[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withdrawn on October 13, 2020.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 http://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 http://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
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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 (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
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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 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 http://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
[[Page 64641]]
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
[[Page 64642]]
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
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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).
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
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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 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 http://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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