largest terrestrial member of the Mustelidae family (weasels, fisher,
mink, marten, and others) and resembles a small bear with a bushy tail
(Service 2018, p. 1). Wolverines possess a number of morphological and
physiological adaptations that allow them to travel long distances and
they maintain large territories in remote areas (Service 2018, p. 1).
They have been described as curious, intelligent, and playful, but
cautious animals, though their social behavior and social organization
has not been well-studied (Service 2018, p. 1). Wolverines have a
distribution that includes the northern portions of Europe, Asia, and
North America. In North America, they are found in Alaska, much of
Canada, and the western-northwestern United States.
During the late 1800s and early 1900s, the wolverine population
declined or was extirpated in much of the contiguous United States
(lower 48 States), which has been largely attributed to unregulated
trapping (Hash 1987, p. 583). Wolverine numbers have recovered to some
extent from this decline and, in the United States, wolverines are
currently found in parts of Washington, Oregon, Idaho, Montana,
Wyoming, California (single male), and Alaska, and as recently as 2010
in Michigan, 2012 in Colorado, and 2016 in Utah. Known reproducing
wolverine populations are found in Washington, Idaho, Montana, and
Wyoming (Service 2018, p. 1).
[[Page 64627]]
Regulatory and Analytical Framework
Regulatory Framework
Section 4 of the Act (16 U.S.C. 1533) and its implementing
regulations (50 CFR part 424) set forth the procedures for determining
whether a species is an ``endangered species'' or a ``threatened
species.'' The Act defines an endangered species as a species that is
``in danger of extinction throughout all or a significant portion of
its range,'' and a threatened species as a species that is ``likely to
become an endangered species within the foreseeable future throughout
all or a significant portion of its range.'' The Act requires that we
determine whether any species is an ``endangered species'' or a
``threatened species'' because of any of the following factors:
(A) The present or threatened destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial, recreational, scientific, or
educational purposes;
(C) Disease or predation;
(D) The inadequacy of existing regulatory mechanisms; or
(E) Other natural or manmade factors affecting its continued
existence.
These factors represent broad categories of natural or human-caused
actions or conditions that could have an effect on a species' continued
existence. In evaluating these actions and conditions, we look for
those that may have a negative effect on individuals of the species, as
well as other actions or conditions that may ameliorate any negative
effects or may have positive effects.
We use the term ``threat'' to refer in general to actions or
conditions that are known to or are reasonably likely to negatively
affect individuals of a species. The term ``threat'' includes actions
or conditions that have a direct impact on individuals (direct
impacts), as well as those that affect individuals through alteration
of their habitat or required resources (stressors). The term ``threat''
may encompass--either together or separately--the source of the action
or condition or the action or condition itself.
However, the mere identification of any threat(s) does not
necessarily mean that the species meets the statutory definition of an
``endangered species'' or a ``threatened species.'' In determining
whether a species meets either definition, we must evaluate all
identified threats by considering the expected response by the species,
and the effects of the threats--in light of those actions and
conditions that will ameliorate the threats--on an individual,
population, and species level. We evaluate each threat and its expected
effects on the species, then analyze the cumulative effect of all of
the threats on the species as a whole. We also consider the cumulative
effect of the threats in light of those actions and conditions that
will have positive effects on the species, such as any existing
regulatory mechanisms or conservation efforts. The Secretary determines
whether the species meets the definition of an ``endangered species''
or a ``threatened species'' only after conducting this cumulative
analysis and describing the expected effect on the species now and in
the foreseeable future.
The Act does not define the term ``foreseeable future,'' which
appears in the statutory definition of ``threatened species.'' Our
implementing regulations at 50 CFR 424.11(d) set forth a framework for
evaluating the foreseeable future on a case-by-case basis. The term
``foreseeable future'' extends only so far into the future as the
Services can reasonably determine that both the future threats and the
species' responses to those threats are likely. In other words, the
foreseeable future is the period of time in which we can make reliable
predictions. ``Reliable'' does not mean ``certain''; it means
sufficient to provide a reasonable degree of confidence in the
prediction. Thus, a prediction is reliable if it is reasonable to
depend on it when making decisions.
It is not always possible or necessary to define foreseeable future
as a particular number of years. Analysis of the foreseeable future
uses the best scientific and commercial data available and should
consider the timeframes applicable to the relevant threats and to the
species' likely responses to those threats in view of its life-history
characteristics. Data that are typically relevant to assessing the
species' biological response include species-specific factors such as
lifespan, reproductive rates or productivity, certain behaviors, and
other demographic factors.
Analytical Framework
The SSA report documents the results of our comprehensive
biological status review for the species, including an assessment of
the potential threats to the species. The SSA report does not represent
a decision by the Service on whether the species should be proposed for
listing as an endangered or threatened species under the Act. It does,
however, provide the scientific basis that informs our regulatory
decisions, which involve the further application of standards within
the Act and its implementing regulations and policies. The following
sections provide summaries of the key results and conclusions from the
SSA report; the full SSA report can be found on the Mountain-Prairie
Region website at https://www.fws.gov/mountain-prairie/ and at https://www.regulations.gov under Docket No. FWS-R6-ES-2016-0106.
To assess wolverine viability, we used the three conservation
biology principles of resiliency, redundancy, and representation
(Shaffer and Stein 2000, pp. 306-310). Briefly, resiliency supports the
ability of the species to withstand environmental and demographic
stochasticity (for example, wet or dry, warm or cold years), redundancy
supports the ability of the species to withstand catastrophic events
(for example, droughts, large pollution events), and representation
supports the ability of the species to adapt over time to long-term
changes in the environment (for example, climate changes). In general,
the more resilient and redundant a species is and the more
representation it has, the more likely it is to sustain populations
over time, even under changing environmental conditions. Using these
principles, we identified the species' ecological requirements for
survival and reproduction at the individual, population, and species
levels, and described the beneficial and risk factors influencing the
species' viability.
The SSA process can be categorized into three sequential stages.
During the first stage, we evaluated the individual species' life-
history needs. The next stage involved an assessment of the historical
and current condition of the species' demographics and habitat
characteristics, including an explanation of how the species arrived at
its current condition. The final stage of the SSA involved making
predictions about the species' responses to positive and negative
environmental and anthropogenic influences. This process used the best
available information to characterize viability as the ability of a
species to sustain populations in the wild over time. We use this
information to inform our regulatory decision.
Distinct Population Segment
Pursuant to the Act, we must consider for listing any species,
subspecies, or, for vertebrates, any distinct population segment (DPS)
of these taxa, if there is sufficient information to indicate that such
action may be warranted. To interpret and implement the DPS provision
of the Act and Congressional guidance, the Service and the National
Marine Fisheries Service published, on February 7, 1996, an interagency
Policy
[[Page 64628]]
Regarding the Recognition of Distinct Vertebrate Population Segments
under the Act (61 FR 4722; February 7, 1996). This policy addresses the
recognition of DPSs for potential listing actions. The policy allows
for more refined application of the Act that better reflects the
biological needs of the taxon being considered, and avoids the
inclusion of entities that do not require its protective measures.
Under our DPS policy, three elements are considered in a decision
regarding the status of a possible DPS as endangered or threatened
under the Act. These are applied similarly for additions to the list of
endangered and threatened species, reclassification, and removal from
the list. They are: (1) Discreteness of the population segment in
relation to the remainder of the taxon; (2) the biological or
ecological significance of the population segment to the taxon to which
it belongs; and (3) the population segment's conservation status in
relation to the Act's standards for listing (i.e., whether the
population segment is, when treated as if it were a species or
subspecies, an endangered or threatened species). Discreteness refers
to the degree of isolation of a population from other members of the
species, and we evaluate this factor based on specific criteria. If a
population segment is considered discrete, we must consider whether the
discrete segment is ``significant'' to the taxon to which it belongs by
using the best available scientific and commercial information. When
determining if a potential DPS is significant, our policy directs us to
sparingly list DPSs while encouraging the conservation of genetic
diversity. If we determine that a population segment is both discrete
and significant, we then evaluate it for endangered or threatened
species status based on the Act's standards.
Both new and updated information and analyses presented in the
wolverine SSA report, summarized below in support of our listing
determination, along with public comment, have prompted us to
reevaluate our previous assessment of the DPS (presented in our 2013
proposed listing rule, which in turn relied on the DPS analysis
completed in our 2010 12-month finding) with respect to wolverine in
the contiguous United States. Below we provide our revised evaluation
of discreteness under the DPS policy of the segment of the North
American wolverine occurring in the contiguous United States.
Distinct Population Segment Analysis for Wolverine in the Contiguous
United States
Analysis of Discreteness
Under our DPS Policy, a population segment of a vertebrate species
may be considered discrete if it satisfies either one of the following
conditions: (1) It is markedly separated from other populations of the
same taxon as a consequence of physical, physiological, ecological, or
behavioral factors (quantitative measures of genetic or morphological
discontinuity may provide evidence of this separation); or (2) it is
delimited by international governmental boundaries within which
differences in control of exploitation, management of habitat,
conservation status, or regulatory mechanisms exist that are
significant in light of section 4(a)(1)(D) of the Act (inadequacy of
existing regulatory mechanisms). Discreteness Based on Marked
Separation
In our February 4, 2013, proposed listing rule (78 FR 7864), we did
not rely on marked separation from other populations to support
discreteness of the contiguous United States wolverine population. As
supported by information in the SSA report, we maintain that there are
no physical, physiological, ecological, or behavioral factors
separating wolverines in the contiguous United States from wolverines
in Canada. We do not consider wolverines in the contiguous United
States to be genetically isolated from wolverines in Canada (McKelvey
et al. 2014; Pilgrim and Schwartz 2018). Therefore, wolverines in the
contiguous United States are not discrete based on marked separation
from other populations of the same taxon.
Discreteness Based on the International Border--Legal Status Conveyed
by National, State, and Provincial Governments; Differences in Control
of Exploitation
Our 2013 proposed rule (78 FR 7864), which incorporated by
reference our DPS analysis from our 2010 12-month finding, found there
was no significant difference between the legal status of wolverines
between Canada and the United States (75 FR 78030; December 14, 2010).
In the wolverine SSA report, we provide an updated assessment of legal
protections and regulatory mechanisms for wolverine in North America
(Service 2018, pp. 70-71, Appendix G). Legal protections in the
contiguous United States include State listing in California and Oregon
(threatened), endangered in Colorado, a candidate species in
Washington, non-game species protections in Idaho and Wyoming, a
species of concern and furbearer with a closed season in Montana, and
protected from collection, importation, and possession in Utah. In
Canada, provincial designations range from endangered to threatened in
eastern provinces, and sensitive/special concern to no ranking in other
provinces (definitions provided by the Committee on the Status of
Endangered Wildlife in Canada, 2014). As was determined in our 2013
proposed listing rule (78 FR 7864), we again find no significant
differences in legal status.
In the 2010 12-month finding (75 FR 78030) and reiterated in our
2013 proposed listing rule (78 FR 7864), we stated that differences in
control of exploitation exist, but favor the contiguous United States
population. Trapping or hunting of wolverines is currently prohibited
in the contiguous United States and regulated as appropriate in Canada
(Service 2018, pp. 68-69). In the wolverine SSA report, we included a
new analysis of trapping in southern Canada and trapping effort along
the U.S.-Canada border, which we found to be limited. Thus, we conclude
that the differences in exploitation are not significant in light of
section 4(a)(1)(D) of the Act (inadequacy of existing regulatory
mechanisms).
Discreteness Based on the International Border--Differences in
Management of Habitat
As we outlined in the proposed 4(d) rule (78 FR 7888) management
activities (e.g., timber harvest, wildland firefighting, prescribed
fire, and silviculture) can modify wolverine habitat, but this
generalist species appears to be little affected by changes to the
vegetative characteristics of its habitat. In addition, most wolverine
habitat occurs at high elevations in rugged terrain that is not
conducive to intensive forms of silviculture and timber harvest.
Habitat management is not a conservation need for wolverine. Therefore,
differences in management of habitat between the United States and
Canada are not significant in light of section 4(a)(1)(D) of the Act.
Discreteness Based on the International Border--Differences in
Conservation Status
In the December 14, 2010, 12-month finding (75 FR 78030), which is
incorporated and discussed in the February 4, 2013, proposed listing
rule (78 FR 7864), we found that the wolverine population in the
contiguous United States met the second DPS discreteness condition
because of
[[Page 64629]]
differences in conservation status as delimited by the U.S.-Canada
international governmental boundary. We found that those differences
were substantial and significant in light of section 4(a)(1)(D) of the
Act. We stated that in the remaining current range in Canada and
Alaska, wolverines exist in well-distributed, interconnected, large
populations. We added that, conversely, wolverine populations in the
remaining United States range appear to be at numbers so low that their
continued existence could be at risk, especially in light of the
threats to the species. In the 2010 finding, we stated that risks come
from three main factors: (1) Small total population size; (2) effective
population size below that needed to maintain genetic diversity and
demographic stability; and (3) the fragmented nature of wolverine
habitat in the contiguous United States that results in smaller,
isolated sky island patches separated by unsuitable habitat. We stated
it was apparent that maintaining wolverines within their native range
in the contiguous United States into the future is likely to require
regulatory mechanisms that are not currently in place. As a result, we
concluded that the contiguous United States population of the wolverine
meets the discreteness criterion in our DPS Policy (61 FR 4722,
February 7, 1996). Consequently, we used the international border
between the United States and Canada to define the northern boundary of
the contiguous United States wolverine DPS in our December 14, 2010,
12-month finding (75 FR 78030) and our February 4, 2013, proposed
listing rule (78 FR 7864). Below we provide a reevaluation of that
determination supported by information presented in the wolverine SSA
report.
Small Total Population Size--Wolverine densities vary across North
America and have been described as naturally low (van Zyll de Jong
1975, p. 434); wolverine populations are naturally uncommon given the
species' large home range, wide-ranging movements, and solitary
characteristics (Service 2018, p. 56). There are many fewer wolverines
in the contiguous United States than there are in Canada and Alaska
(Committee on the Status of Endangered Wildlife in Canada (COSEWIC)
2014, p. 36; Inman et al. 2013, p. 282; Service 2018, p. 71), but this
is more a reflection of the amount of suitable habitat available within
the contiguous United States (both currently and historically) for a
species that needs large exclusive territories, than it is a reflection
of poor conservation status. Wolverines in Canada are considered to
occur as a single large group as they are easily able to move between
areas of suitable habitat and because wolverine habitat is relatively
contiguous (Harrower 2017, pers. comm.). However, wolverines in the
contiguous United States are considered to be a metapopulation
connected with wolverine populations in Canada (Inman et al. 2013, p.
277). Wolverines currently occupy areas in the contiguous United States
where they were once extirpated and continue to repopulate the
contiguous United States after decades of unregulated trapping,
hunting, and poisoning (Service 2018, p. iv). The same holds true for
Canada, where wolverines are being detected in areas once extirpated
(COSEWIC 2014, p. v).
These movement patterns are supported by recent genetic information
that indicates wolverines from Canada have slowly repopulated the
contiguous United States over the past century since the era of
unregulated persecution (Service 2018, pp. 45-50). This point is
discussed in detail below in the Genetic Diversity section of this
withdrawal. We stated in the December 14, 2010, finding that
differences in population sizes between the contiguous United States
and Canada were reflective of a difference in conservation status (75
FR 78030). However, based on new information, we now conclude that the
contiguous United States wolverines represent a peripheral population
at the southern extent of the North American wolverine range. Thus, we
now consider the small population size of wolverines in the United
States to be a natural result of habitat fragmentation and not
reflective of a difference in conservation status (see Habitat
Fragmentation below for more detail). Therefore, any difference in
population size on the contiguous United States side of the
international border is not a significant difference in conservation
status in light of section 4(a)(1)(D) of the Act as it applies to
discreteness.
Effective Population Size--Effective population size
(Ne) is defined as ``the size of an idealized population
that would experience the same amount of genetic drift and inbreeding
as the population of interest (Service 2018, Box 2). In popular terms,
Ne is the number of individuals in a population that
contribute offspring to the next generation'' (Hoffman et al. 2017, p.
507). Effective population size can be interpreted differently
depending on how it's defined and used, and the concept of effective
population size (Ne) (see review by Wang et al. 2016) and,
relatedly, minimum viable population, has been a topic of debate,
particularly the 50/500 rule (population size of 50 for short-term, and
500 for long-term genetic health). Importantly, the concept and
guidelines for genetically effective population size were developed for
a single, isolated population (Laikre et al. 2016, p. 280). The term
``effective population size'' is not a meaningful term unless
additional context is provided relative to which concept of population
size is being evaluated (Ewens 1990, p. 309). Demographic factors are
needed when interpreting actual population size from an effective
population size; thus, there is no justification for a fixed,
genetically derived minimum viable population size value of `500' as
each case is unique and is dependent on such factors as sex ratio,
subpopulations, dispersal, and immigration (Ewens 1990, pp. 311-313).
As noted above, we do not consider the wolverine population in the
contiguous United States to be genetically isolated from wolverines on
the other side of the international border in Canada. In the wolverine
SSA report, we provide a contextual discussion of the effective
population size concept, particularly in the context of genetic studies
related to the phylogeographic history of the North American wolverine
(Service 2018, pp. 45-50), which was not well described in the 2013
proposed rule. In sum, the spatial distribution of genetic variability
currently observed in wolverines in North America appears to be a
reflection of a complex history in which population abundance has
fluctuated since the time of the last glaciation with insufficient time
passing since human persecution, since at least the 1700s, to allow for
full recovery of wolverine densities (Cardinal 2004, pp. 23-24;
Zigouris et al. 2012, p. 1,554). This history and the fact that
wolverines in the contiguous United States occupy the southern
periphery of its entire North American range are important
considerations. The wolverine SSA report also presents information from
genetic and observational studies that provide support for wolverine
movement across the international border of the contiguous United
States and Canada (Aubry et al. 2016, pp. 16, 20; Lucid et al. 2016, p.
184; Service 2018, pp. 9-23). Thus, we consider wolverines that occupy
the contiguous United States to be genetically continuous with
wolverines in adjacent Canadian provinces.
Wolverines travel (disperse) through areas outside high-elevation,
forested habitats. For example, tracked movements of a male wolverine,
M56, from Wyoming into Colorado and its subsequent discovery in North
Dakota, indicate extensive travel outside of
[[Page 64630]]
modeled primary wolverine habitat (i.e., Inman et al. 2013), including
through arid grasslands and shrubland habitats of the Wyoming Basin
ecoregion (Packila et al. 2017, entire). This animal's movement also
supports some level of connectivity (and potential gene flow) between
currently occupied habitat (Wyoming) and unoccupied habitat within the
wolverine's historical range (Colorado) (Packila et al. 2017, p. 404).
Similarly wolverines in the North Cascades region have moved from
Washington and Idaho into British Columbia, and from Montana to British
Columbia and Alberta (Service 2018, p. 45). Based on genetic analyses,
the male wolverine currently occupying an area within the Sierra Nevada
Mountains of California also represents evidence of connectivity
between wolverine populations of the Rocky and Sierra Nevada Mountain
Ranges (Moriarty et al. 2009, p. 154). Within the Southwestern Crown of
the Continent (SWCC) in northwestern Montana, cross-valley movements of
wolverines have been detected, which researchers believe is an
indication of good connectivity in this region (SWCC Working Group
2016, pers. comm.).
A preliminary mitochondrial DNA analysis was prepared for wolverine
samples collected during the winters of 2015-2016 and 2016-2017 as part
of the Western States Wolverine Conservation Project--Coordinated
Occupancy Survey (Pilgrim and Schwartz 2018, entire). All 45 wolverines
identified from samples collected in Idaho, Montana, and Wyoming match
haplotype Wilson-A, which is common throughout the Rocky Mountains,
Alaska, and Canada, while all 5 wolverines identified from samples
collected in Washington match haplotype Wilson-C (Pilgrim and Schwartz
2018, p. 3). Previous analyses of recent or modern (1989-2012) samples
from the Cascades Range in northern Washington and southern British
Columbia, as presented in McKelvey et al. (2014, p. 328), were
characterized as haplotype C, and one historical (defined in this study
as pre-1930) sample as haplotype A (McKelvey et al. 2014, p. 327).
Outside of this region, haplotype C has been found only in Alberta,
Saskatchewan, and Nunavut provinces (McKelvey et al. 2014, p. 330).
Based on mitochondrial DNA, McKelvey et al. (2014, p. 330) concluded
that modern (defined in their study as 1989-2012) wolverine populations
in the contiguous United States are the result of recolonization
(following persecution during a period of unregulated hunting or
trapping and poisoning) from the north. The additional mitochondrial
analysis from samples collected in 2015, 2016, and 2017 provides
further support that all contiguous United States historical (pre-1900)
and recent wolverine populations are likely descendants of immigrants
from Canada.
The 2013 proposed rule presented an effective population size
estimate for wolverines in the contiguous United States from a
publication by Schwartz et al. (2009), which estimated a summed
effective population size of 35, with credible limits from 28 to 52
(Schwartz et al. 2009, p. 3,226). As described in the wolverine SSA
report, the study used wolverine samples from the main part of the
Rocky Mountain wolverine populations and did not include subpopulations
from two other mountain regions in Montana, and samples were missing
from other parts of the wolverine range in Idaho and other areas of
Montana. Thus, the analysis missed wolverine subpopulations and
individuals, which would underestimate the results for this type of
analysis. Furthermore, a small effective population size would be more
of a concern if the population was in isolation; however, wolverines in
the contiguous United States are not genetically or physically isolated
from wolverines in Canada.
To summarize, the currently known spatial distribution of genetic
variability in wolverines in North America appears to be a reflection
of a complex history where population abundance has fluctuated since
the time of the last glaciation and insufficient time has passed since
human persecution for a full recovery of wolverine densities (Cardinal
2004, pp. 23-24; Zigouris et al. 2012, p. 1,554). Zigouris et al.
(2012, p. 1,545) noted that the genetic diversity reported in Cegelski
et al. (2006) and Kyle and Strobeck (2001, 2002) for the southwestern
edge of the North American range represented only part of the diversity
in the northern populations of wolverines. Zigouris et al. (2012, p.
1,545) posit that the irregular distribution of wolverines in the
southwestern periphery and the genetic diversity observed in those
analyses is a result of population bottlenecks that were caused by
range contractions from a panmictic (random mating) northern core
population approximately 150 years ago coinciding with human
persecution. Recent dispersals of wolverines into Colorado (2009),
California (2008), and Utah (2014) provide evidence for connectivity
and the potential for gene flow between Northern Rocky Mountain
populations and areas where wolverines were extirpated. As noted above,
there is also recent evidence of wolverine movement across the
international border. Furthermore, our analysis of trapping levels in
the wolverine SSA report does not support previous assumptions that
trapping in Canada near the border acts as a barrier to wolverine
movement into the contiguous United States (Service 2018, pp. 68-69).
Finally, very few successful migrants are needed per generation to
maintain at least 95 percent of the genetic variation in the next 100
generations (approximately 750 years) in the contiguous United States
(Cegelski et al. 2006, p. 209).
We conclude that this level of migration from the north has already
been occurring following the end of intense persecution of this
species; wolverines are currently observed in previously occupied areas
within the contiguous United States. Given the recent observations of
wolverines moving vast distances over varied terrain and across the
U.S.-Canada border, our recent assessment of the low levels of trapping
mortality in Canada near the border, and further confirmation of Canada
as the source of wolverine genetics present in contiguous United States
wolverines, we believe that wolverines in the contiguous United States
are not separated genetically from the larger population in Canada.
Wolverines in the contiguous United States exhibit genetic and
phenotypic similarities with wolverines in Canada that implies
connectivity with Canada. As such, we conclude that it is not
biologically appropriate to consider the low effective population size
of wolverines on the contiguous United States side of the border as a
difference in conservation status that is significant in light of
section 4(a)(1)(D) of the Act as it applies to discreteness. For
additional information related to wolverine genetic diversity and
effective population size, see Genetic Diversity below and the
wolverine SSA report (Service 2018, pp. 45-50).
Habitat Fragmentation--In our 2010 12-month finding (incorporated
into the 2013 proposed listing rule), we stated that wolverine habitat
in the contiguous United States consists of small, isolated islands of
high-elevation habitat separated from each other by low valleys of
unsuitable habitat. We also described that these `habitat islands' are
represented by areas containing spring snow, citing Copeland et al.
(2010). We concluded that the fragmented nature and distribution of
wolverine habitat in the contiguous United States results in a
population that is highly vulnerable to extirpation because of lack of
[[Page 64631]]
connectivity between subpopulations, and this also makes them more
vulnerable to external threats (75 FR 78030; December 14, 2010).
Our previous analysis of wolverine habitat fragmentation relied
upon the assumption that wolverines are constricted to habitats that
contain deep, persistent spring snow cover and, therefore, are more or
less confined to areas that were defined by the Copeland et al. (2010)
spring snow cover model. However, wolverines are observed in and move
through areas without snow cover (e.g., male wolverines dispersing to
California and Colorado), and female wolverines have successfully
denned in areas outside previously modeled projections of deep,
persistent spring snow cover (e.g., Webb et al. 2016; Persson 2017,
pers. comm.; Jokinen 2018, pers. comm.).
We now conclude that it is not accurate to categorize the occupied
habitat of wolverines in the contiguous United States as `habitat
islands.' As discussed above, wolverine populations in the contiguous
United States represent the southern periphery of a much larger range
of the North American wolverine due to naturally occurring landscape
features such as high elevation and topographic roughness of mountain
regions. Thus, the distribution of persistent spring snow cover in
mountainous regions does not represent the only determining habitat
feature for wolverines. The availability of prey and avoidance of
predators are also important elements of wolverine habitat (Inman et
al. 2012, p. 785; Scrafford et al. 2017, p. 34)). As described in the
SSA report, wolverines use a unique and productive ecological niche
that allows them to occupy high-elevation regions across the
northwestern portion of the contiguous United States (Service 2018, pp.
27, 38). Finally, as noted above, wolverine movement in the contiguous
United States is not constrained by high-elevation habitat or snow
cover, and wolverines can easily move and disperse long distances in
the western United States (e.g., SWCC Working Group 2016, pers. comm.;
Packila et al. 2017, entire). Therefore, habitat fragmentation in the
context of availability of persistent spring snow cover or loss of
connectivity in the contiguous United States is not an appropriate
difference in conservation status in light of section 4(a)(1)(D) of the
Act as it applies to discreteness.
Discreteness Based on the International Border--Differences in
Regulatory Mechanisms
Because there aren't significant differences in control of
exploitation, legal conservation status, and management of habitat, nor
other threats to the wolverine requiring regulatory mechanisms to
address them, we conclude that there are not differences in regulatory
mechanisms between the United States and Canada that are significant in
light of section 4(a)(1)(D).
Conclusion on Discreteness
Based on our updated analysis described above and supported by
information in the wolverine SSA report, the contiguous United States
population of wolverine does not meet the discreteness criterion in our
DPS Policy (61 FR 4722; February 7, 1996). As a result, the contiguous
United States population of wolverines does not qualify as a DPS and is
not a listable entity under section 3(16) of the Act. After determining
that a vertebrate population is not discrete, we are not required to
complete an analysis to determine if the population in question is
significant according to our DPS Policy.
DPS Conclusion
Based on the best available information, we conclude that the
population of wolverines in the contiguous United States is not
discrete in relation to the remainder of the species in North America.
As a result, the population of wolverines in the contiguous United
States is not a listable entity under section 3(16) of the Act.
The DPS Policy sets forth a three-step process for determining
whether a vertebrate population as a separate entity warrants listing:
(1) Determine whether the population is discrete; (2) if the population
is discrete, determine whether the population is significant to the
taxon as a whole; and (3) if the population is both discrete and
significant, then evaluate the conservation status of the population to
determine whether it is endangered or threatened. Although we have
determined that wolverines in the contiguous United States do not
qualify as a DPS and, therefore, are not a listable entity, we provide
below a status determination of the wolverine population in the
contiguous United States. The DPS Policy neither requires nor prohibits
completion of a status determination once we have determined that a
population does not qualify as a DPS. Nevertheless, in this instance,
we concluded that completing an assessment--and detailing the nature,
scope, and likely effect of the threats to the population and the
species--would provide us and the public with useful information
regarding wolverines occupying the contiguous United States.
Summary of Biological Status and Threats
In preparing the SSA report for the wolverine, we reviewed
available reports and peer-reviewed literature, incorporated survey
information, and contacted species experts to collect additional
unpublished information for the North American subspecies (Gulo gulo
luscus), including Canada and Alaska. We identified uncertainties and
data gaps in our assessment of the current and future status of the
species. We also evaluated the appropriate analytical tools to address
these gaps and conducted discussions with species experts and prepared
updated maps of the known species' range in North America. In some
instances, we used publications and other reports (primarily from
Fenno-Scandinavia) of the Eurasian subspecies (G. g. gulo) in
completing this assessment.
Since the publication of the February 4, 2013, proposed listing
rule (78 FR 7864), several new wolverine studies have been published
(e.g., Aronsson 2017, Aronsson and Persson 2016, Aronsson et al. 2017,
Heinemeyer et al. 2019, Jokinen et al, 2019, Magoun et al. 2017,
Persson et al. 2017, Stewart et al. 2016, Webb et al. 2016, see
additional list of citations in the wolverine SSA report), which have
added to our understanding of wolverine biology while also highlighting
new insights into identifying key species' needs and their interactions
with both abiotic and biotic factors. This new information is
particularly relevant for a difficult-to-study animal like the
wolverine.
Using the species, individual, and population needs identified for
the wolverine and location results from surveys and studies, we
conducted a geospatial analysis to estimate the current potential
extent of occurrence for the North American wolverine in North America
including the contiguous United States (Figure 1; Service 2018).
``Current potential extent'' represents the perimeter of the outermost
geographic limits based on all (available) occurrence records (that is,
the maximum extent of occurrences) of a species minus those areas where
we believe the species has been extirpated (Service 2017). We then
evaluated this area and previous estimates of potentially suitable
habitat in the western-northwestern United States to
[[Page 64632]]
assess the species' current conditions within that region. Our future-
condition analysis includes the potential conditions that the species
or its habitat may face, that is, the most probable scenario if those
conditions are realized in the future. This most probable scenario
includes consideration of the sources that have the potential to most
likely impact the species at the population or rangewide scales in the
future, including potential cumulative impacts.
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Our analysis of potential future effects to the North American
wolverine and its habitat that are associated with climate change
(probabilistic estimates for temperature and precipitation) is
presented in the SSA report and summarized here. This analysis was
based on downscaled (high resolution local climate information derived
from global climate models) climate model projections, including a
detailed study of two regions in the western United States--Glacier
National Park (currently occupied by reproducing wolverines) and Rocky
Mountain National Park (occupied by a single male wolverine from 2009
to at least 2012, but not known to be currently occupied). The two
regions studied include a high-latitude area near tree line within
Glacier National Park, where tree line occurs at ~ 1,800 to 2,100 m
(5,906 to 6,890 ft) that is currently occupied by wolverines; and a
lower latitude area within Rocky Mountain National Park, where tree
line occurs at higher elevation (~ 3,500 m (11,483 ft)) (Ray et al.
2017, p. 2). These sites were selected to bracket the range of latitude
and elevation wolverines currently occupy in the contiguous United
States (Ray et al. 2017, p. 2).
For the purpose of this assessment, we generally define viability
as ``consisting of self-sustaining populations that are well
distributed throughout the species' range,'' and where ``[s]elf-
sustaining populations are those that are sufficiently abundant and
have sufficient genetic diversity to display the array of life history
strategies and forms that will provide for their persistence and
adaptability in the planning area over time'' (Committee of Scientists
1999, p. 38). We use a timeframe of approximately 38 to 50 years for
assessing future effects to wolverine viability. This timeframe
captures consideration of the projected future conditions related to
trapping/harvesting, climate change, or other potential cumulative
impacts (Service 2018, p. 73). Beyond this range, climate
[[Page 64633]]
modeling uncertainty increases substantially. We believe this is a
reasonable timeframe to consider as it includes the potential for
observing these effects over several generations of the wolverine.
As discussed above in Analytical Framework, we consider what the
species needs to maintain viability by characterizing the status of the
species in terms of resiliency, redundancy, and representation (Wolf et
al. 2015, entire). Resiliency is having sufficiently large populations
for the species to withstand stochastic events (arising from random
factors). We can measure resiliency based on metrics of population
health; for example, birth versus death rates and population size.
Resilient populations are better able to withstand disturbances such as
random fluctuations in birth rates (demographic stochasticity),
variations in rainfall (environmental stochasticity), or the effects of
anthropogenic activities.
Redundancy is having a sufficient number of populations for the
species to withstand catastrophic events (such as a rare destructive
natural event or episode involving many populations). Redundancy is
about spreading the risk and can be measured through the duplication
and distribution of populations across the range of the species. The
greater the number of populations a species has distributed over a
larger landscape, the better it can withstand catastrophic events.
Representation is having the breadth of genetic makeup of the
species to adapt to changing environmental conditions. Representation
can be measured through the genetic diversity within and among
populations and the ecological diversity (also called environmental
variation or diversity) of populations across the species' range. The
more representation, or diversity, a species has, the more it is
capable of adapting to changes (natural or human caused) in its
environment. In the absence of species-specific genetic and ecological
diversity information, we evaluate representation based on the extent
and variability of habitat characteristics within the geographical
range.
Life-History Needs
Wolverines are capable of moving and dispersing over great
distances over short periods of time. Wolverine populations are
characterized by naturally low densities in North America. The species
is highly territorial, with very little overlap between same-sex
adults. Wolverines occupy a variety of habitats, but generally select
habitat in locations away from human settlements. Wolverines consume a
variety of food resources, and seasonal switching of prey is commonly
observed. As with other Arctic mammals, wolverines have the ability to
adapt to both warm and cold ambient temperatures and solar radiation
through both physiological and behavioral responses, such as
vasodilation, increase in skin temperature, seasonal adjustments in fur
insulation, and micro- and macro-habitat selection.
Wolverine reproduction includes the following characteristics:
polygamous behavior (i.e., male mates with more than one female each
year), delayed implantation (up to 6 months), a short gestation period
(30-40 days), denning behavior, and an extended period of maternal
care. The reproductive behavior in wolverines is temporally adapted to
take advantage of the availability of food resources, limited
interspecific competition, and snow cover in the winter.
Since the publication of the Service's 2013 proposed rule to list
the distinct population segment of the North American wolverine in the
contiguous United States (78 FR 7864; February 4, 2013), several new
wolverine studies have been published (e.g., Aronsson 2017, Aronsson
and Persson 2016, Aronsson et al. 2017, Heinemeyer et al. 2019, Jokinen
et al, 2019, Magoun et al. 2017, Persson et al. 2017, Stewart et al.
2016, Webb et al. 2016, see additional list of citations in the
wolverine SSA report). These studies have improved our understanding of
wolverine biology while also highlighting new insights into identifying
key species' needs and their interactions with both abiotic and biotic
factors. Of particular importance relative to life history needs and
wolverine reproductive behavior, wolverine populations and wolverine
dens have been observed outside previously modeled projections of
spring snow cover.
Overall, the best available information indicates that within the
contiguous United States the wolverine's physical and ecological needs
include:
(1) Large territories in relatively inaccessible landscapes, at
high elevation (1,800 to 3,500 m (5,906 to 11,483 ft));
(2) access to a variety of food resources, which vary with seasons;
and
(3) physical/structural features (e.g., talus slopes, rugged
terrain) linked to reproductive behavioral patterns.
Current Condition
Current Potential Extent of Occurrence
As noted above, using the best available information on current
distribution and recent occurrences, we created maps to describe an
area of ``current potential extent of occurrence'' (current potential
extent) of wolverine for the western-northwestern contiguous United
States (Service 2018, pp. 12-13, 15). The current potential extent
represents the perimeter of the outermost geographic limits based on
all (available) occurrence records (that is, the maximum extent of
occurrences) for the wolverine minus those areas where we believe the
species has been extirpated (Service 2018, pp. 11-12). The current
potential extent area identified in Figure 2 encompasses approximately
280,316 km\2\ (69,267,592 acres (ac)) (Service 2018, p. 12). We also
prepared a current potential extent map for all of North America,
including Canada and Alaska, for a total estimated current potential
extent of 8,114,878 km\2\ (2,005,230,024 ac) (Service 2018, p. 12 and
Figure 1 of this document). The current potential extent area in the
contiguous United States represents approximately 3.5 percent of the
total current potential extent of wolverines in North America (Service
2018, p. 13 and Figure 2 of this document).
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Population Abundance and Density
Areas in the western contiguous United States have been previously
identified as suitable for wolverine survival (long-term survival; used
by resident adults) or primary habitat, reproduction (used by
reproductive females), and dispersal (female and male) of wolverines
(see methodology in Inman et al. 2013, pp. 279-280). From these
results, the researchers estimated potential and current distribution
and abundance of wolverines in the western contiguous United States.
They estimated current population size of wolverines to be 318
individuals (range 249-626) located within the Northern Continental
Divide (Montana) and areas within the following ecoregions: Salmon-
Selway (Idaho, portion of eastern Oregon), Central Linkage (primarily
Idaho, Montana), Greater Yellowstone (Montana, Idaho, Wyoming), and
Northern Cascades (Washington) (Inman et al. 2013, p. 282). Potential
wolverine population capacity based on habitat modeling was estimated
to be 644 individuals (range: 506-1,881) (Inman et al. 2013, p. 282);
however, we do not have information indicating wolverine abundance in
the contiguous United States.
In the wolverine SSA report, we provide a discussion of recent
studies of wolverine detections and observations in the western United
States (Service 2018, pp. 51-56); however, no comprehensive surveys
have been conducted across the entire area defined as the species'
maximum extent of occurrence (Service 2018, p. 14; Figure 2) or current
potential extent of occurrence (Figure 2 above) in the contiguous
United States. Below we
[[Page 64635]]
provide a summary of recent wolverine observations and detections in
the western United States.
A recent study (2007-2015) has demonstrated that the Cascades
region of Washington and Canada supports a resident wolverine
population (Aubry et al. 2016, p. 40). For the first time in recent
history, a breeding female wolverine was detected south of I-90 in the
south Cascades of Washington, as well as her potential mate, indicating
wolverines may be extending their current range in that area (Flatt
2018, p. 1). Wolverines have been detected in the Eagle Cap Wilderness
Area in the Wallowa Mountains of northeastern Oregon in 2011-2012,
2016, and 2017 (Magoun et al. 2013, p. 17; Magoun 2017, pers. comm.).
In California, a single male wolverine occurs in the Truckee area as of
March 2017 (Shufelberger 2017, pers. comm.).
Since 2010, survey and monitoring efforts in the Idaho Panhandle
and adjoining areas of Washington, Montana, and British Columbia,
Canada, have detected five individual male wolverines (Service 2018, p.
52). One male was also detected in British Columbia, north of Canadian
Highway 3 (Lucid et al. 2016, p. 184), which some consider to be a
barrier to wildlife passage (IDFG 2017, pers. comm.). This male was
most recently detected in Idaho, on March 6, 2013 (Lucid et al. 2016,
p. 175). One likely wolverine den was located in the Saint Joe
Mountains in Idaho (Lucid et al. 2017, p. 12).
Results from a pilot study to evaluate wolverine occupancy in
Wyoming indicated at least three individual wolverines (at five
stations) with at least one individual in the Gros Ventre and Wind
River mountain ranges, and at least two individuals in the Southern
Absaroka mountain range (Inman et al. 2015, p. 9). Occupancy modeling
estimated a probability of occupancy for sampled sites of 62.9 percent
(Inman et al. 2015, p. 8).
Building on the results of the Wyoming pilot study, the Western
Association of Fish and Wildlife Agencies (WAFWA), in coordination with
Tribal partners, formed a multi-State, multi-agency working group
(Western States Wolverine Working Group) to design and implement the
Western States Wolverine Conservation Project (WSWCP)-Coordinated
Occupancy Survey. The primary objectives of the WSWCP include: (1)
Implement a monitoring program to define a baseline wolverine
distribution and genetic characteristics of the metapopulation across
Montana, Idaho, Wyoming, and Washington; (2) model and maintain the
connectivity of the wolverine metapopulation in the western United
States; and (3) develop policies to address socio-political needs to
assist wolverine population expansion as a conservation tool, including
translocation of wolverines (IDFG 2016, pers. comm.; Montana FWP 2016,
pers. comm.; WGFD 2016, pers. comm.).
The Wyoming Game and Fish Department began implementation of the
survey in Wyoming in the Greater Yellowstone Ecosystem region and the
Bighorn Mountains in the winter of 2015-2016 (WGFD 2016, pers. comm.).
That initial survey detected at least three unique wolverines in the
Wind River and southern Absaroka Mountain Ranges (WGFD 2016, pers.
comm.).
The monitoring effort was expanded in the winter of 2016-2017 in
four States (Washington, Idaho, Montana, and Wyoming), and our review
of the results indicate that wolverines were detected in all four
States (Service 2018, p. 53). From this study, a total of 43 unique
individuals were identified, 20 males and 23 females (Pilgrim et al.
2018, no page number).
We also received additional wolverine observations from State and
Federal agencies in northwestern Wyoming. A wolverine was detected by
camera in northern Grand Teton National Park, and a member of the
public reported wolverine tracks in southwestern Grand Teton National
Park while skiing, which was confirmed by a Forest Service biologist
(Service 2018, p. 53). Both of these observations occurred in March
2017. South of this area in the Wyoming Range (about 4 miles east of
Alpine, Wyoming), a wolverine was detected by camera in May 2017
(Service 2018, p. 53).
Wolverine densities vary across North America and have been
described as naturally low and wolverine populations as naturally
uncommon given the species' large home range, wide-ranging movements,
and solitary characteristics (Service 2018, p. 56). In the contiguous
United States, density estimates (number of wolverines per 1,000 km\2\
(386 mi\2\)) ranged from 3.5 for the Greater Yellowstone region (2001-
2008) (areas above 2,150 m (7,054 ft) (latitude-adjusted elevation),
4.5 for central Idaho (1992-1995), to 15.4 for northwestern Montana
(1972-1977) (Service 2018, p. 57).
We note here that in our 2013 proposed listing rule for the
wolverine (78 FR 7864), we discussed the occurrences of two dispersing
individuals in California and Colorado (the Colorado wolverine was
later killed in North Dakota). We know of one male wolverine in
California that has consistently occupied an area much farther north in
the Sierra Nevada Mountains, and we have no evidence of any other
wolverines currently in the State. We have no recent records of
wolverines in Arizona. Aubry et al. (2007, p. 2,150) identified the
year 1860 as the most recent verifiable documentation of wolverines in
northern New Mexico. We know of no wolverines currently occupying
Colorado. As presented in Aubrey et al. (2007, p. 2,151; Figure 1),
prior to 1900, the most recent verifiable record for wolverine in New
Mexico was 1860 and 1887 for Nebraska; no records were found for
Arizona, Texas, Oklahoma. This was also true for most midwestern and
mid-Atlantic States (Aubrey et al., 2007, p. 2,152, Figure 1).
Additionally, historical range maps shown in Seton (1909, p. 947; Map
51), Aubrey et al. (2007, p. 2,152; Figure 1), and the assessment and
status review for the wolverine in Canada (COSEWIC 2014, p. 12; Figure
3) do not extend the distribution of wolverines into these regions.
Our updated analysis of wolverine occurrence in the contiguous
Unted States is based on a more scientifically robust and spatially
explicit assessment of the current areas occupied by wolverines in the
contiguous United States, which was prepared based on verifiable
wolverine records and comments received by reviewers of a draft of the
wolverine SSA report (see the Historical Range and Distribution section
of the wolverine SSA report for more on the information used to assess
the maximum extent of occurrences (`historical range') and current
extent of occurrence (Service 2018, pp. 9-16; Figures 2-4)). Using the
current potential extent of occurrence, as presented in Figures 1 and 2
above, provides a more accurate reflection of the areas currently
occupied by wolverines in the contiguous United States supported by the
best available information.
Alaska and Canada
In the wolverine SSA report, we provide a summary of population
abundance in Alaska and Canada where wolverines are more abundant than
in the contiguous United States (Service 2018, pp. 57-60). Much of what
we know about wolverine occurrences and abundance has been gathered
from trapping records (see summary in Service 2018, pp. 53-56).
In Alaska and Yukon, density estimates presented by Inman et al.
(2012, p. 789) range from 3 to about 14 wolverines per 1,000 km\2\ (386
mi\2\), using a number of methods. For
[[Page 64636]]
example, Royle et al. (2011, p. 609) estimated wolverine densities for
southeastern Alaska (Tongass National Forest; 2008) from 8.2 to 9.7 per
1,000 km\2\ (386 mi\2\) (using mark-recapture), where the higher
estimate incorporates a positive, trap-specific behavioral response.
Density of wolverines were recently reported as an estimated 5-10
wolverines per 1,000 km\2\ (386 mi\2\) (based on snow tracking) for
southcentral Alaska, and approximately 10 per 1,000 km\2\ (386 mi\2\)
(based on DNA mark-recapture methods) for southeastern Alaska (Golden
2017, pers. comm.). A wolverine occupancy study in 2015 within an area
of central Alaska reported a density estimate of 9.48 wolverines per
1,000 km\2\ (386 mi\2\) (Alaska Department of Fish and Game (ADF&G)
2015, p. 7).
Wolverine density estimates for Canada vary across regions, from 5
to 10 per 1,000 km\2\ (386 mi\2\) in northern mountain and boreal
regions to 1 to 4 per 1,000 km\2\ (386 mi\2\) in southern boreal areas
(COSEWIC 2014, p. 27). More recently, Clevenger et al. (2017, entire)
presented a density estimate (using spatial capture/recapture models)
for the Kootenay region of British Columbia of 0.78 wolverines per
1,000 km\2\ (386 mi\2\), for 3 study years (2014-2016), which they
reported as lower than expected (Clevenger et al. 2017, p. 6).
Researchers in Canada are currently conducting a landscape level
analysis to estimate the size and sustainable harvest for wolverine
populations within British Columbia (Weir 2017, pers. comm.).
According to the most recent COSEWIC Assessment and Status Report
on the Wolverine, Gulo gulo in Canada (COSEWIC 2014, entire), Canada's
western subpopulation has been estimated at 15,688 to 23,830 adults,
which we recognize is an estimate based on several assumptions, such as
consistent trapping effort and uniform densities across the species'
range (COSEWIC 2014, p. 36). In Alaska, estimates of populations are
not available and are best evaluated based on density with recent
density estimates ranging from 5 to 10 wolverines per 1,000 km\2\ (386
mi\2\) for Alaska (Parr 2017, pers. comm.). In Alaska, which, like
Canada, allows regulated hunting and trapping of wolverines, an average
of 590 wolverines have been taken each year over the past 6 years
(Service 2018, p. 68). The consistent harvest levels in these regions
suggest relatively stable wolverine populations in Alaska that more
likely than not number in the thousands of individuals in order to
sustain such level of harvest.
We do not have reliable current population estimates for wolverines
in the contiguous United States. As discussed above, the only estimate
available is from 2013, when researchers, using spatial modeling
methods, estimated the then-current population size of wolverines to be
318 (range: 249-626) (Inman et al. 2013, p. 282). Potential wolverine
population capacity in the contiguous United States based on habitat
modeling was estimated to be 644 individuals (range: 506-1,881) (Inman
et al. 2013, p. 282). However, these capacity estimates did not
consider spatial characteristics related to behavior, such as
territoriality (home range), of wolverine populations. Given all the
assumptions, differing methods of estimation, limitations, and
uncertainties of the available estimates of North American wolverines
(as discussed in the wolverine SSA report (Service 2018, pp. 50-56)),
we believe caution should be used relative to comparing the number of
wolverines in the contiguous United States to the remainder of the
taxon. However, even assuming the high population estimate from 2013
for the contiguous United States (n=626) and the low estimate of
wolverines in western Canada from 2014 (15,688 adults), the contiguous
United States conservatively contains approximately 4 percent of the
total wolverines within these two regions. This estimate does not
account for wolverines in Alaska, for which we have no population
estimate, but, based on a rough estimate of land area for the State
occupied by wolverines and estimated wolverine densities of between 5
to 10 animals per 1000 km\2\ (386 mi\2\) (Parr 2017, pers. comm.), it
is reasonable to assume there are thousands of wolverines in the State.
The actual percentage of wolverines in the contiguous United States
compared to the overall taxon (Canada and Alaska included) is still
significantly less than 4 percent of the overall North American
wolverine population. Wolverine densities vary across North America and
have been described as being naturally low, due in large part to the
species having large home ranges, wide-ranging movements, and solitary
characteristics (Service 2018, p. 56). It is important to understand
that the amount of suitable habitat in the contiguous United States
identified both in historical and current distribution maps (see, for
example, 1909, p. 947; Map 51), Aubrey et al. (2007, p. 2,152) does not
support the larger numbers of wolverines and higher densities found in
Canada and Alaska (see Figure 3 in the wolverine SSA report (Service
2018, p. 15)).
Summary of Factors Affecting the North American Wolverine
As mentioned above in Regulatory Framework, a species may be
determined to be an endangered or threatened species due to one or more
of the five factors described in section 4(a)(1) of the Act: (A) The
present or threatened destruction, modification, or curtailment of its
habitat or range; (B) overutilization for commercial, recreational,
scientific, or educational purposes; (C) disease or predation; (D) the
inadequacy of existing regulatory mechanisms; or (E) other natural or
manmade factors affecting its continued existence. Listing actions may
be warranted based on any of the above threat factors, singly or in
combination. Potential stressors evaluated for wolverine in the
contiguous United States include effects from roads (Factors A and E);
disturbance due to winter recreational activity (Factors A and E);
other human disturbance (Factors A and E); effects from wildland fire
(Factor A); disease (Factor C); predation (Factor C); overutilization
(trapping) (Factor B); genetic diversity (Factor E); small-population
effects (Factor E); and climate change (Factors A and E). A summary of
the potential stressors affecting wolverine in the contiguous United
States is presented below; for a full description of our evaluation of
the effects of these stressors, refer to the wolverine SSA report
(Service 2018, pp. 57-101).
Effects from Roads: Wolverines are associated with habitat found in
high-elevation areas, but are known to disperse over great distances.
Major highways can present mortality risks to dispersing individuals
and affect immigration to open territories, but roads do not represent
absolute barriers to wolverine movements. Wolverines den during winter
months in locations that are often inaccessible or restricted to
motorized vehicles, though secondary roads and trails are used for
winter recreational activity. Although we recognize there are likely
additional events that have not been reported, we estimated the total
number of wolverine mortalities due to roads from 1972 to 2016 (44
years) in North America was 20, at least 11 of which are from Canada
(Service 2018, p. 60). In the SSA report, we calculated a low
proportion of major highways in both modeled primary habitat and a low
mean density of roads at high elevations where wolverines have been
observed, with the exception of the southern Rocky Mountains (Service
2018, p. 60). We therefore determine that the effects from roads
present a low stressor to wolverines at the individual and population
level in
[[Page 64637]]
most of its current area of occupancy within the contiguous United
States.
Disturbance due to Winter Recreational Activity: Wolverine behavior
patterns, such as denning, rearing of young, movement and dispersal,
and foraging/scavenging, may be affected by recreational activities
(COSEWIC 2014, p. 42), although several wolverines have been captured
for research on or near ski areas (e.g., Teton Mountains) (Montana FWP
2017, pers. comm.). In Norway, one study found, at the home-range
scale, a minimal threshold distance of approximately 1.5 km (0.93 mi)
for wolverine den sites from private roads and/or recreational cabins
(May et al. 2012, p. 201). Another study found that in an area of
active recreation (Columbia Mountains, Canada), female wolverines were
negatively associated with helicopter and backcountry skiing in their
winter models (Krebs et al. 2007, pp. 2,187-2,188). In summer months,
Copeland et al. (2007, p. 2,210) reported that wolverines in their
study area of central Idaho were not uncommonly found near maintained
trails and active campgrounds, which suggests some level of tolerance
to human presence/recreational activity.
The Wolverine-Winter Recreation Study represents an ongoing project
to evaluate the potential effects of backcountry winter recreation
(e.g., backcountry skiers, heli-skiers, cat-skiers, snowmobilers) on
wolverines in central Idaho and areas in the western Yellowstone region
(Island Park area and Teton Mountains) (Heinemeyer 2016, pers. comm.;
Heinemeyer 2019, entire; Heinemeyer and Squires 2015, p. 3). Early
analysis of the data suggested that wolverines demonstrate a behavioral
response to recreation activities, such as increased movement rates and
a reduction in resting periods in areas of high-recreation activity,
especially high-recreation days (Saturday and Sunday) (Heinemeyer and
Squires 2013, pp. 5, 7-8). However, this research also found that
wolverines maintained their home ranges within areas with relatively
high winter-recreation activity over several years of monitoring,
including some areas found to contain the highest recreational
activities (Heinemeyer 2016, pers. comm.). The study has not been able
to determine whether these resident wolverines are reproductively
successful due to the limited monitoring information available for
reproductive females (Heinemeyer 2016, pers. comm.).
A final Winter Recreation Study report found that wolverines
maintained multi-year home ranges in areas that support relatively
intensive winter recreation, suggesting that wolverines are able to
tolerate winter recreation at some scales (Heinemeyer et al. 2017, p.
iv; Heinemeyer et al. 2019, p. 16). Wolverines responded negatively to
increasing intensity of winter recreation, with off-road and dispersed
recreation having a greater effect than recreation that was
concentrated on access routes (Heinemeyer et al. 2017, p. 34;
Heinemeyer et al. 2019, p. 13). Wolverine avoidance of roads and
groomed areas used by winter recreationists was found to be less than
estimated for dispersed recreation, suggesting that wolverines may be
less sensitive to predictable winter-recreational use patterns
(Heinemeyer et al. 2017, p. 40; Heinemeyer et al. 2019, p. 15). Habitat
selection in females evaluated in the multi-year study was complex, and
likely driven by a combination of abiotic (snow, cold) and biotic
factors (predator avoidance, food availability) (Heinemeyer et al.
2017, p. 36; Heinemeyer et al. 2019, p. 16). This study did not assess
demographic effects, fitness effects, or population level effects of
winter recreation on wolverines (Heinemeyer et al. 2019, p. 17 and 19).
Conservation measures currently being implemented that address the
effects of roads in the Teton Mountains include winter closures in
certain areas (generally from November 1 through May 1), including road
closures in the Bridger-Teton and Caribou-Targhee National Forests and
in Grand Teton National Park (Service 2018, p. 67, Appendix F). These
closures are being implemented to help minimize disturbance to wildlife
(e.g., migration pathways). State Wildlife Action Plans prepared for
individual western States identify recreation management strategies
within wolverine habitats. For example, in Oregon, the Oregon
Department of Fish and Wildlife Conservation Strategy identifies
management of winter-recreation use as a conservation action to avoid
impacts to wolverines (ODFW 2016). In Montana's State Wildlife Action
Plan, conservation actions for the wolverine are identified to address
potential impacts from recreation, such as consideration of seasonal
closures during denning season (Montana FWP 2015, p. 63). The Idaho
Department of Fish and Game Management Plan for the Conservation of
Wolverines in Idaho also includes conservation strategies related to
developing a better understanding of the relationships between
wolverine behavior and winter recreation activities (IDFG 2014, p. 35),
and the State continues to support the Wolverine-Winter Recreation
Study. Appendix G in the SSA report provides additional details on
individual State conservation strategies. Although we do not rely on
these conservation measures to support our decision on listing status,
they do provide some protections to address potential impacts to
wolverine from disturbance from winter recreational activity and
mortality from roads.
Based on the studies summarized above, wolverine behavior
(movement) is potentially affected by winter recreational activity.
However, wolverines can maintain residency in high winter-recreational
use areas (Heinemeyer et al. 2017, p. iv; Heinemeyer et al. 2019, p.
16). Based on the best available scientific and commercial information,
the effect of winter recreational activity represents a low stressor to
wolverines in the contiguous United States at the individual and
population level.
Other Human Disturbance: Infrastructure, such as pipelines, active
logging or clearcuts, seismic lines, and activities associated with
mining (e.g., producing mines, mines under development, mineral
exploration areas), may also affect individual wolverine behavior
(e.g., avoidance) or loss or modification of wolverine habitat. In the
SSA report, we summarize a recently published study of habitat
selection of wolverines in response to human disturbance in western
Canadian forested habitat (Service 2018, p. 62). That study found that
wolverines avoided interior areas of some logged areas, but also found
that wolverines were attracted to all-season road sections with borrow
pits (Scrafford et al. 2017, pp. 32-34). The authors concluded that
wolverine selection patterns relative to industrial activity and
infrastructure in their study area represented a balance between
exposure to predators and foraging opportunities (Scrafford et al.
2017, p. 32). Based on the best available scientific and commercial
information, we find that these human disturbance effects are likely to
be small or narrow in scope and scale for wolverines in the contiguous
United States.
Effects from Wildland Fire: Wildland fire can produce both direct
and indirect effects to wildlife. Direct effects include injury and
mortality as well as escape or emigration movement away from fires
(Lyon et al. 2000, pp. 17-21). We are unaware of any studies evaluating
direct effects of wildland fire to wolverines. Wildland fire is likely
to temporarily displace wolverines, which could affect home range
dynamics. Given that
[[Page 64638]]
wolverines can travel long distances in a short period of time,
individuals would be expected to move away from fire and smoke
(Luensmann 2008, p. 14). In addition, because young wolverines are born
in underground or otherwise sheltered dens during winter months and in
locations where wildland fire risk is low due to snow cover or
increased moisture (Luensmann 2008, p. 14), the potential effects of
fire at that critical life stage is very low (Luensmann 2008, p. 14).
Indirect effects of wildland fire can include habitat-related effects
or effects to prey and competitors/predators; however, we are unaware
of empirical studies evaluating these potential effects as they relate
to wolverines.
Given the diversity of habitats occupied by wolverines, their
opportunistic foraging habitats and seasonal switching of food sources,
their occupancy of high elevations, and extensive mobility, wildland
fire represents a limited indirect and direct stressor, in scope and
scale, to wolverine habitat and its prey in the contiguous United
States range (Service 2018, pp. 63-64) such that it would not be
expected to have population or species-level impacts.
Disease: We are unaware of comprehensive surveys evaluating the
prevalence of diseases in wolverines in the contiguous United States.
Other than a parasitic pneumonia mortality event and a single rabies
case, we are not aware of any other studies documenting impacts of
disease to wolverines in North America (Service 2018, p. 65). At this
time, based on the best available scientific and commercial
information, we do not find that disease is a population- or species-
level stressor to the wolverine in the contiguous United States
(Service 2018, pp. 64-65).
Predation: A number of potential natural predators have been
identified for wolverines within its North American range, including
intraspecific predation (Service 2018, p. 65). However, we have no
information that suggests predation represents a significant stressor
to the wolverine at the population level. At the individual level, we
recognize that wolverines likely avoid areas of potential predation
risk from wolves and other potential predators (Service 2018, p. 65).
Thus, indirect effects of predators may result in predator avoidance
behavior of individual wolverines through habitat selection. However,
the best scientific and commercial information available indicates that
predation is not a stressor for the wolverine (Service 2018, p. 65).
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes: During the late 1800s and early 1900s, the
wolverine population declined or was extirpated in much of the
contiguous United States, which has been attributed in large part to
unregulated persecution (Service 2018, p. 1). Similar range reductions
and extirpations of some wolverine populations were observed in parts
of Canada during this time period (van Zyll de Jong 1975, entire;
COSEWIC 2014, p. iv). However, after unregulated harvest of wolverines
ceased, the numbers of wolverines in Canada and the contiguous United
States began to recover from this decline (e.g., Aubry et al., 2007, p.
2,151; Aubry et al., 2012, entire; Aubry et al. 2016, pp. 14-15; Magoun
et al. 2013, p. 27).
In Montana, wolverines were a legally harvested furbearer up until
2012 (Service 2018, p. 65). There is, however, no evidence to suggest
that the harvest of wolverines in Montana at historical rates (about 10
animals per year) was detrimental to wolverine populations (Service
2018, pap. 65-66 and Appendix G). Furthermore, States within the
wolverine range in the contiguous United States have adopted protective
regulations to prevent unauthorized take and are implementing other
measures to limit incidental mortality of wolverines (Service 2018, p.
66). There is currently no allowable trapping or harvesting of
wolverines in the contiguous United States, although incidental
trapping, shooting, and poisoning mortalities have been documented
(Service 2018, pp. 65-69).
In Alaska, wolverine trapping and hunting is controlled by seasons
and bag limits, with about 550 animals harvested each year (ADF&G
2017a). This level of harvest has been fairly consistent since 2010
(Service 2018, Table 7).
Trapping and harvesting of wolverines occurs over much of the range
in Canada (COSEWIC 2014, pp. 10, 29-35). Specifically, wolverines are
harvested in the northern and western territories--Manitoba,
Saskatchewan, Alberta, British Columbia, Yukon, Northwest Territories,
and Nunavut (COSEWIC 2014, p. 43). The population of wolverines in
British Columbia is estimated to be 2,700-4,760 and 1,500-2,000 animals
in Alberta (COSEWIC 2014, p. 36). In the wolverine SSA report, we
evaluated trapping of wolverines in British Columbia and Alberta
regions of southern Canada in an effort to document potential impacts
to dispersing wolverines along the U.S.-Canada border (Service 2018,
pp. 68-69). This type of analysis was not conducted for the 2013
proposed listing rule (78 FR 7864; February 4, 2013) or for our 2014
withdrawal (79 FR 47522; August 13, 2014). The results of our spatial
analysis for British Columbia indicates a total of 77 wolverines were
trapped in wildlife management units within 110 km (68.35 mi) of the
U.S.-Canada border in the period 2007-2015, or an average of 8.5
animals per year (Service 2018, pp. 68-69). We used this distance since
it is similar to both the average maximum distance per dispersal
movement of 102 km (63 mi) for male wolverines in the Greater
Yellowstone region of Montana (Inman et al. 2012, p. 784), and a
reported 100-km (62-mi) dispersal distance for a juvenile male for
Ontario, Canada (COSEWIC 2014, p. 24, citing unpublished data from
Dawson et al. 2013). For Alberta, we identified a total of 15
wolverines harvested by trappers and data presented in other studies
within 110 km (68.35 mi) of the U.S.--Canada border in the period 1989-
2014 (average of less than 1.0 animal per year) (Service 2018, p. 68).
Based on this new analysis, legal trapping effort along the U.S.-
Canada border does not represent a barrier to wolverine movement and
dispersal along the international border. As discussed below and in the
DPS analysis above, results from genetic analyses provide further
evidence of movement and dispersal of wolverines across the
international border (see Genetic Diversity below).
In summary, overutilization does not currently represent a stressor
to the wolverine in the contiguous United States at the individual,
population, or species level. Wolverine populations in the contiguous
United States are currently protected under several State laws and
regulations. Regulated hunting and trapping activities for wolverines
are currently suspended or closed entirely for animals that occupy
western States of the contiguous United States, though occasional
incidental trapping can occur. Current trapping in Alaska and Canada
appears to be sustainable and wolverine populations along the Alaska--
Canada border are continuous with the Yukon region of Canada, which
suggests a rescue effect (animals from a higher population density area
moving to areas of lower population density, preventing local
extirpation) for Canadian populations along this international boundary
(COSEWIC 2014, p. 37). Trapping or harvesting of wolverines along the
contiguous U.S.-Canada border does not represent a barrier or stressor
to wolverines migrating into the contiguous United States at the
individual or population level.
[[Page 64639]]
Genetic Diversity: The geographical genetic structure of wolverines
is believed to be largely structured around the strong female
philopatry characteristic of this species (Rico et al. 2015, p. 2) and
the species' polygamous behavior. Results from Scandinavia indicate
that wolverine population distributions are primarily limited by
dispersal of the more philopatric sex (females) (Aronsson 2017, p. 13).
The extensive and often asymmetrical movement of male wolverines from
core populations to the periphery of their range can result in the
addition of nuclear genetic material to these edges (Zigouris et al.
2012, p. 1,553). Thus, the dispersal pattern for male wolverines may
help explain why allelic richness (i.e., nuclear DNA, which is
inherited from both parents) can be similar across regions, but
haplotype richness (mitochondrial DNA, which is maternally inherited)
is lower at the periphery of the species' range (Zigouris et al. 2012,
p. 1,553).
Studies evaluating the genetic structure of wolverines, primarily
within its core range in North America, were presented in Chappell et
al. (2004) and Kyle and Strobeck (2001, 2002). Using microsatellite
markers, Kyle and Strobeck (2002) and Zigouris et al. (2012) found
greater genetic structure of wolverines toward the eastern and southern
peripheries of their North American distribution, likely due to a west-
to-east recolonization during the Holocene (Zigouris et al. 2013, p.
9). Similarly, based on an evaluation of mitochondrial DNA, which is
used primarily for an evaluation of phylogenetic structure and
phylogeography, McKelvey et al. (2014, p. 330) concluded that modern
wolverine populations in the contiguous United States are the result of
recolonization (following persecution during a period of unregulated
hunting or trapping and poisoning) from the north.
Genetic diversity and population genetic structure of a larger
sample size of wolverines were examined by Cegelski et al. (2006,
entire) for the southern extent of their North American range using
both microsatellite markers and mitochondrial DNA. They concluded that
the wolverine populations in the contiguous United States were not
sources for dispersing individuals into Canada (Cegelski et al. 2006,
p. 208). They found that there was significant differentiation between
most of the populations in Canada and the United States (Cegelski et
al. 2006, p. 208). However, they cautioned that their statistical
analysis may not have been able to detect ``effective migrants'' and
that sample size can affect the detection of dispersers (Cegelski et
al. 2006, p. 208). They concluded that some migration of wolverines was
occurring between the Rocky Mountain Front region (northwestern
Montana) and Canada as well as among wolverine populations in the
United States, with the exception of Idaho (Cegelski et al. 2006, p.
208).
This study also conducted model simulations of the number of
effective wolverine breeders necessary to maintain genetic variation
(heterozygosity) in their sampled population of the contiguous United
States in the absence of gene flow (Cegelksi et al. 2006, p. 201). They
indicated that two effective migrants from either Canada or Wyoming
into the Rocky Mountain Front population would be needed (per
generation, 7.5 years) to maintain the levels of genetic diversity in
that population, and one effective migrant was needed to maintain
levels of diversity in the Gallatin, Crazybelt, or Idaho populations
(Cegelski et al. 2006, p. 209). They also found that to maintain at
least 95 percent of the genetic variation in the next 100 generations
(we estimate this to be approximately 750 years, based on generation
time) 200 to 300 wolverine breeding pairs were needed in the Wyoming
and Rocky Mountain Front populations, respectively, and 200 breeding
pairs were needed in the Gallatin, Crazybelts, and Idaho wolverine
populations (Cegelski et al., 2006, pp. 208-209). The authors concluded
that migration is essential for maintaining diversity in wolverine
populations in the contiguous United States since effective population
size may never be reached due to the naturally low population densities
of wolverines (Cegelski et al. 2006, p. 209).
More recently, an analysis of mitochondrial DNA was prepared for
wolverine samples collected during the winters of 2015-2016 and 2016-
2017 as part of the Western States Wolverine Conservation Project-
Coordinated Occupancy provides further support that all contiguous
United States historical (pre-1900) and recent wolverine populations
are likely descendants of immigrants from Canada and suggest continued
connectivity between the contiguous United States and Canadian
wolverine populations (Pilgrim and Schwartz 2018, entire).
Effective population size (Ne) is defined as ``the size
of an idealized population that would experience the same amount of
genetic drift and inbreeding as the population of interest. In popular
terms, Ne is the number of individuals in a population that
contribute offspring to the next generation'' (Hoffman et al. 2017, p.
507; see also Service 2018, Box 2). It represents a metric for
quantifying rates of inbreeding and genetic drift and is often used in
conservation management to set genetic viability targets (Olsson et al.
2017, p. 1). It is not the same as the more commonly used metric,
census population size (N), but is often assumed to represent the
genetically effective population size.
In his review of the minimum viable population size concept, Ewens
(1990, entire) emphasized that the term ``effective population size''
is not a meaningful term unless additional context is provided relative
to which concept of population size is being evaluated (Ewens 1990, p.
309). He introduced the concept of mutation effective population size,
defined as the size of population defined by its capacity to maintain
genetic variation (Ewens 1990, p. 307), which is different than actual
population size (Ewens 1990, p. 309). Demographic factors such as sex
ratio, subpopulations, dispersal, and immigration are needed when
interpreting actual population size from an effective population size;
thus, there is no justification for a fixed, genetically derived
minimum viable population size value of `500' as each case is unique
(Ewens 1990, p. 310). A review of the minimum viable population concept
by Flather et al. (2011, entire) also found that any ``rule of thumb''
used for minimum viable population will likely be a poor estimate for
that population (Flather et al. 2011, pp. 311, 313). Minimum viable
population estimates therefore vary considerably both within and among
species and are sensitive to the timeframe in which data are collected
(Flather et al. 2011, p. 314).
An effective population size analysis for wolverines in the
contiguous United States was presented in Schwartz et al. (2009, p.
3,225) using wolverine samples from the main part of the Rocky
Mountains populations (e.g., central and eastern Idaho, Montana,
northwestern Wyoming). Subpopulations from the Crazy and Belt Mountains
in Montana were excluded from this analysis based on suggestion by
Cegelski et al. (2003) that they represented separate groups (Schwartz
et al. 2009, p. 3,225). The summed effective population size was
estimated at 35, with credible limits from 28-52, and the summed values
for the three timeframes was reported as follows: Ne 1989-
1994 = 33, credible limits 27-43; Ne 1995-2000 = 35,
credible limits 28-57; Ne 2001-2006 = 38, credible limits
33-59 (Schwartz et al. 2009, p. 3,226). Thus, the two later time-frames
evaluated indicate an
[[Page 64640]]
(increasing) effective population size with credible limits above 50.
Of direct relevance to potential gene flow and genetic structure at
the landscape level, wolverines travel (disperse) through areas outside
high-elevation, forested habitats. For example, tracked dispersal
movements of a male wolverine, M56, from Wyoming into Colorado and its
subsequent discovery in North Dakota, indicate extensive travel outside
of modeled primary wolverine habitat (i.e., Inman et al. 2013),
including through arid grasslands and shrubland habitats of the Wyoming
Basin ecoregion (Packila et al. 2017, entire). This animal's movement
also supports some level of connectivity (and gene flow) between
currently occupied habitat (Wyoming) and unoccupied habitat within the
wolverine's historic range (Colorado) (Packila et al. 2017, p. 404).
Similarly, wolverines have recently moved from Washington and Idaho
into British Columbia, and earlier from Montana to British Columbia and
Alberta (Service 2018, p. 45). Based on genetic analyses, the male
wolverine currently occupying an area within the Sierra Nevada
Mountains of California also represents evidence of connectivity
between wolverine populations of the Rocky and Sierra Nevada Mountain
Ranges (Moriarty et al. 2009, p. 154). Wolverines have been detected
making cross-valley movements in the Southwestern Crown of the
Continent (SWCC) in northwestern Montana, which researchers believe is
an indication of good connectivity in this region (SWCC Working Group
2016, pers. comm.).
It can be difficult to make inferences about the relationship
between population size and point estimates of genetic diversity
without continued genetic monitoring and an understanding of the
demographic history of a species' population (Hoffman et al. 2017, p.
507), including factors that have historically influenced and continue
to influence movement patterns and connectivity. Additionally, the
extensive dispersal movements of both male and female wolverines can
produce gene flow among diverged populations, making it difficult to
distinguish, without additional sampling and analysis, between long-
distance dispersal and fragmentation based on the patchy distribution
of some haplotypes (Zigouris et al. 2013, p. 10). Genetic diversity can
be a reflection of favorable adaptations (natural selection) and is
necessary for species to locally adapt to environmental stressors or to
facilitate range shifts (Zigouris et al. 2012, p. 1,544). Genetic
distinctiveness in peripheral populations may therefore play a role in
both maintaining and generating biological diversity for a species
(Zigouris et al. 2012, p. 1,544; citing results presented in Channell
and Lomolino 2000, p. 84). Relatedly, genetic variation that is
adaptive is a better predictor of the long-term success of populations
as compared to overall genetic variation (Hoffman et al. 2017, p. 510).
The challenge is to be able to determine whether genetic variation is
adaptive and is a reflection of remnants of high genetic diversity from
ancestral populations, or whether that variation is a reflection of
accumulated deleterious, nonadaptive genes due to genetic drift in
small populations (Hoffman et al. 2017, p. 509).
In summary, the currently known spatial distribution of genetic
variability in wolverines in North America appears to be a reflection
of a complex history where population abundance has fluctuated since
the time of the last glaciation and insufficient time has passed since
human persecution for a full recovery of wolverine densities (Cardinal
2004, pp. 23-24; Zigouris et al. 2012, p. 1,554). Zigouris et al.
(2012, p. 1,545) noted that the genetic diversity reported in Cegelski
et al. (2006) and Kyle and Strobeck (2001, 2002) for the southwestern
edge of the North American range represented only part of the diversity
in the northern populations of wolverines. Zigouris et al. (2012, p.
1,545) posit that the irregular distribution of wolverines in the
southwestern periphery and the genetic diversity observed in those
analyses is a result of population bottlenecks that were caused by
range contractions from a panmictic (random mating) northern core
population approximately 150 years ago coinciding with human
persecution. As described here, recent dispersals of wolverines into
Colorado, California, and Utah provide evidence for connectivity and
the potential for gene flow between Northern Rocky Mountain populations
and areas where wolverines were extirpated.
As noted above in this section (and in the Distinct Population
Segment section), there is recent evidence of wolverines traveling
across the international border. Furthermore, our analysis of trapping
levels in the wolverine SSA report (summarized in Overutilization for
Commercial, Recreational, Scientific, or Educational Purposes above)
does not support previous assumptions that trapping in Canada near the
border acts as a barrier to wolverine movement into the contiguous
United States. Cegelski et al. (2006, p. 209) determined that very few
successful migrants are needed per generation to maintain at least 95
percent of the genetic variation in the next 100 generations
(approximately 750 years) in the contiguous United States (Cegelski et
al. 2006, p. 209). We have no reason to believe that this level of
migration from the north has not already been occurring following the
end of intense persecution of this species to repopulate previously
occupied areas within the contiguous United States. This repopulation
has occurred without human-assisted introductions and with unregulated
trapping from about the 1930s to 1970 in Montana. Given the recent
observations of dispersing wolverines moving vast distances over varied
terrain and movement of wolverines across the U.S.-Canada border, our
recent assessment of the low levels of trapping mortality in Canada
near the border, and further confirmation of Canada as the source of
wolverine genetics present in contiguous United States wolverines, we
conclude that wolverines in the contiguous United States are not
separated genetically from the larger population in Canada.
Furthermore, even if they were separated genetically, the multiple
generations it would take for genetic isolation to potentially result
in significantly lower genetic diversity and for the deleterious
effects of decreased genetic diversity to then manifest into negative
population-level effects is likely beyond the foreseeable future used
for this determination (38 to 50 years, see Future Condition section
below). As such, we conclude that loss of genetic diversity is not a
stressor for wolverines in the contiguous United States now or within
the foreseeable future.
Small Population Effects: As described above in Population
Abundance and Density, the number of wolverines in the contiguous
United States is relatively small compared to the remainder of the
range in Canada and Alaska, in large part due to limited suitable
habitat and previous persecution and unregulated trapping pressures. As
described above in Genetic Diversity, we now consider wolverines in the
contiguous United States to be genetically connected to wolverines in
Canada (McKelvey et al. 2014; Pilgrim and Schwartz 2018) and wolverines
in the contiguous United States are not separated from the larger North
American wolverine population to the North (Canada and Alaska). In
previous proposed rules and findings, we have discussed small
population
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size as a vulnerability that places wolverines in the contiguous United
States at risk of extirpation. However, those assertions were
predicated on a belief that wolverines in the contiguous United States
were effectively isolated regionally within the United States and
isolated from Canada, thereby increasing the risk of deleterious
genetic effects (countered above in Genetic Diversity) and
susceptibility to stochastic events and limited rescue effect
(migrants) from Canada. With further genetic evidence of the
recolonization of once-extirpated areas of the contiguous United States
by wolverines from Canada post-unregulated trapping over the last
approximately 100 years, history has demonstrated the resiliency of the
North American wolverine population to recover from extreme persecution
and unprecedented direct mortality. We do not currently foresee any
stochastic or catastrophic events that could result in a similar
population-level effect on wolverines in the contiguous United States.
It is no longer accurate to consider contiguous United States
wolverines in isolation from the rest of North American wolverines;
rather, it is more accurate to consider the contiguous United States
wolverines a portion of a much larger and proven resilient North
American wolverine population. We conclude that small population
effects are not a stressor for wolverines in the contiguous United
States now or within the foreseeable future.
Climate Change: In the SSA report, included in our discussion of
future conditions, we provide a summary of current trends related to
observed climate change effects, such as increased temperatures and
changes in precipitation patterns, in areas that encompass the current
potential extent of occurrence for the wolverine. We are not aware of
any adverse effects of these observed changes to the wolverine in the
contiguous United States. The potential effects of future climate
change (projections) are fully considered in our future condition
analysis in the wolverine SSA report (Service 2018, pp. 73-99). We
summarize the results of that analysis in the Future Condition section
of this document below.
Summary of Current Condition
Wolverine populations in much of North America are still recovering
from large losses of individuals from intensive hunting and unregulated
persecution pressures in the late 1880s into the mid-20th century
(Service 2018, p. 104). The distribution of wolverines within suitable
habitat provides a more appropriate method for estimating population
status than using abundance of animals, although there is limited
rangewide survey information. Based on the best available information,
wolverines continue to be detected within suitable habitat within the
western-northwestern contiguous United States including Washington,
Oregon, Idaho, Montana, and Wyoming (Service 2018, p. 71). Studies are
currently under way to provide a better assessment of the species'
current distribution and genetic characteristics of these populations.
The best available information does not indicate the portion of the
North American wolverine population in the contiguous United States is
currently negatively impacted by lower genetic diversity, and there is
no evidence that wolverine numbers in the contiguous United States are
declining.
We prepared a map of the current potential extent of occurrence to
illustrate the species' current distribution in the contiguous United
States (Figure 2). We estimate this area represents approximately 3.5
percent of the wolverine's current potential extent in North America
(Service 2018, p. 71). We determined that 72 percent of our current
potential extent of the wolverine in the contiguous United States is
found on lands owned or managed by the Federal Government (Service
2018, p. 72 and Appendix D). We also evaluated previously modeled
wolverine primary habitat in the contiguous United States (Inman et al.
2013, entire) and estimated that 96 percent of this area is owned or
managed by Federal agencies and 41 percent of this area is located in
designated wilderness areas (Service 2018, p. 72). In our SSA report,
we provide a detailed summary of regulatory mechanisms and conservation
measures affecting wolverines related to State and Federal land
management in the contiguous United States (Service 2018, Appendix G).
We evaluated several potential stressors that may be affecting
wolverine populations or its habitat, including effects from roads,
disturbance due to winter recreation and other activities, effects from
wildland fire, disease and predation, overutilization for (primarily)
commercial purposes, genetic diversity, and small-population effects.
We determined that the effects of roads (evaluated by number of
miles, density, and location) and disturbance represent low-level
stressors to the wolverine in the contiguous United States. Wildland
fire was determined to be a short-term stressor to wolverine habitat
and its prey. Disease and predation, genetic diversity, and small
population size are not considered stressors to the wolverine.
Trapping or hunting of wolverines is currently prohibited in the
contiguous United States. Incidental trapping of wolverines is
infrequent in the contiguous United States and, in Idaho and Montana,
education programs are being implemented to reduce this stressor.
Wolverines are harvested in several Canadian provinces and near the
U.S.-Canada border with management and monitoring oversight based on
spatial and temporal elements. We evaluated historical trapping
information to assess potential impacts to dispersing wolverines into
the United States. Based on the best available commercial and
scientific information, overutilization does not represent a stressor
to the wolverine in the contiguous United States.
We also determined that the wolverines in the contiguous United
States are connected to and an extension of the Canadian population
that is not genetically isolated nor considered a small population that
may be more vulnerable to stressors.
Future Condition
The foreseeable future timeframe evaluated in our SSA analysis is
approximately 38 to 50 years, which captures consideration of the
projected future conditions related to trapping/harvesting, climate
change, or other potential cumulative impacts (Service 2018, p. 73). We
use a timeframe of approximately 38 to 50 years because, beyond this
range, climate modeling uncertainty increases substantially. We believe
this is a reasonable timeframe to consider as it includes the potential
for observing these effects over several generations of the wolverine.
Evaluations of future conditions for species have an inherent level of
uncertainty relative to demographic risks, particularly those related
to climate change projections. After considering the current conditions
for the wolverine and its habitat, we determine that climate change
effects (i.e., significantly elevated temperatures resulting in decline
in snowpack) that may modify suitable habitat, including reproductive
denning habitat, could also change the scope of the wildland fire
stressor and is the most likely future scenario to potentially have an
effect on wolverines at the population level in the contiguous United
States. Based on our review of the best available information, we
determined that there were no other plausible future scenarios that
were likely to have population-level impacts
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to wolverine in the contiguous United States (Service 2018, p. 73). As
described in detail in the wolverine SSA report (Service 2018, pp. 57-
72), the effects of disease, predation, overutilization (trapping),
genetic diversity, small-population effects, and effects of wildland
fire are expected to continue to be at low levels in the future but are
not expected to result in population-level effects to wolverine.
Climate Change Effects
In the wolverine SSA report, we considered climate changes that may
affect environmental conditions upon which the wolverine relies. As
defined by the Intergovernmental Panel on Climate Change (IPCC), the
term ``climate'' refers to the mean and variability of different types
of weather conditions over time (IPCC 2013, p. 1,450). Thus, the term
``climate change'' refers to a change in the mean or the variability of
relevant properties, which persists for an extended period, typically
decades or longer, due to natural conditions (e.g., solar cycles) or
human-caused changes in the composition of atmosphere or in land use
(IPCC 2013a, p. 1,450).
Multiple lines of evidence, not just projections derived from
quantitative models, should be examined when conducting climate
vulnerability assessments (Michalak et al. 2017, entire). Thus, we
evaluated projected effects from climate change in the western United
States relative to both abiotic (e.g., temperature, precipitation, snow
cover) and biotic (e.g., phenology, behavior) factors. Refer to the
wolverine SSA report for a complete discussion of our analysis of the
effects of climate change to wolverine in the contiguous United States
(Service 2018, pp. 73-99). We summarize the results of that analysis
below.
Summary of Future Condition
Abiotic Factors: Observed trends and future climate model
projections indicate warming temperatures for much of the western
United States, including areas within the current potential extent of
the wolverine (Service 2018, pp. 75-81). The degree of future warming
varies by region and is dependent upon the future emission scenario
used during the modeling process. Future precipitation trends are less
certain for many regions, in part, due to naturally high inter-annual
variability; some regions are projected to experience greater winter
precipitation (Service 2018, p. 81). Wolverines have been found to have
a wide range in their physiological critical temperature depending on
season and undergo seasonal changes in fur insulation to adapt to
warmer temperatures in summer (Service 2018, p. 81). Wolverines also
exhibit changes in behavior, such as moving to higher elevations in
summer months (Service 2018, p. 81). Wolverines continue to occupy
areas that have exhibited increases in temperature (e.g., California,
parts of Montana and Washington) due to effects of climate change;
however, no empirical studies have evaluated these physiological and
behavioral adaptations, including sublethal effects, relative to
warming temperatures (Service 2018, p. 81).
Biotic Factors: In addition to evaluating changes in abiotic
factors, biotic interactions should be considered in evaluating
species' response to climate change (reviewed by Post 2013). Although
abiotic changes drive ecological processes, the alterations in biotic
interactions (e.g., competition among conspecifics, interactions with
competitors, resources, and predators) represent the ecological
responses that result from those changes (Post 2013, p. 1). Changes in
certain abiotic factors, such as snow and ice cover, should also be
considered in an ecological context since they represent habitat for
many species (Post 2013, p. 11).
The results presented in the wolverine SSA report indicate biotic
effects resulting from climate change, varying from phenological
changes to shifts in vegetation and vegetation succession (Service
2018, pp. 81-82). We are unaware of studies that have directly
evaluated these types of effects to the North American wolverine or its
habitat. Given the relatively large area and varied habitats occupied
by wolverines in the contiguous United States, the projected shifts in
vegetation are likely to be relatively narrow in scope and scale
relative to potential effects to wolverines. Furthermore, we have no
information to suggest that wolverines selectively use any specific
vegetation type, and some projected changes in vegetation may be
advantageous for wolverine prey (Service 2018, p. 82).
Climate Change and Potential for Cumulative Effects
Threats can work in concert with one another to cumulatively create
conditions that may impact the wolverine or its habitat beyond the
scope of each individual threat (Service 2018, p. 82). Given an
expected increase in temperature in the western United States, the best
available information indicates that, if there are any cumulative
impacts in the future, the most likely population-level effects on
wolverine in the contiguous United States could be: (1) Changes in
snowpack from the combination of increased temperature and changes in
precipitation patterns, or (2) changes in snowpack and increase in
wildland fire potential (Service 2018, p. 83).
Snowpack/Snow Cover: The effects of climate change on snow
persistence has been suggested as an important negative impact on
wolverine habitat and populations by the mid-21st century and was the
primary basis of our 2013 proposed rule to list the North American
wolverine in the contiguous United States (78 FR 7864; February 4,
2013). In light of the court decision remanding our consideration of
our withdrawal of the 2013 proposed rule relative to climate change
effects to wolverine, the Service pursued a refined methodology to
provide insights into the potential impacts of climate change on snow
persistence (Service 2018, p. 85; Ray et al. 2017, entire).
The Service engaged the National Oceanic and Atmospheric
Administration and University of Colorado in Boulder, Colorado, to
evaluate and model fine-scale persistence of snow in occupied and
potential wolverine habitat in the contiguous United States. The
primary objective of this study was to refine spatial and temporal
scale of snow modeling efforts and improve the scientific understanding
of the extent of spring snow retention currently and into the future
under a changing climate (Ray et al. 2017, p. 9). The objectives of the
study included (Ray et al. 2017, p. 10):
Use of fine-scale models to analyze the topographic
effects of snow, including slope and aspect (compass direction that
slope faces).
Use of a range of plausible future climate change
scenarios to assess snow persistence.
Analysis of extremes and year-to-year variability by
selecting representative wet, dry, and near normal years (using
observed conditions) and then modeling changes for those base years
under several future climate scenarios.
Assessment of changes in snow persistence by elevation.
The study was designed to parallel as much as possible and thereby
refine the previous assessment of snow cover persistence in the western
United States presented in McKelvey et al. (2011). However, an exact
replication of the McKelvey et al. (2011) study was not possible given
the time, funding, and computational constraints needed to develop a
fine-scale assessment. The current study was limited to two study
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areas (approximately 1,500 to 3,000 km \2\ (579 to 1,158 mi \2\) each)
in the northern and southern Rocky Mountains (see Service 2018,
Appendix H). These two National Parks bound the Northern and Southern
part of the wolverine historic range, and were selected because they
encompass the latitudinal and elevational range of wolverines within
the contiguous United States. Glacier National Park is representative
of a high-latitude and relatively low-elevation area currently occupied
by wolverines. The Rocky Mountain National Park region is a lower
latitude and higher elevation area within the wolverine's historical
range, which was recently occupied by a wolverine from 2009 to at least
2012. See the wolverine SSA report for a summary of the methods used in
Ray et al. (2017) (Service 2018, pp. 86-87).
Comparison with McKelvey et al. (2011): Although the methods used
in this study have similarities with those presented in McKelvey et al.
(2011), there are several key differences.
Ray et al. (2017) used a finer spatial resolution model
than McKelvey et al. (2011) (0.0625 km\2\ vs. 37 km\2\) (see Service
2018, Appendix I for a comparison figure) that also incorporated slope
and aspect.
The grid cells represented in McKelvey et al. (2011) were
assumed to be flat (i.e., north-facing slopes treated as identical to
south-facing slopes).
McKelvey et al. (2011) focused on May 1 snow depth as a
proxy for May 15 snow disappearance, while Ray et al. (2017) focused
directly on May 15 snow disappearance and produced results for the
presence or absence of deeper snow (nominally greater than or equal to
0.5 m (20 in) depth) on May 1 and April 15. Ray et al. (2017)
originally focused on May 15 to compare to the McKelvey et al. (2011)
study, and June 1 to bracket the snowmelt season. However, the April 15
and April 30 dates were added to the evaluation of snow-covered areas
to align with temporal reproductive patterns of the wolverine (see Use
of Dens and Denning Behavior discussion in the Reproduction and Growth
section of the wolverine SSA report (Service 2018, pp. 23-28)).
Because of the increased resolution of this study, Ray et
al. (2017) were able to consider whether any areas of snow with depth
greater than 0.5 m (20 in) will persist in these areas in the future at
time periods encompassing the end of the wolverine denning period.
Additional comparisons are outlined in the wolverine SSA report
(Service 2018, p. 88; Table 8) and our rationale supporting the use of
snow depth greater than 0.5 m (20 in) is documented in the wolverine
SSA report (Service 2018, p. 87) and in Ray et al. (2017; Table 5-2).
Interpretation of results and additional analysis relative to
wolverine den site scale: Recent studies of wolverine populations and
distribution in Sweden have observed wolverine populations and
reproductive den sites outside areas modeled with persistent spring
snow cover (Aronsson and Persson 2016, p. 266; Persson 2017, pers.
comm.). Another recent study, from Canada, concluded wolverines are
adaptable and do not require large areas of deep spring snowpack for
successful reproduction, and may select small areas covered with deep
snow at a finer scale than can be detected using satellite imagery
(Webb et al. 2016, p. 1,468). Jokinen et al (2019) reported seven
wolverine den sites in hollow mounds (caused by the uplifted root
masses from fallen Black Spruce trees) in the boreal forest of Alberta.
These areas were largely devoid of spring snow cover (mean distance
from dens to nearest spring snow cover was 15.2 km) and the authors
stated wolverines appear to be using ``locally-available denning
structures in the lowland boreal forest, despite a lack of deep snow,
persistent spring snow cover, or large boulders documented in other
studies.'' Regardless as to whether or not wolverines are obligated to
den in areas of deep snow, the Service was interested in exploring the
question, ``If snow cover is required for wolverine denning, will there
be a sufficient amount of significant snow cover in the future in areas
wolverines have historically used for denning in the contiguous United
States?'' The Service integrated future Distributed Hydrology Soil
Vegetation Model projections (2000-2013 averages) of snow-covered area
(greater than 0.5 m (20 in) depth) on May 1 for Glacier National Park
and Rocky Mountain National Park with new information obtained from a
spatial analysis of documented den sites in the contiguous United
States. This analysis indicated 31 of 34 documented den sites in the
contiguous United States were located in areas with slopes less than 25
degrees. Avalanche risk increases significantly in areas with slope
greater than 25 degrees (Scott 2017, pers. comm.) and thus wolverines
maybe avoiding these areas for denning due to this risk (Service 2018,
p. 91).
The Service calculated areal estimates for future snow covered area
in both study sites and limited these estimates to elevation bands
wolverines have used historically for denning and for areas with slopes
less than 25 degrees. This approach resulted in providing the most
conservative estimates of future snow covered area in the areas
wolverines are most likely to use for denning.
Using the projections prepared by Ray et al. (2017), the wolverine
SSA report presents the spatial distribution of significant snow-
covered area with slopes less than 25 degrees and within the elevation
bands expected to be used by wolverines for denning for three future
climate scenarios in each study area (Service 2018, pp. 92-98). The
three scenarios for Glacier National Park and Rocky Mountain National
Park were chosen to span the range of Global Climate Model uncertainty
regarding temperature and precipitation, and by extension significant
snow-covered area (Service 2018, p. 93). A detailed description of
methods describing the process of Global Climate Model selection can be
found in Ray et al. (2017, pp. 35-38). We found that large portions of
the study areas meet all three criteria--greater than 0.5 m (20 in)
snow depth on May 1, at elevation 1,514-2,252 m (4,967-7,389 ft) for
Glacier National Park or 2,700 to 3,600 m (8,858 to 11,811 ft) for
Rocky Mountain National Park, and with slopes less than 25 degrees--
across both study sites in the future (See map legends in Figures 10-15
in the SSA report, (Service 2018, pp. 94-98)).
We also determined that large tracts (several hundred km\2\/mi\2\)
of significant snow (greater than 0.5 m (20 in) in depth) are projected
in close proximity to documented historical den sites across all three
climate scenarios (Service 2018, pp. 94-95). This analysis is limited
to Glacier National Park because this is the only area where new snow-
covered area projections and historical den locations were both
available. Wolverines would not have to travel far, or at all, relative
to either distance or elevation to reach areas with significant snow-
covered area for denning in the future (Service 2018, pp. 94-95).
Based on the best available information, we have no reason to
believe wolverines are confined to previously modeled spring snow
covered areas. Furthermore, there is no quantitative data documenting
spring snow patch size or depth to the denning needs of wolverines.
Even if wolverines must have spring snow for denning, which we do not
believe to be true, the wolverines in the lower 48 will likely have
access to areas with significant spring snow cover in the future. Based
on the new information presented above and in the wolverine SSA report,
we do not believe wolverines need spring snow cover for denning.
Nevertheless, new information suggests that spring
[[Page 64644]]
snow cover will not be a limiting resource for wolverines in the
contiguous United States in the future. Therefore, based on the best
available information, we do not consider the effects of changes in
snowpack from the combination of increased temperature and changes in
precipitation patterns to be a threat to the wolverine.
Wildland Fire
The wolverine SSA report includes a discussion of available
information on the relationship of predicted future climate conditions
on wildland fire projections in the western United States (Service
2018, pp. 99-100). In summary, based on these projections, wildland
fire risk is likely to increase in the western United States, with
future patterns and trends of wildland fire dependent on several
factors (e.g., degree of warming and drought conditions, fuel and soil
moisture, wildland fire management practices, elevation) and geographic
region (Service 2018, p. 100). However, given the diversity of habitats
occupied by wolverines, their occupancy of high elevations, extensive
mobility, and the positive effect wildland fire may have on wolverine
prey species, wildland fire represents a limited stressor, in scope and
scale, to wolverine habitat and its prey in the contiguous United
States range (Service 2018, pp. 63-64).
To summarize, based on the best available information, the
cumulative effects of wildland fire and climate change (e.g., snowpack)
will continue to represent a low impact to the wolverine and its
habitat into the mid-21st century, based on climate change projections
(Service 2018, p. 100).
Other Cumulative Effects
Finally, we note here that the effects of climate change on
snowpack are projected to negatively affect the season lengths for
winter recreational activities, such as skiing and snowmobiling,
shortening the winter recreation season (Service 2018, pp. 100-101). A
shorter winter recreation season would likely decrease the amount of
winter recreation related disturbance occurring in wolverine habitat
and fewer effects to wolverines. Alternatively, even though winter
recreation seasons will be shorter, we could see more winter human
activity at higher elevations due to snow loss at lower elevations.
However, even at current levels, we do not consider winter recreational
activities to be a threat to wolverine in the contiguous United States.
For further discussion of winter recreational activities see the
wolverine SSA report (Service 2018, pp. 100-101).
Summary of Future Conditions
Climate change model projections for the range of the wolverine
within the contiguous United States indicate increases in temperature
by the mid-21st century as compared to early to mid-20th century values
(Service 2018, p. 101). The degree of future warming varies by region;
area specific discussions are included in the SSA report (Service 2018,
pp. 73-80). Precipitation patterns into the future are less clear as
the climate models show significant disagreement in their many regional
projections. Although drought conditions in the western United States
are not unusual, drought duration and intensity have the potential to
be exacerbated by projected temperature increases. Projected
temperature and precipitation changes will affect future snow cover and
the persistence of snow on the landscape.
Snow cover is projected to decline in response to warming
temperatures and changing precipitation patterns, but this varies by
elevation, topography, and by geographic region (Service 2018, p. 101).
Simulations of natural snow accumulation at winter recreation locations
have found that, overall, higher elevation areas (e.g., Rocky
Mountains, Sierra Nevada Mountains) are more resilient to projected
changes in temperature and precipitation as compared to lower
elevations (Wobus et al. 2017, p. 12). In general, models indicate
higher elevations will retain more snow cover than lower elevations,
particularly in early spring (April 30/May 1) (Service 2018, p. 101).
In the wolverine SSA report, we present results from several recent
climate models projecting snowpack declines in the western United
States (Service 2018, pp. 83-100). More specifically, we reviewed a new
analysis (Ray et al. 2017, entire) that modeled future snow persistence
for Glacier and Rocky Mountain National Parks (areas that encompass the
latitudinal and elevational range of the wolverine in the contiguous
United States) at high spatial resolution and at the den-site scale
(Service 2018, pp. 85-98). The results indicate large areas (several
hundred km\2\/mi\2\ for each site) of future snow (greater than 0.5 m
(20 in) in depth) will persist on May 1 (end of the denning season) at
elevations currently used by wolverines for denning (Service 2018, pp.
93-98). This is true, on average, across the range of climate models
used out to approximately year 2055.
Within their North American range, wolverines are found in a
variety of habitats within primarily high-elevation areas of the
western-northwestern United States, and exhibit wide-ranging movements
(Service 2018, p. 102). Wolverines select den sites for differing
characteristics depending on location, and natal den locations are
generally associated with snow cover; however, many natal dens have
been observed outside of the circumpolar boundary of the snow model
presented in Copeland et al. (2010) (Service 2018, p. 103),
particularly in Scandinavia. In addition, reproductive success of
wolverines has not been evaluated relative to the depth and persistence
of snow cover at the den site scale, or in combination with these or
other important key life-history characteristics, including avoidance
and/or protection from predators, prey availability, availability of
food-caching habitat.
We also considered temperature and precipitation projections from
climate change models in conjunction with wildland fire risk. This risk
is likely to increase across the western United States, but patterns
and trends are dependent on several factors (e.g., degree of warming
and drought conditions, fuel and soil moisture) and geographic region
(Service 2018, p. 102) and wildland fire represents a limited stressor,
in scope and scale, to wolverine habitat and prey as described above in
Effects from Wildland Fire.
Overall Assessment
The wolverine's current potential extent of occurrence includes the
western-northwestern United States (see Figure 2), large areas of
Canada, and Alaska (Service 2018, p. 16). The wolverine is found in a
variety of habitats in North America, but generally occurs in high-
elevation, relatively inaccessible locations (Service 2018, p. 102). In
the contiguous United States, potentially suitable habitat (i.e.,
primary habitat), as determined by the physical and ecological features
and the ecological needs of the wolverine, is estimated at 164,125
km\2\ (63,369 mi\2\) (Inman et al. 2013, p. 281). Based on our review
of available relevant literature, we identified the physical and
ecological needs of the species as follows: large territories in
relatively inaccessible landscapes, at high elevation (1,800 to 3,500 m
(5,906 to 11,483 ft)) within the contiguous United States; access to a
variety of food resources, that varies with seasons; and reproductive
behavior linked to both temporal and physical features (Service 2018,
p. 104). These needs are currently met for wolverines in the contiguous
United States and are expected to be met in the future (i.e., in 38-50
years) (Service 2018, p. 104).
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We recognize there is limited information available for the
wolverine, including population estimates and abundance trends. In the
contiguous United States, the structure of the wolverine population is
represented as a metapopulation, although its genetic structure
relative to its entire North American range has not been
comprehensively evaluated (Service 2018, p. 102). Wolverine populations
in Alaska are considered to be continuous with populations in the Yukon
and British Columbia provinces of Canada based on genetic studies
(COSEWIC 2014, p. 37). Similarly, studies of wolverines in the North
Cascades region have documented recent movement of wolverines from
Washington into British Columbia (Aubry et al. 2016, pp. 16, 20) and
from Idaho (Lucid et al. 2016, p. 184) to British Columbia, and earlier
from Montana to British Columbia and Alberta (e.g., Newby and Wright
1955, p. 252).
We present in our SSA report a detailed discussion of wolverine
reproductive behavior. Based on the best available information,
wolverines select den sites for different characteristics depending on
location. Dens located under snow cover may be related to wolverine
distribution based on other life-history traits, including
morphological, demographic, and behavioral adaptations that allow them
to successfully compete for food resources (Inman 2013, pers. comm.).
Structure (e.g., uprooted trees, boulders and talus fields) appears to
be an important requirement for natal den sites. However, reproductive
success of wolverines has not been evaluated relative to the depth and
persistence of snow cover, or in combination with these or other
important characteristics, including prey availability and predator
avoidance. Recent studies of wolverine populations and distribution in
Sweden have observed wolverine populations and reproductive den sites
outside areas modeled with persistent spring snow cover (Aronsson and
Persson 2016; Persson 2017, pers. comm.). Another recent study
concluded that wolverines are adaptable and do not require large areas
of deep spring snowpack for successful reproduction, and may select
small areas covered with deep snow at a finer scale than can be
detected using satellite imagery (Webb et al. 2016, p. 1,468). Most
recently, wolverine dens have been documented in boreal Alberta,
Canada, several kilometers away from spring snow cover, in hollow
mounds caused by fallen spruce trees (Jokinen et al, 2019). We would
not expect fallen trees, and the potentially suitable denning sites
created by them, to be a limiting resource in wolverine habitat.
We identified several potential stressors that may be affecting the
species and its habitat currently or in the future, including impacts
associated with climate change effects. Based on the best available
information, demographic risks to the species from either known or most
likely potential stressors (i.e., disturbance due to winter
recreational activities, other human disturbances, effects of wildland
fire, disease, predation, overutilization, genetic diversity, small-
population effects, climate change, and cumulative effects) are low
based on our evaluation of the best available information as it applies
to current and potential future conditions for the wolverine and in the
context of the attributes that affect the needs of the species (Service
2018, p. 103).
Climate change model projections for the range of the wolverine
within the contiguous United States indicate increases in temperature
by the mid-21st century as compared to early to mid-20th century values
(Service 2018, p. 103). Our evaluation of climate change indicates that
snow cover is projected to decline in response to warming temperatures
and changing precipitation patterns, but this varies by elevation,
topography, and by geographic region (Service 2018, p. 103). In
general, models indicate higher elevations will retain more snow cover
than lower elevations, particularly in early spring (April 30/May 1)
(Service 2018, p. 103). Although the persistence of spring snow has not
yet been determined to be critical to wolverine survival in North
America, our review of projected snow persistence (to approximately
2055) within the Northern and Southern Rocky Mountains, indicates
several hundred km\2\/mi\2\ of deep snow will persist on May 1 at
elevations used by the wolverine for denning (Service 2018, p. 103).
Legal protections of the wolverine in the contiguous United States
include State listing in California and Oregon (as threatened);
Colorado (as endangered); candidate species status in Washington;
protected as a non-game species in Idaho and Wyoming; a species of
concern and furbearer with a closed season in Montana; and protected
from collection, importation, and possession in Utah (Service 2018, p.
107). Trapping or hunting of wolverines is currently prohibited in the
contiguous United States. Trapping effort along the U.S.-Canada border
does not represent a barrier to wolverine movement and dispersal along
the international border (Service 2018, p. 103).
Management actions for conservation of the wolverine and its
habitat are included within State Wildlife Action Plans, the Management
Plan for the Conservation of Wolverines in Idaho (IDFG 2014), and USDA
Forest Service Land and Resource Management Plans (Service 2018,
Appendix G). Various provisions of these plans include, but are not
limited to, winter road closures, fire management, and land acquisition
or conservation easements. These management measures, currently and in
the future, will alleviate effects associated with potential impacts
related to stressors. However, we do not rely on the management
measures and conservation efforts contained in these plans to support
our listing decision. In addition, the WAFWA Wildlife Chiefs Wolverine
Subcommittee is providing a forum for western States to work
collaboratively with each other and with the Service and other partners
for conserving wolverines found in the western-northwestern United
States, and, to date, approximately $1.5 million of funding has been
applied towards conservation and management actions for the wolverine
(e.g., Western States Wolverine Conservation Project) (McDonald 2017,
pers. comm.).
Determination of Species Status
Section 4 of the Act (16 U.S.C. 1533) and its implementing
regulations (50 CFR part 424) set forth the procedures for determining
whether a species meets the definition of ``endangered species'' or
``threatened species.'' The Act defines an ``endangered species'' as a
species that is ``in danger of extinction throughout all or a
significant portion of its range,'' and a ``threatened species'' as a
species that is ``likely to become an endangered species within the
foreseeable future throughout all or a significant portion of its
range.'' The Act requires that we determine whether a species meets the
definition of ``endangered species'' or ``threatened species'' because
of any of the following factors:
(A) The present or threatened destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial, recreational, scientific, or
educational purposes;
(C) Disease or predation;
(D) The inadequacy of existing regulatory mechanisms; or
(E) Other natural or manmade factors affecting its continued
existence.
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Determination of Status Throughout All of Its Range in the Contiguous
United States
Since the publication of the February 4, 2013, proposed rule (78 FR
7864) and reinstatement of that proposed rule on October 18, 2016 (81
FR 71670), we prepared a comprehensive assessment of the current and
future status of wolverines in the contiguous United States as
presented in the wolverine SSA report (Service 2018, entire). New
information from recent surveys and a reevaluation of the species'
current range, new genetic information, new studies of wolverine
reproductive behavior and denning habitat, and results from detailed
modeling of future spring snow persistence are included in the
wolverine SSA report and contribute to our current understanding of the
species. The wolverine SSA report also provides a comprehensive summary
of wolverine life history and ecology, including an assessment of
wolverine physiology, and an analysis of new information on wolverine
trapping pressure in Canada near the United States-Canada border, as
well as analyses of new information relevant to other potential threats
to the species. We have carefully assessed the best scientific and
commercial information available regarding the past, present, and
future threats to North American wolverines in the contiguous United
States including effects from roads (Factors A and E); disturbance due
to winter recreational activity (Factors A and E); other human
disturbance (Factors A and E); effects from wildland fire (Factor A);
disease (Factor C); predation (Factor C); overutilization (trapping)
(Factor B); genetic diversity (Factor E); small-population effects
(Factor E); and climate change (Factors A and E). We also assessed the
adequacy of existing regulatory mechanisms (Factor D).
Consideration of Cumulative Effects--Threats can work in concert
with one another to cumulatively create conditions that may impact the
wolverine or its habitat beyond the scope of each individual threat.
See the Climate Change and Potential for Cumulative Effects section
above and the wolverine SSA report for an in-depth analysis of
cumulative effects (Service 2018, pp. 82-101). We note that by using
the SSA framework to guide our analysis of the scientific information
documented in the SSA report, we have not only analyzed individual
effects on the species, but we have also analyzed their potential
cumulative effects. We incorporate the cumulative effects into our SSA
analysis when we characterize the current and future condition of the
species. Our assessment of the current and future conditions
encompasses and incorporates the threats individually and cumulatively.
Because the SSA framework considers not just the presence of the
factors, but to what degree they collectively influence risk to the
entire species, our assessment integrates the cumulative effects of the
factors.
Our future-condition analysis in the wolverine SSA report includes
the potential conditions that the species or its habitat may face, that
is, the most probable scenario if those conditions are realized in the
future. This most probable scenario includes consideration of the
sources that have the potential to most likely impact the species at
the population or rangewide scales in the future, including potential
cumulative impacts. Given an expected increase in temperature in the
western United States, the best available information indicates that,
if there are any cumulative impacts in the future, the most likely to
have population-level effects on wolverine in the contiguous United
States could be: (1) Changes in snowpack from the combination of
increased temperature and changes in precipitation patterns, or (2)
changes in snowpack and increase in wildland fire potential (Service
2018, p. 83). The best available information does not indicate that the
effects of trapping and mortality from roads will act cumulatively with
effects of climate change, and those stressors are expected to remain
low-level impacts into the future. We provide a detailed analysis of
climate change and the potential for cumulative effects in the
wolverine SSA report (Service 2018, pp. 82-102). Based on the best
available information, the cumulative effects of wildland fire and
climate change (e.g., snowpack) will continue to represent a low impact
to the wolverine and its habitat into the mid-21st century, based on
climate change projections.
Resilience, Representation, and Redundancy--In order to
characterize a species' viability and demographic risks, we consider
the concepts of resilience, representation, and redundancy. We also
consider known and potential stressors that may negatively impact the
physical and biological features that the species needs for survival
and reproduction. Stressors are expressed as risks to its demographic
features such as abundance, population and spatial structure, and
genetic or ecological diversity. We consider the level of impact a
stressor may have on a species along with the consideration of
demographic factors (e.g., whether a species has stable, increasing, or
decreasing trends in abundance, population growth rates, diversity of
populations, and loss or degradation of habitat).
Wolverine populations in much of North America are still recovering
from large losses of individuals from unregulated hunting and
persecution pressures in the late 1880s into the mid-20th century
(Service 2018, p. 104). Surveys conducted in the winter of 2015-2016
and 2016-2017 continue to document its presence within portions of its
historical range in the western contiguous United States
(representation) (Service 2018, p. 104).
Redundancy, the ability to rebound after stochastic perturbation,
can be characterized by the distribution and connectivity of
populations. In considering wolverine in the contiguous United States,
individuals are found in alpine, boreal, and subalpine habitats, with
breeding populations in four western States. Additionally, wolverines
in the contiguous United States are connected to wolverine populations
in Canada along the U.S.-Canada border, which contributes to current
and future redundancy (Service 2018, p. 104).
Resiliency, the ability to withstand stochastic events, can be
characterized by numbers of individuals and abundance trends. As
indicated above, actual current population size, growth rate, and
current population trends are unknown for wolverines in the contiguous
United States due to the lack of abundance information. However,
according to recent estimates, Canada's western subpopulation (which is
connected to wolverines in the contiguous United States) has been
estimated at 15,688 to 23,830 adult wolverines, with expansion of
wolverines into historically occupied areas in both Canada and the
contiguous United States with movement across both international
borders (Service 2018, pp. 54, 105). The 2014 Committee on the Status
of Endangered Wildlife in Canada report concluded that a climate-driven
decline in wolverine populations in North America is not evident at
this time in much of its range (COSEWIC 2014, p. 22). Wolverine
populations in Canada are considered stable (Service 2018, p. 105). We
also note that density estimates indicate no declining trend in
wolverine populations in Alaska (Service 2018, p. 105). We recognize
that there is limited information on populations (representation) or
genetic diversity (resiliency and representation) for the wolverine in
the contiguous United States, and no comprehensive studies to indicate
what a viable (or
[[Page 64647]]
minimal) wolverine population size should be across its North American
range. However, the best available information does not indicate either
increasing or declining numbers of the wolverine in North America,
including the contiguous United States. Further, at this time, the best
available information does not indicate that the species' abundance is
significantly impacted by the stressors evaluated (singly or
cumulatively), and this situation is unlikely to change in the future,
supporting current and future resiliency.
As discussed in the wolverine SSA report, both direct and
cumulative effects of climate change (e.g., higher temperatures, loss
of snow cover, wildland fire) may affect the resilience of the
wolverine in the future by creating an environment that is less
favorable to its physiological and ecological needs (Service 2018, p.
105). We are unaware of studies of the wolverine that have formally
evaluated the species' responses (e.g., reproductive success or
survival) to warming temperatures or other climate change effects.
As described in the wolverine SSA report, the best available
information indicates confirmed observations of wolverines denning in
areas with patchy snow cover in Alaska, Canada, and Scandinavia
(Service 2018, p. 105). Further, using fine-scale snow modeling, we
estimated that large areas of spring snow (May 1) will remain within
Glacier National Park, where wolverines are known to den (Service 2018,
p. 105). Given their high rate of movement, large dispersal distances,
including travel through areas not covered with snow, and other life-
history traits (e.g., behavioral plasticity) observed in wolverines, we
do not predict a significant loss of individual and population
resiliency to the species in the future (i.e., 38-50 years) within its
North America range, including the contiguous United States (Service
2018, p. 105).
Currently, we are unaware of any documented specific risks for the
wolverine related to a substantial change or loss of diversity in life-
history traits, population demographics, morphology, behavior, or
genetic characteristics that can be used to characterize species
representation (the ability to adapt to change). Rates of dispersal or
gene flow are not known to have changed, and recent evidence supports
continued connectivity with contiguous United States wolverines and
wolverines in Canada. Additionally, there is no currently available
information to indicate that the current abundance of the wolverine
across its current potential extent in the contiguous United States is
at a level that is causing inbreeding depression or that loss of
genetic variation is affecting representation or that would affect
representation in the future (Service 2018, p. 105). Nor is there any
information to indicate that this species is unable to adapt or adjust
to changing conditions (e.g., potential reduction in snow cover). We do
not expect a reduction in representation of the wolverines in the
contiguous United States in the future. We have determined that the
needs of the species are provided within the contiguous United States
currently and into the future. Thus, after assessing the best available
information, we conclude that the North American wolverine in the
contiguous United States is not in danger of extinction throughout all
of its range (endangered) nor is it likely to become so in the
foreseeable future (threatened).
Because we determined that the North American wolverine in the
contiguous United States is not in danger of extinction or likely to
become so in the foreseeable future throughout all of its range, we
will consider whether there are any significant portions of its range
in which it is in danger of extinction or likely to become so in the
foreseeable future.
Determination of Status Throughout a Significant Portion of Its Range
Under the Act and our implementing regulations, a species may
warrant listing if it is in danger of extinction or likely to become so
in the foreseeable future throughout all or a significant portion of
its range. Having determined that the wolverine is not in danger of
extinction or likely to become so in the foreseeable future throughout
all of its range, we now consider whether it may be in danger of
extinction or likely to become so in the foreseeable future in a
significant portion of its range--that is, whether there is any portion
of the species' range for which it is true that both (1) the portion is
significant; and, (2) the species is in danger of extinction now or
likely to become so in the foreseeable future in that portion.
Depending on the case, it might be more efficient for us to address the
``significance'' question or the ``status'' question first. We can
choose to address either question first. Regardless of which question
we address first, if we reach a negative answer with respect to the
first question that we address, we do not need to evaluate the other
question for that portion of the species' range.
In undertaking this analysis for the North American wolverine in
the contiguous United States, we choose to address the status question
first--we consider information pertaining to the geographic
distribution of both the species and the threats that the species faces
to identify any portions of the range where the species is endangered
or threatened.
For the North American wolverine in the contiguous United States,
we considered whether the threats are geographically concentrated in
any portion of the species' range at a biologically meaningful scale.
We examined the following threats: effects from roads, disturbance due
to winter recreational activity, other human disturbance, wildland
fire, disease, predation, overutilization (trapping), genetic
diversity, small-population effects, climate change, and cumulative
impacts of these potential threats (Service 2018, entire). All of these
potential stressors are relatively evenly distributed geographically
throughout the range of the wolverine in the contiguous United States.
We found no concentration of threats in any portion of the wolverine's
range at a biologically meaningful scale. Therefore, no portion of the
species' range can provide a basis for determining that the species is
in danger of extinction now or likely to become so in the foreseeable
future in a significant portion of its range, and we find the species
is not in danger of extinction now or likely to become so in the
foreseeable future in any significant portion of its range. This is
consistent with the courts' holdings in Desert Survivors v. Department
of the Interior, No. 16-cv-01165-JCS, 2018 WL 4053447 (N.D. Cal. Aug.
24, 2018), and Center for Biological Diversity v. Jewell, 248 F. Supp.
3d, 946, 959 (D. Ariz. 2017).
Determination of Status
We have reviewed the best available scientific and commercial
information regarding the past, present, and future threats to the
North American wolverine in the contiguous United States and we have
determined that, if it were to be a listable entity, it does not meet
the definition of an endangered species or a threatened species in
accordance with sections 3(6) and 3(20) of the Act. Furthermore, we
have determined that the population of wolverines in the contiguous
United States is not a DPS. As a consequence of these determinations,
we are withdrawing our proposed rule to list the distinct population
segment of the North American wolverine occurring in the contiguous
United States as a threatened species.
[[Page 64648]]
References Cited
A complete list of references cited in this document and the
wolverine SSA report are available on the internet at https://www.regulations.gov at Docket No. FWS-R6-ES-2016-0106 and upon request
from the Montana Ecological Services Office (see FOR FURTHER
INFORMATION CONTACT).
Authors
The primary authors of this document are the staff members of the
Montana Ecological Services Office and the Mountain-Prairie Regional
Office.
Signing Authority
The Director, U.S. Fish and Wildlife Service, approved this
document and authorized the undersigned to sign and submit the document
to the Office of the Federal Register for publication electronically as
an official document of the U.S. Fish and Wildlife Service. Aurelia
Skipwith, Director, U.S. Fish and Wildlife Service, approved this
document on August 10, 2020, for publication.
Authority
The authority for this action is the Endangered Species Act of
1973, as amended (16 U.S.C. 1531 et seq.).
Madonna Baucum,
Regulations and Policy Chief, Division of Policy, Economics, Risk
Management, and Analytics of the Joint Administrative Operations, U.S.
Fish and Wildlife Service.
[FR Doc. 2020-19538 Filed 10-9-20; 8:45 am]
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