Endangered and Threatened Wildlife and Plants; Revised 12-Month Finding on a Petition To List the Upper Missouri River Distinct Population Segment of Arctic Grayling as an Endangered or Threatened Species, 49383-49422 [2014-19353]
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Department of the Interior
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Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Revised 12-Month Finding
on a Petition To List the Upper Missouri River Distinct Population Segment
of Arctic Grayling as an Endangered or Threatened Species; Proposed
Rule
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R6–ES–2013–0120;
4500030113]
Endangered and Threatened Wildlife
and Plants; Revised 12-Month Finding
on a Petition To List the Upper
Missouri River Distinct Population
Segment of Arctic Grayling as an
Endangered or Threatened Species
AGENCY:
Fish and Wildlife Service,
Interior.
Notice of 12-month petition
finding.
ACTION:
We, the U.S. Fish and
Wildlife Service (Service), announce a
revised 12-month finding on a petition
to list the Upper Missouri River distinct
population segment (Upper Missouri
River DPS) of Arctic grayling
(Thymallus arcticus) as an endangered
or threatened species under the
Endangered Species Act of 1973, as
amended (Act). After review of the best
available scientific and commercial
information, we find that listing the
Upper Missouri River DPS of Arctic
grayling is not warranted at this time.
The best available scientific and
commercial information indicates that
habitat-related threats previously
identified, including habitat
fragmentation, dewatering, thermal
stress, entrainment, riparian habitat
loss, and effects from climate change,
for the Upper Missouri River DPS of
Arctic grayling have been sufficiently
ameliorated and that 19 of 20
populations of Arctic grayling are either
stable or increasing. This action
removes the Upper Missouri River DPS
of the Arctic grayling from our
candidate list. Although listing is not
warranted at this time, we ask the
public to submit to us any new
information that becomes available
concerning the threats to the Upper
Missouri River DPS of Arctic grayling or
its habitat at any time.
DATES: The finding announced in this
document was made on August 20,
2014.
SUMMARY:
This finding is available on
the Internet at https://
www.regulations.gov at Docket Number
FWS–R6–ES–2013–0120. Supporting
documentation we used in preparing
this finding is available for public
inspection, by appointment, during
normal business hours at the U.S. Fish
and Wildlife Service, Montana
Ecological Services Office, 585 Shepard
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ADDRESSES:
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Way, Suite 1, Helena, MT 59601. Please
submit any new information, materials,
comments, or questions concerning this
finding to the above street address.
FOR FURTHER INFORMATION CONTACT: Jodi
Bush, Field Supervisor, Montana
Ecological Services Office (see
ADDRESSES); telephone 406–449–5225. If
you use a telecommunications device
for the deaf (TDD), please call the
Federal Information Relay Service
(FIRS) at 800–877–8339.
SUPPLEMENTARY INFORMATION:
Background
Section 4(b)(3)(B) of the Act (16
U.S.C. 1531 et seq.) requires that, for
any petition to revise the Federal Lists
of Endangered and Threatened Wildlife
and Plants that contains substantial
scientific or commercial information
that listing the species may be
warranted, we make a finding within 12
months of the date of receipt of the
petition. In this finding, we will
determine that the petitioned action is:
(1) Not warranted, (2) warranted, or (3)
warranted, but the immediate proposal
of a regulation implementing the
petitioned action is precluded by other
pending proposals to determine whether
species are endangered or threatened,
and expeditious progress is being made
to add or remove qualified species from
the Federal Lists of Endangered and
Threatened Wildlife and Plants. We
must publish these 12-month findings
in the Federal Register.
Previous Federal Actions
We have published a number of
documents on Arctic grayling since
1982, and have been involved in
litigation over previous findings. We
describe previous federal actions that
are relevant to this document below.
We published our first status review
for the Montana Arctic grayling
(Thymallus arcticus montanus), then
thought to be a subspecies of Arctic
grayling, in a Federal Register
document on December 30, 1982 (47 FR
58454). In that document, we designated
the purported subspecies, Montana
Arctic grayling, as a Category 2 species.
At that time, we designated a species as
Category 2 if a listing as endangered or
threatened was possibly appropriate,
but we did not have sufficient data to
support a proposed rule to list the
species.
On October 9, 1991, the Biodiversity
Legal Foundation and George
Wuerthner petitioned us to list the
fluvial (riverine) populations of Arctic
grayling in the Upper Missouri River
basin as an endangered species
throughout its historical range in the
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coterminous United States. We
published a notice of a 90-day finding
in the January 19, 1993, Federal
Register (58 FR 4975), concluding the
petitioners presented substantial
information indicating that listing the
fluvial Arctic grayling of the Upper
Missouri River in Montana and
northwestern Wyoming may be
warranted. This finding also noted that
taxonomic recognition of the Montana
Arctic grayling (Thymallus arcticus
montanus) as a subspecies (previously
designated as a category 2 species) was
not widely accepted, and that the
scientific community generally
considered this population a
geographically isolated member of the
wider species (T. arcticus).
On July 25, 1994, we published
notification of a 12-month finding in the
Federal Register (59 FR 37738),
concluding that listing the DPS of
fluvial Arctic grayling in the Upper
Missouri River was warranted but
precluded by other higher priority
listing actions. This DPS determination
predated our DPS policy (61 FR 4722,
February 7, 1996), so the entity did not
undergo a DPS analysis as described in
the policy. The 1994 finding placed
fluvial Arctic grayling of the Upper
Missouri River on the candidate list and
assigned it a listing priority of 9,
indicating that the threats were
imminent but of moderate to low
magnitude.
On May 31, 2003, the Center for
Biological Diversity and Western
Watersheds Project (Plaintiffs) filed a
complaint in U.S. District Court in
Washington, DC, challenging our 1994
‘‘warranted but precluded’’
determination for the DPS of fluvial
Arctic grayling in the Upper Missouri
River basin. On May 4, 2004, we
elevated the listing priority number of
the fluvial Arctic grayling to 3 (69 FR
24881), indicating threats that were
imminent and of high magnitude. On
July 22, 2004, the Plaintiffs amended
their complaint to challenge our failure
to emergency list this population. We
settled with the Plaintiffs in August
2005, and we agreed to submit a revised
determination on whether this
population warranted listing as
endangered or threatened to the Federal
Register on or before April 16, 2007.
On April 24, 2007, we published a
revised 12-month finding on the
petition to list the Upper Missouri River
DPS of fluvial Arctic grayling (72 FR
20305) (‘‘2007 finding’’). In this finding,
we determined that fluvial Arctic
grayling of the upper Missouri River did
not constitute a species, subspecies, or
DPS under the Act. Therefore, we found
that the upper Missouri River
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population of fluvial Arctic grayling was
not a listable entity under the Act, and,
as a result, listing was not warranted.
With that document, we withdrew the
fluvial Arctic grayling from our
candidate list.
On November 15, 2007, the Center for
Biological Diversity, Federation of Fly
Fishers, Western Watersheds Project,
George Wuerthner, and Pat Munday
filed a complaint (CV–07–152, in the
District Court of Montana) to challenge
our 2007 finding. We settled this
litigation on October 5, 2009. In the
stipulated settlement, we agreed to: (a)
Publish, on or before December 31,
2009, a document in the Federal
Register soliciting information on the
status of the upper Missouri River
Arctic grayling; and (b) submit, on or
before August 30, 2010, a new 12-month
finding for the upper Missouri River
Arctic grayling to the Federal Register.
On October 28, 2009, we published in
the Federal Register a notice of intent
to conduct a status review of Arctic
grayling (Thymallus arcticus) in the
upper Missouri River system (74 FR
55524). To ensure the status review was
based on the best available scientific
and commercial data, we requested
information on the taxonomy, biology,
ecology, genetics, and population status
of the Arctic grayling of the upper
Missouri River system; information
relevant to consideration of the
potential DPS status of Arctic grayling
of the upper Missouri River system;
threats to the species; and conservation
actions being implemented to reduce
those threats in the upper Missouri
River system. That document further
specified that the status review might
consider various DPS designations that
include different life histories of Arctic
grayling in the upper Missouri River
system and different DPS
configurations, including fluvial,
adfluvial (lake populations), or all life
histories of Arctic grayling in the upper
Missouri River system.
On September 8, 2010, we published
a revised 12-month finding on the
petition to list the Upper Missouri River
DPS of Arctic grayling (75 FR 54708)
(‘‘2010 finding’’). In this finding, we
determined that fluvial and adfluvial
Arctic grayling of the upper Missouri
River did constitute a DPS under the
Act. Further, we found that a DPS
configuration including both adfluvial
and fluvial life histories was the most
appropriate for the long-term
conservation of Arctic grayling because
genetic evidence indicated that fluvial
and adfluvial life-history forms did not
represent distinct evolutionary lineages.
We concluded by finding that the Upper
Missouri River DPS of Arctic grayling
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was warranted for listing under the Act,
but precluded by other higher priority
listing actions.
On September 9, 2011, we reached an
agreement with plaintiffs in Endangered
Species Act Section 4 Deadline Litig.,
Misc. Action No. 10–377 (EGS), MDL
Docket No. 2165 (D. D.C.) (known as the
‘‘MDL case’’) on a schedule to publish
proposed listing rules or not-warranted
findings for the species on our
candidate list. This agreement
stipulated that we would submit for
publication in the Federal Register
either a proposed listing rule for the
Upper Missouri River DPS of Arctic
grayling, or a not-warranted finding, no
later than the end of Fiscal Year 2014.
On November 26, 2013, we published
a document in the Federal Register (78
FR 70525) notifying the public that we
were initiating a status review of the
Upper Missouri River DPS of Arctic
grayling to determine whether the entity
meets the definition of an endangered or
threatened species under the Act. That
document requested general information
(taxonomy, biology, ecology, genetics,
and status) on the Arctic grayling of the
upper Missouri River system, as well as
information on the conservation status
of, threats to, planned and ongoing
conservation actions for, habitat
selection of, habitat requirements of,
and considerations concerning the
possible designation of critical habitat
for the Arctic grayling of the upper
Missouri River system.
This document constitutes a revised
12-month finding (‘‘2014 finding’’) on
whether to list the Upper Missouri River
DPS of Arctic grayling (Thymallus
arcticus) as endangered or threatened
under the Act, and fulfills our
commitments under the MDL case.
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For more detail on taxonomy and
species description, see the 2010 finding
(75 FR 54708).
Distribution
Arctic grayling are native to Arctic
Ocean drainages of Alaska and
northwestern Canada, as far east as
Hudson’s Bay, and westward across
northern Eurasia to the Ural Mountains
(Scott and Crossman 1998, pp. 301–302;
Froufe et al. 2005, pp. 106–107; Weiss
et al. 2006, pp. 511–512). In North
America, they are native to northern
Pacific Ocean drainages as far south as
the Stikine River in British Columbia
(Nelson and Paetz 1991, pp. 253–256;
Behnke 2002, pp. 327–331).
For a full discussion on the global
distribution of Arctic grayling, see the
2010 finding (75 FR 54709–54710).
Here, we focus on the distribution of
Arctic grayling within the conterminous
United States.
Distribution in the Conterminous
United States
Two disjunct groups of Arctic
grayling were native to the
conterminous United States: One in the
upper Missouri River basin in Montana
and Wyoming (currently extant only in
Montana); and another in Michigan that
was extirpated in the late 1930s (Hubbs
and Lagler 1949, p. 44), and has not
been detected since.
During the status review process, the
Service received information indicating
that Arctic grayling may have also been
native to areas outside the Upper
Missouri River basin in Montana and
Wyoming. This information included
multiple historical newspaper clippings
and several reports from early Army
expeditions purporting that Arctic
grayling were captured in the
Yellowstone River drainage in Montana
Species Information
and the Snake River drainage in Idaho
Taxonomy and Species Description
(Shea 2014, entire). Some of these
reports even included descriptions of
The Arctic grayling (Thymallus
arcticus) is a fish belonging to the family captured fish. However, none of the
descriptions mentions the colorful, sailSalmonidae (salmon, trout, charr,
like dorsal fin of Arctic grayling, a
whitefishes), subfamily Thymallinae
prominent feature that clearly
(graylings), and it is represented by a
single genus, Thymallus. Arctic grayling distinguishes Arctic grayling from other
salmonids. In addition, a similar species
have elongate, laterally compressed,
resembling Arctic grayling (i.e.,
trout-like bodies with deeply forked
mountain whitefish) is native to both
tails, and adults typically average 300–
the Yellowstone River drainage and
380 millimeters (mm) (12–15 inches
Snake River drainage. Mountain
(in.)) in length. Coloration can be
whitefish were sometimes referred to as
striking, and varies from silvery or
‘‘grayling’’ in some areas of the West
iridescent blue and lavender, to dark
(Ellis 1914, p. 75). Thus, it is likely that
blue (Behnke 2002, pp. 327–328). A
early reports of Arctic grayling
prominent morphological feature of
Arctic grayling is the sail-like dorsal fin, occurring outside the upper Missouri
River basin were mountain whitefish
which is large and vividly colored with
rows of orange to bright green spots, and misidentified as Arctic grayling.
Therefore, without information to the
often has an orange border (Behnke
contrary, we consider Arctic grayling to
2002, pp. 327–328).
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be native only to the upper Missouri
River basin in Montana and Wyoming
and to Michigan.
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Native Distribution of Arctic Grayling in
the Upper Missouri River Basin
The first Euro-American ‘‘discovery’’
of Arctic grayling in North America is
attributed to members of the Lewis and
Clark Expedition, who encountered the
species in the Beaverhead River in
August 1805 (Nell and Taylor 1996, p.
133). Vincent (1962, p. 11) and Kaya
(1992, pp. 47–51) synthesized accounts
of Arctic grayling occurrence and
abundance from historical surveys and
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contemporary monitoring to determine
the historical distribution of the species
in the upper Missouri River system
(Figure 1). We base our conclusions on
the historical distribution of Arctic
grayling in the upper Missouri River
basin on these two reviews. Arctic
grayling were widely but irregularly
distributed in the upper Missouri River
system above the Great Falls in Montana
and in northwest Wyoming within the
present-day location of Yellowstone
National Park (Vincent 1962, p. 11).
They were estimated to inhabit up to
2,000 kilometers (km) (1,250 miles (mi))
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of stream habitat until the early 20th
century (Kaya 1992, pp. 47–51). Arctic
grayling were reported in the mainstem
Missouri River, as well as in the Smith,
Sun, Jefferson, Madison, Gallatin, Big
Hole, Beaverhead, and Red Rock Rivers
(Vincent 1962, p. 11; Kaya 1992, pp. 47–
51; USFWS 2007; 72 FR 20307, April
24, 2007). Anecdotal accounts report
that the species may have been present
in the Ruby River, at least seasonally
(Magee 2005, pers. comm.), and were
observed there as recently as the early
1970s (Holton, undated).
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Fluvial Arctic grayling were
historically widely distributed in the
upper Missouri River basin, but a few
adfluvial populations also were native
to the basin. For example, Arctic
grayling are native to Red Rock Lakes,
in the Centennial Valley (Vincent 1962,
pp. 112–121; Kaya 1992, p. 47). Vincent
(1962, p. 120) stated that Red Rock
Lakes were the only natural lakes in the
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upper Missouri River basin accessible to
colonization by Arctic grayling, and
concluded that Arctic grayling there
were the only native adfluvial
population in the basin. However,
Arctic grayling were also native to Elk
Lake (in the Centennial Valley; Kaya
1990, p. 44) and a few small lakes in the
upper Big Hole River drainage, based on
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recent genetic information (Peterson and
Ardren 2009, p. 1768).
The distribution of native Arctic
grayling in the upper Missouri River
went through a dramatic reduction in
the first 50 years of the 20th century,
especially in riverine habitats (Vincent
1962, pp. 86–90, 97–122, 127–129; Kaya
1992, pp. 47–53). The native
populations that formerly resided in the
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Smith, Sun, Jefferson, Beaverhead,
Gallatin, and mainstem Missouri Rivers
are considered extirpated, and the only
remaining native fluvial population is
found in the Big Hole River and some
of its tributaries (Kaya 1992, pp. 51–53).
The fluvial form currently occupies less
than 10 percent of its historical range in
the Missouri River system (Kaya 1992,
p. 51). Other native populations in the
upper Missouri River occur in two
small, headwater lakes in the upper Big
Hole River system (Miner and
Mussigbrod Lakes); the upper Ruby
River (recently reintroduced from Big
Hole River stock); the Madison River
upstream from Ennis Reservoir; Elk
Lake in the Centennial Valley (recently
reintroduced from Red Rock Lakes
stock); and the Red Rock Lakes in the
Centennial Valley (Everett 1986, p. 7;
Kaya 1992, p. 53; Peterson and Ardren
2009, pp. 1762, 1768; see Figure 1).
Introduced Lake-Dwelling Arctic
Grayling in the Upper Missouri River
Basin
From 1898 through the 1960s, an
estimated 100 million Arctic grayling
were stocked across Montana and other
western States. The sources of these
stockings varied through time as
different State, Federal, and private
hatchery operations were created, but
the ultimate source for all hatcheries in
Montana appears to be stock from two
Montana populations: Centennial Valley
and Madison River (Peterson and
Ardren 2009, p. 1767; Leary 2014,
unpublished data; MFISH 2014a). Arctic
grayling derived from these two sources
were stocked on top of every known
native Arctic grayling population in the
upper Missouri River basin. In addition,
Arctic grayling were stocked in multiple
high elevation lakes, some of which
likely were historically fishless.
There are 20 known, introduced
Arctic grayling populations that exist in
the upper Missouri River basin. These
20 populations, along with the 6
populations existing in native habitat,
comprise the listable entity (total of 26
populations) of Arctic grayling in the
upper Missouri River basin. However,
six of these introduced populations are
considered to have low conservation
value because they occupy unnatural
habitat, are not self-sustaining, or are
used as captive brood reserves. These
six populations are Axolotl Lake, Green
Hollow Lake, Sunnyslope Canal, Tunnel
Lake, South Fork Sun River, and Elk
Lake. The Axolotl and Green Hollow
populations are captive brood reserves
maintained in natural lakes for
reintroduction purposes. Sunnyslope
Canal is a fluvial population that occurs
in unnatural habitat (irrigation canal).
Tunnel Lake is stocked with ‘‘rescued’’
fish from Sunnyslope Canal, but lacks a
spawning tributary and is consequently
not self-sustaining (SSA 2014). South
Fork Sun River is a small fluvial
population that resides in about 1⁄4 mile
of stream during the summer and is not
considered self-sustaining (SSA 2014).
The Elk Lake population is a genetic
replicate of the Centennial Valley
population, but no documented
spawning has occurred to date (Jaeger
2014a, pers. comm.); thus this
population is not currently considered
self-sustaining. For these reasons, we
primarily focus our analysis on the
populations considered to have high
conservation value; those populations
that are self-sustaining, in natural
habitats, and wild.
The 14 known remaining introduced,
lake-dwelling (adfluvial) Arctic grayling
populations within the upper Missouri
River basin are likely the result of
historical stocking (Table 1). In our 2010
finding, we considered and discussed
the conservation value of these
populations. Based on the information
available at that time, we considered
these introduced populations to not
have conservation value for multiple
reasons. Below, we list each of the
reasons for this conclusion as provided
in the 2010 finding, and provide an
updated assessment and conclusion
about the potential conservation value
of these populations, based on new
information obtained since 2010.
TABLE 1—GEOGRAPHIC DISTRIBUTION, GENETIC STATUS, AND SOURCE OF INTRODUCED ADFLUVIAL ARCTIC GRAYLING
POPULATIONS IN THE UPPER MISSOURI RIVER BASIN
Drainage
Genetic
analysis
completed?
Source a
Citation
Agnes Lake ................
Odell Lake ..................
Bobcat Lake ...............
Schwinegar Lake .......
Pintlar Lake ................
Deer Lake ..................
Emerald Lake .............
Grayling Lake .............
Hyalite Lake ...............
Diversion Lake ...........
Gibson Reservoir .......
Lake Levale ...............
Park Lake ...................
Grebe Lake ................
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Big Hole ...
Big Hole ...
Big Hole ...
Big Hole ...
Big Hole ...
Gallatin .....
Gallatin .....
Gallatin .....
Gallatin .....
Sun ..........
Sun ..........
Sun ..........
Missouri ...
Madison ...
No ............
Yes ...........
Yes ...........
No ............
Yes ...........
Yes ...........
Yes ...........
Yes ...........
Yes ...........
Yes b ........
Yes b ........
Yes b ........
No ............
Yes ...........
Madison/Centennial ...
Centennial ..................
Centennial ..................
Madison/Centennial.c
Madison/Centennial ...
Madison/Centennial ...
Madison/Centennial ...
Madison/Centennial ...
Madison/Centennial ...
Big Hole .....................
Big Hole .....................
Big Hole .....................
Madison/Centennial.c
Centennial ..................
MFISH 2014a.
Peterson and Ardren 2009, p. 1766; Leary 2014, unpublished data.
Peterson and Ardren 2009, p. 1766; Leary 2014, unpublished data.
Leary 2014, unpublished data.
Leary 2014, unpublished data.
Leary 2014, unpublished data.
Leary 2014, unpublished data.
Leary 2014, unpublished data.
Horton 2014a, pers. comm.; Magee 2014, pers. comm.
Horton 2014a, pers. comm.; Magee 2014, pers. comm.
Horton 2014a, pers. comm.; Magee 2014, pers. comm.
Peterson and Ardren 2009, p. 1766; Varley 1981, p. 11.
a Origin of source stock was determined by genetic analysis and through analysis of historical stocking records and scientific literature, in some
cases. Where multiple sources are cited, fish from each population were known to be stocked, although the genetic contribution of each donor
population to the current population structure is unknown.
b These populations are the result of reintroductions using known sources of Montana origin.
c Schwinegar and Park Lakes Arctic grayling populations are likely from Montana-origin sources due to proximity to other lakes with known
Montana origin; however, definitive evidence is lacking.
1. The Service interprets the Act to
provide a statutory directive to conserve
species in their native ecosystems (49
FR 33885, August 27, 1984) and to
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conserve genetic resources and
biodiversity over a representative
portion of a taxon’s historical
occurrence (61 FR 4722, February 7,
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1996). Since most of the introduced
populations of Arctic grayling were of
unknown genetic origin and in lakes
that were likely historically fishless,
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these populations were considered in
2010 to be outside the species’ native
range, and we concluded that they did
not appear to add conservation value to
the DPS.
Since 2010, new genetic information
from 7 of the 14 introduced populations
indicates there are moderate to high
levels of genetic diversity within and
among these populations, and indicates
these populations were derived from
native sources within the upper
Missouri River basin (Leary 2014,
unpublished data; Table 1). In addition,
stocking records show common stocking
sources for introduced populations that
were genotyped (as described
previously) and the two populations
that were not genotyped (the remaining
3 populations were reintroductions of
known Montana origin sources; Table
1). Thus, it appears that all 14
introduced Arctic grayling populations
contain moderate to high levels of
genetic diversity of Arctic grayling in
the upper Missouri River basin that was
not captured within the DPS
designation in the 2010 finding.
The Service’s current interpretation of
the Act is consistent with that in the
2010 finding; we believe it is important
to conserve species in their native
ecosystems and to conserve genetic
resources and biodiversity over a
representative portion of a taxon’s
historical occurrence. In light of the new
genetics information gained since 2010
(Leary 2014, unpublished data), we also
believe it is important to acknowledge
the moderate to high levels of genetic
diversity within the introduced
populations in the upper Missouri River
basin and the potential adaptive
capabilities represented by this
diversity. All Arctic grayling
populations (introduced or not)
currently within the upper Missouri
River basin are derived from a common
ancestor and have a distinct
evolutionary trajectory relative to the
historical founding populations in
Canada and Alaska. Thus, Arctic
grayling originating from and currently
within the upper Missouri River basin
represent the southernmost assemblage
of the species, facing similar selection
pressures and evolving independent of
more northern populations.
The introduced Arctic grayling
populations in the upper Missouri River
basin occupy, for the most part, highelevation habitats that are high-quality
because of intact riparian areas and a
consistent supply of cool water. Given
the predicted effects of climate change
in the West (see discussion under
‘‘Climate Change’’ in Factor A below),
these types of habitats are the same
habitats that the Service would explore
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for long-term conservation of Arctic
grayling, if needed, because they may
serve as thermal refugia as temperatures
rise and provide greater redundancy in
case of catastrophic events.
2. In 2010, the Service concluded
there did not appear to be any formally
recognized conservation value for the
introduced populations of Arctic
grayling in the upper Missouri River
basin because they were not being used
in conservation or restoration programs.
This conclusion was based on an
interpretation of a National Marine
Fisheries Service final policy on the
consideration of hatchery-origin fish in
Endangered Species Act listing
determinations for Pacific salmon and
steelhead (anadromous Oncorhynchus
spp.) (NMFS 2005, entire).
Until recently, the genetic structure
and source of these introduced
populations were unknown.
Populations with a high likelihood of
being Montana origin were used for
conservation purposes (e.g.,
reintroductions) as a precautionary
approach to Arctic grayling
conservation. Now that the amount of
genetic diversity within and among the
introduced Arctic grayling populations
and their source(s) are known, it is
probable these introduced populations
could be used in future conservation
actions as source stock, if needed.
3. In 2010, the Service indicated there
were concerns that introduced, lakedwelling Arctic grayling populations
could pose genetic risks to the native
fluvial population (i.e., Big Hole
Population) as cited in the Montana
Fluvial Arctic Grayling Restoration Plan
(‘‘Restoration Plan,’’ 1995, p. 15). In the
Restoration Plan, Arctic grayling
populations in Agnes, Schwinegar,
Odell, Miner and Mussigbrod lakes were
identified as potential threats to the
genetic integrity of the Big Hole River
population because of hydrologic
connectivity between these lakes and
the Big Hole River and the potential for
genetic mixing.
Recently, genetic analyses have
confirmed reproductive isolation among
extant Arctic grayling populations in the
upper Missouri River basin and within
the Big Hole River watershed (Peterson
and Ardren 2009, p. 1770; Leary 2014,
unpublished data). In addition, multiple
historical stockings have occurred in the
Big Hole River from other sources
within the upper Missouri River basin.
Recent genetic analysis found no
evidence of a significant genetic
contribution from historical stocking on
the current genetic structure of Arctic
grayling in the Big Hole River (Peterson
and Ardren 2009, p. 1768). Thus, we
now conclude that the concern that
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lake-dwelling populations within the
Big Hole River watershed could pose
genetic risks to the Big Hole River
fluvial population appears unfounded.
4. In 2010, the Service concluded that
introduced populations of Arctic
grayling in the upper Missouri River
basin had no conservation value
because these populations apparently
had been isolated from their original
source stock for decades without any
supplementation from the wild and
were established without any formal
genetic consideration to selecting and
mating broodstock.
It is now apparent from our review of
historical stocking records that many of
these introduced populations received
multiple stockings from the same source
or multiple stockings from several
different sources over a wide range of
years (MFISH 2014a, unpublished data).
Additionally, most individual stockings
involved a large number of eggs or fry
(up to 1 million for some stockings).
Cumulatively, this information suggests
several points. First, stockings that used
a large number of eggs or fry necessitate
that gametes from multiple brood fish
were used per stocking, given the
physical constraints of number of eggs
per unit body size of female Arctic
grayling. Second, stockings in most of
the introduced populations occurred
over many years (up to 60 years in some
cases). This indicates different cohorts
of Arctic grayling had to be used, since
the generation time of Arctic grayling is
approximately 3.5 years in the upper
Missouri River basin (references in
Dehaan et al. 2014, p. 10). Lastly, the
new genetic analyses from seven of the
introduced Arctic grayling populations
indicate moderate to high levels of
genetic diversity within the
populations. This result could likely
only be obtained from the founding of
these populations using large numbers
of brood fish and gametes over multiple
years. Mutation is unlikely to have
accounted for these levels of genetic
diversity over a relatively short time
period of isolation (Freeman and Herron
2001, p. 143).
For perspective, Montana Fish,
Wildlife, and Parks has developed
guidelines for the establishment and
maintenance of Arctic grayling
broodstock. To adequately capture most
of the genetic variation in a source
population, the crossing of a minimum
of 25 male and 25 female Arctic grayling
is currently recommended (Leary 1991,
p. 2151). It is likely that the historical
stockings used to found the introduced
Arctic grayling populations in the upper
Missouri River basin equaled or
exceeded this through stocking large
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numbers of eggs or fry over multiple
years.
5. In 2010, the Service concluded that
the source populations used to found
the introduced Arctic grayling
populations in the upper Missouri River
drainage were not well documented
(Peterson and Ardren 2009, p. 1767), so
we could not be certain of whether these
Arctic grayling were of local origin.
Since 2010, new genetic information
(Leary 2014, unpublished data) and
review of historical stocking records
(MFISH 2014a, unpublished data)
indicate the founding populations used
for stocking are local and believed
representative of the Upper Missouri
River DPS of Arctic grayling, and
contain moderate to high levels of
genetic diversity.
6. In 2010, the Service concluded the
primary intent of culturing and
introducing Arctic grayling populations
within the upper Missouri River basin
was to provide recreational fishing
opportunities in high mountain lakes,
and that, therefore, these introduced
populations had no conservation value.
Since 2010, review of the historical
literature indicates adfluvial Arctic
grayling populations were presumably
stocked both for recreational fishing and
conservation purposes (Brown 1943, pp.
26–27; Nelson 1954, p. 341; Vincent
1962, p. 151). Following the drought in
the 1930s, conservation stockings of
Arctic grayling were advocated because
most rivers and streams were
dewatered, prompting fish managers to
introduce Arctic grayling into habitats
with a more consistent supply of cool
water (e.g., high-elevation mountain
lakes; Brown 1943, pp. 26–27; Nelson
1954, p. 341; Vincent 1962, p. 151).
In conclusion, introduced populations
of Arctic grayling established within the
upper Missouri River basin, whether
they were originally established for
recreational fishing or conservation
purposes, captured moderate to high
levels of genetic diversity of upper
Missouri River basin Arctic grayling.
The potential adaptive capabilities
represented by this genetic diversity
have conservation value, particularly in
a changing climate. These populations
reside in high-quality habitat, the same
habitat the Service would look to for
long-term conservation, if needed. Thus,
the introduced populations of Arctic
grayling within the upper Missouri
River basin have conservation value,
and, therefore, we include them in our
analysis of a potential DPS of Arctic
grayling.
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Origins, Biogeography, and Genetics of
Arctic Grayling in North America
North American Arctic grayling are
most likely descended from Eurasian
Thymallus that crossed the Bering land
bridge during or before the Pleistocene
glacial period (Stamford and Taylor
2004, pp. 1533, 1546). There were
multiple opportunities for freshwater
faunal exchange between North America
and Asia during the Pleistocene, but
genetic divergence between North
American and Eurasian Arctic grayling
suggests that the species could have
colonized North America as early as the
mid-late Pliocene (more than 3 million
years ago) (Stamford and Taylor 2004, p.
1546). Genetic studies of Arctic grayling
using mitochondrial DNA (mtDNA,
maternally inherited DNA located in
cellular organelles called mitochondria)
and microsatellite DNA (repeating
sequences of nuclear DNA) have shown
that North American Arctic grayling
consist of at least three major lineages
that originated in distinct Pleistocene
glacial refugia (Stamford and Taylor
2004, p. 1533). These three groups
include a South Beringia lineage found
in western Alaska to northern British
Columbia, Canada; a North Beringia
lineage found on the North Slope of
Alaska, the lower Mackenzie River, and
to eastern Saskatchewan; and a Nahanni
lineage found in the lower Liard River
and the upper Mackenzie River drainage
in northeastern British Columbia and
southeastern Yukon (Stamford and
Taylor 2004, pp. 1533, 1540). Arctic
grayling from the upper Missouri River
basin were tentatively placed in the
North Beringia lineage because a small
sample (three individuals) of Montana
Arctic grayling shared a mtDNA
haplotype (form of the mtDNA) with
populations in Saskatchewan and the
lower Peace River, British Columbia
(Stamford and Taylor 2004, p. 1538).
The existing mtDNA data suggest that
Missouri River Arctic grayling share a
common ancestry with the North
Beringia lineage, but other genetic
markers (e.g., allozymes, microsatellites)
and biogeographic history indicate that
Missouri River Arctic grayling have
been physically and reproductively
isolated from northern populations for
millennia. Pre-glacial colonization of
the Missouri River basin by Arctic
grayling was possible because the river
flowed to the north and drained into the
Arctic-Hudson Bay prior to the last
glacial cycle (Cross et al. 1986, pp. 374–
375; Pielou 1991, pp. 194–195). Low
mtDNA diversity observed in a small
number of Montana Arctic grayling
samples and a shared ancestry with
Arctic grayling from the North Beringia
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lineage suggest a more recent, postglacial colonization of the upper
Missouri River basin. In contrast,
microsatellite DNA show substantial
divergence between Montana and
Saskatchewan (i.e., same putative
mtDNA lineage) (Peterson and Ardren
2009, entire). Differences in the
frequency and size distribution of
microsatellite alleles between Montana
populations and two Saskatchewan
populations indicate that Montana
Arctic grayling have been isolated long
enough for mutations (i.e., evolution) to
be responsible for the observed genetic
differences.
Additional comparison of 21 Arctic
grayling populations from Alaska,
Canada, and the Missouri River basin
using 9 of the same microsatellite loci
as Peterson and Ardren (2009, entire)
further supports the distinction of
Missouri River Arctic grayling relative
to populations elsewhere in North
America (USFWS, unpublished data).
Analyses of these data using two
different methods clearly separates
sample fish from 21 populations into
two clusters: One cluster representing
populations from the upper Missouri
River basin, and another cluster
representing populations from Canada
and Alaska (USFWS, unpublished data).
These new data, although not yet peer
reviewed, support the interpretation
that the previous analyses of Stamford
and Taylor (2004, entire)
underestimated the distinctiveness of
Missouri River Arctic grayling relative
to other sample populations, likely
because of the combined effect of small
sample sizes and the lack of variation
observed in the Missouri River for the
markers used in that study (Stamford
and Taylor 2004, pp. 1537–1538). Thus,
these recent microsatellite DNA data
suggest that Arctic grayling may have
colonized the Missouri River before the
onset of Wisconsin glaciation (more
than 80,000 years ago).
Genetic relationships among native
and introduced populations of Arctic
grayling in Montana have recently been
investigated (Peterson and Ardren 2009,
entire). Introduced, lake-dwelling
populations of Arctic grayling trace
some of their original ancestry to the
Centennial Valley (Peterson and Ardren
2009, p. 1767), and stocking of hatchery
Arctic grayling did not have a large
effect on the genetic composition of the
extant native populations (Peterson and
Ardren 2009, p. 1768). Differences
between native populations of the two
Arctic grayling ecotypes (adfluvial,
fluvial) are not as large as differences
resulting from geography (i.e., drainage
of origin). For example, native adfluvial
Arctic grayling populations from
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different lakes are genetically different
(Peterson and Ardren 2009, p. 1766).
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Habitat
Arctic grayling generally require clear,
cold water, and are characterized as
belonging to a ‘‘coldwater’’ group of
salmonids, which also includes bull
trout (Salvelinus confluentus) and
Arctic char (Salvelinus alpinus) (Selong
et al. 2001, p. 1032). Arctic grayling
optimal thermal habitat is between 7 to
17 °C (45 to 63 °F), but becomes
unsuitable above 20 °C (68 °F) (Hubert
et al. 1985, p. 24). Arctic grayling fry
may be more tolerant of high water
temperature than adults (LaPerriere and
Carlson 1973, p. 30; Feldmeth and
Eriksen 1978, p. 2041).
Having a broad, nearly circumpolar
distribution, Arctic grayling occupy a
variety of habitats including small
streams, large rivers, lakes, and even
bogs (Northcote 1995, pp. 152–153;
Scott and Crossman 1998, p. 303). They
may even enter brackish water (less than
or equal to 4 parts per thousand salt
content) when migrating between
adjacent river systems (West et al. 1992,
pp. 713–714). Native populations are
found at elevations ranging from near
sea level, such as in Bristol Bay, Alaska,
to high-elevation montane valleys (more
than 1,830 meters (m) or 6,000 feet (ft)),
such as the Big Hole River and
Centennial Valley in southwestern
Montana. Despite this broad
distribution, Arctic grayling have
specific habitat requirements that can
constrain their local distributions,
especially water temperature and
channel gradient. At the local scale,
Arctic grayling prefer cold water and are
often associated with spring-fed habitats
in regions with warmer climates
(Vincent 1962, p. 33). Arctic grayling are
generally not found in swift, highgradient streams, and Vincent (1962, pp.
36–37, 41–43) characterized typical
Arctic grayling habitat in Montana (and
Michigan) as low-to-moderate gradient
(less than 4 percent) streams and rivers
with low-to-moderate water velocities
(less than 2 feet/sec (60 centimeters/
sec)). Juvenile and adult Arctic grayling
in streams and rivers spend much of
their time in pool habitat (Kaya 1990
and references therein, p. 20; Lamothe
and Magee 2003, pp. 13–14).
Breeding
Arctic grayling typically spawn in the
spring or early summer, depending on
latitude and elevation (Northcote 1995,
p. 149). In Montana, Arctic grayling
generally spawn from late April to midMay by depositing adhesive eggs over
gravel substrate without excavating a
nest (Kaya 1990, p. 13; Northcote 1995,
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p. 151). In general, the reproductive
ecology of Arctic grayling differs from
other salmonid species (trout and
salmon) in that Arctic grayling eggs tend
to be comparatively small; thus, they
have higher relative fecundity (females
have more eggs per unit body size).
Males establish and defend spawning
territories rather than defending access
to females (Northcote 1995, pp. 146,
150–151). The time required for
development of eggs from embryo until
they emerge from stream gravel and
become swim-up fry depends on water
temperature (Northcote 1995, p. 151). In
the upper Missouri River basin,
development from embryo to fry
averages about 3 weeks (Kaya 1990, pp.
16–17). Small, weakly swimming fry
(typically 1–1.5 centimeters (cm) (0.4–
0.6 in.) at emergence) prefer lowvelocity stream habitats (Armstrong
1986, p. 6; Kaya 1990, pp. 23–24;
Northcote 1995, p. 151).
Arctic grayling of all ages feed
primarily on aquatic and terrestrial
invertebrates captured on or near the
water surface, but also will feed
opportunistically on fish and fish eggs
(Northcote 1995, pp. 153–154; Behnke
2002, p. 328). Feeding locations for
individual fish are typically established
and maintained through size-mediated
dominance hierarchies where larger
individuals defend favorable feeding
positions (Hughes 1992, p. 1996).
General Life History Diversity
Migratory behavior is a common lifehistory trait in salmonid fishes such as
Arctic grayling (Armstrong 1986, pp. 7–
8; Northcote 1995, pp. 156–158; 1997,
pp. 1029, 1031–1032, 1034). In general,
migratory behavior in Arctic grayling
and other salmonids results in cyclic
patterns of movement between refuge,
rearing-feeding, and spawning habitats
(Northcote 1997, p. 1029).
Arctic grayling may move to refuge
habitat as part of a regular seasonal
migration (e.g., in winter), or in
response to episodic environmental
stressors (e.g., high summer water
temperatures). In Alaska, Arctic grayling
in rivers typically migrate downstream
in the fall, moving into larger streams or
mainstem rivers that do not completely
freeze (Armstrong 1986, p. 7). In Arctic
rivers, fish often seek overwintering
habitat influenced by groundwater
(Armstrong 1986, p. 7). In some
drainages, individual fish may migrate
considerable distances (greater than 150
km or 90 mi) to overwintering habitats
(Armstrong 1986, p. 7). In the Big Hole
River, Montana, similar downstream
and long-distance movement to
overwintering habitat has been observed
in Arctic grayling (Shepard and Oswald
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49391
1989, pp. 18–21, 27). In addition, Arctic
grayling in the Big Hole River may move
downstream in proximity to colder
tributary streams in summer when
thermal conditions in the mainstem
river become stressful (Lamothe and
Magee 2003, p. 17).
In spring, mature Arctic grayling leave
overwintering areas and migrate to
suitable spawning sites. In river
systems, this typically involves an
upstream migration to tributary streams
or shallow riffles within the mainstem
(Armstrong 1986, p. 8; Shepard and
Oswald 1989; p. 18). Arctic grayling in
lakes typically migrate to either the inlet
or outlet to spawn (Armstrong 1986, p.
8; Kaya 1989, p. 474; Northcote 1995 p.
148). In some situations, Arctic grayling
exhibit natal homing, whereby
individuals spawn in or near the
location where they were born
(Northcote 1995 pp. 157–160; Boltz and
Kaeding 2002, p. 22); however, it is
unclear what factors may be influencing
the extent of this phenomenon.
Fry from river populations typically
seek feeding and rearing habitats in the
vicinity of where they were spawned
(Armstrong 1986, pp. 6–7; Kaya and
Jeanes 1995, p. 455; Northcote 1995, p.
156), while those from lake populations
migrate downstream (inlet spawners) or
upstream (outlet spawners) to the
adjacent lake. Following spawning,
adults move to appropriate feeding areas
if they are not adjacent to spawning
habitat (Armstrong 1986, pp. 7–8;
Shepard and Oswald 1989; p. 18).
Juvenile Arctic grayling may undertake
seasonal migrations between feeding
and overwintering habitats until they
reach maturity and add the spawning
migration to this cycle (Northcote 1995,
pp. 156–157).
Life History Diversity in Arctic Grayling
in the Upper Missouri River Basin
Two general life-history forms or
ecotypes of native Arctic grayling occur
in the upper Missouri River Arctic:
Fluvial and adfluvial. Fluvial fish use
river or stream (lotic) habitat for all of
their life cycles and may undergo
extensive migrations within river
habitat, up to 50 miles in the Big Hole
River in Montana (Shepard and Oswald
1989, p. 18). Adfluvial fish live in lakes
and migrate to tributary streams to
spawn. These same life-history forms
also are expressed by Arctic grayling
elsewhere in North America (Northcote
1997, p. 1030). Historically, the fluvial
life-history form predominated in the
Missouri River basin above the Great
Falls, perhaps because there were only
a few lakes accessible to natural
colonization of Arctic grayling that
would permit expression of the
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adfluvial ecotype (Kaya 1992, p. 47).
The fluvial and adfluvial life-history
forms of Arctic grayling in the upper
Missouri River do not appear to
represent distinct evolutionary lineages.
Instead, they appear to represent an
example of adaptive radiation (Schluter
2000, p. 1), whereby the forms
differentiated from a common ancestor
and developed traits that allowed them
to exploit different habitats. The
primary evidence for this conclusion is
genetic data that indicate that within the
Missouri River basin the two ecotypes
are more closely related to each other
than they are to the same ecotype
elsewhere in North America (Redenbach
and Taylor 1999, pp. 27–28; Stamford
and Taylor 2004, p. 1538; Peterson and
Ardren 2009, p. 1766). Historically,
there may have been some genetic
exchange between the two life-history
forms as individuals strayed or
dispersed into different populations
(Peterson and Ardren 2009, p. 1770), but
the genetic structure of current
populations in the upper Missouri River
basin is consistent with reproductive
isolation.
The fluvial and adfluvial forms of
Arctic grayling appear to differ in their
genetic characteristics, but there appears
to be some plasticity in behavior where
individuals from a population can
exhibit a range of behaviors. Arctic
grayling fry in Montana can exhibit
heritable, genetically-based differences
in swimming behavior between fluvial
and adfluvial ecotypes (Kaya 1991, pp.
53, 56–58; Kaya and Jeanes 1995, pp.
454, 456). Progeny of Arctic grayling
from the fluvial ecotype exhibited a
greater tendency to hold their position
in flowing water relative to progeny
from adfluvial ecotypes (Kaya 1991, pp.
53, 56–58; Kaya and Jeanes 1995, pp.
454, 456). Similarly, young Arctic
grayling from inlet and outlet spawning
adfluvial ecotypes exhibited an innate
tendency to move downstream and
upstream, respectively (Kaya 1989, pp.
478–480). All three studies (Kaya 1989,
entire; 1991, entire; Kaya and Jeanes
1995, entire) demonstrate that the
response of fry to flowing water
depended strongly on the life-history
form (ecotype) of the source population,
and that this behavior has a genetic
basis. However, behavioral responses
also were mediated by environmental
conditions (light—Kaya 1991, pp. 56–
57; light and water temperature—Kaya
1989, pp. 477–479), and some progeny
of each ecotype exhibited behavior
characteristic of the other; for example
some individuals from the fluvial
ecotype moved downstream rather than
holding position, and some individuals
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from an inlet-spawning adfluvial
ecotype held position or moved
upstream (Kaya 1991, p. 58). These
observations indicate that some
plasticity for behavior exists, at least for
very young Arctic grayling.
The ability of the fluvial ecotype to
give rise to a functional population of
the adfluvial ecotype has been
demonstrated. Most extant adfluvial
Arctic grayling populations in the
Upper Missouri River originated from
fluvial-dominated sources (see Table 1;
Kaya 1992, p. 53; Jeanes 1996, pp. 54).
However, the ability of the adfluvial
ecotype to give rise to a functional
population of fluvial ecotype is less
certain. Circumstantial support for
reduced plasticity in adfluvial Arctic
grayling comes from observations that
adfluvial fish stocked in river habitats
almost never establish populations
(Kaya 1990, pp. 31–34). However, we
note that adfluvial Arctic grayling retain
some life-history flexibility—at least in
lake environments—as naturalized
populations derived from inletspawning stocks have established
outlet-spawning demes (a deme is a
local populations that shares a distinct
gene pool) in Montana and in
Yellowstone National Park (Kruse 1959,
p. 318; Kaya 1989, p. 480). In addition,
a small percentage of young adfluvial
Arctic grayling exposed to flow
exhibited fluvial-like characteristics
(e.g., station-holding or upstream
movement) in a laboratory experiment
designed to assess movement tendencies
of adfluvial and fluvial Arctic grayling
in flowing water (Kaya 1991, p. 56).
These results indicate some plasticity
exists in adfluvial Arctic grayling that
may allow some progeny of adfluvial
individuals to express a fluvial life
history. Nonetheless, the frequent
failure of introductions of adfluvial
Arctic grayling into fluvial habitats
suggest a cautionary approach to the
loss of particular life-history forms is
warranted.
Age and Growth
Age at maturity and longevity in
Arctic grayling varies regionally and is
probably related to growth rate, with
populations in colder, northern
latitudes maturing at later ages and
having a greater lifespan (Kruse 1959,
pp. 340–341; Northcote 1995 and
references therein, pp. 155–157). Arctic
grayling in the upper Missouri River
typically mature at age 2 (males) or age
3 (females), and individuals greater than
age 6 are rare (Kaya 1990, p. 18; Magee
and Lamothe 2003, pp. 16–17). The
majority of the Arctic grayling spawning
in two tributaries in the Centennial
Valley, Montana, were age 3, and the
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oldest individuals aged from a larger
sample were age 6 (Nelson 1954, pp.
333–334). Arctic grayling spawning in
Red Rock Creek were mostly ages 2 to
5, but some individuals were age 7
(Mogen 1996, pp. 32–34).
Generally, growth rates of Arctic
grayling are greatest during the first
years of life then slow dramatically after
maturity. Within that general pattern,
there is substantial variation among
populations from different regions.
Arctic grayling populations in Montana
(Big Hole River and Red Rock Lakes)
have very high growth rates relative to
those from British Columbia, Asia, and
the interior and North Slope of Alaska
(Carl et al. 1992, p. 240; Northcote 1995,
pp. 155–157; Neyme 2005, p. 28).
Distinct Vertebrate Population Segment
Under the Service’s Policy Regarding
the Recognition of Distinct Vertebrate
Population Segments Under the
Endangered Species Act (61 FR 4722;
February 7, 1996), three elements are
considered in the decision concerning
the establishment and classification of a
possible DPS. These are applied
similarly for additions to or removal
from the Federal List of Endangered and
Threatened Wildlife. These elements
include:
(1) The discreteness of a population in
relation to the remainder of the species
to which it belongs;
(2) The significance of the population
segment to the species to which it
belongs; and
(3) The population segment’s
conservation status in relation to the
Act’s standards for listing, delisting, or
reclassification (i.e., is the population
segment endangered or threatened).
Discreteness
Under the DPS policy, a population
segment of a vertebrate taxon 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.
(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.
Arctic grayling native to the upper
Missouri River are isolated from all
other populations of the species, which
inhabit the Arctic Ocean, Hudson Bay,
and north Pacific Ocean drainages in
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Asia and North America. Arctic grayling
native to the upper Missouri River occur
as a disjunct group of populations
approximately 800 km (500 mi) to the
south of the next-nearest Arctic grayling
population in central Alberta, Canada.
Missouri River Arctic grayling have
been isolated from other populations for
at least 10,000 years based on historical
reconstruction of river flows at or near
the end of the Pleistocene (Cross et al.
1986, p. 375; Pileou 1991, pp. 10–11).
Genetic data confirm Arctic grayling in
the Missouri River basin have been
reproductively isolated from
populations to the north for millennia
(Everett 1986, pp. 79–80; Redenbach
and Taylor 1999, p. 23; Stamford and
Taylor 2004, p. 1538; Peterson and
Ardren 2009, pp. 1764–1766; USFWS,
unpublished data). Consequently, we
conclude that Arctic grayling native to
the upper Missouri River are markedly
separated from other native populations
of the taxon as a result of physical
factors (isolation), and therefore meet
the first criterion of discreteness under
the DPS policy. As a result, Arctic
grayling native to the upper Missouri
River are considered a discrete
population according to the DPS policy.
Because the entity meets the first
criterion (markedly separated), an
evaluation with respect to the second
criterion (international boundaries) is
not needed.
Significance
If a population segment is considered
discrete under one or more of the
conditions described in the Service’s
DPS policy, its biological and ecological
significance will be considered in light
of Congressional guidance that the
authority to list DPSs be used
‘‘sparingly’’ while encouraging the
conservation of genetic diversity. In
making this determination, we consider
available scientific evidence of the
discrete population segment’s
importance to the taxon to which it
belongs. Since precise circumstances are
likely to vary considerably from case to
case, the DPS policy does not describe
all the classes of information that might
be used in determining the biological
and ecological importance of a discrete
population. However, the DPS policy
describes four possible classes of
information that provide evidence of a
population segment’s biological and
ecological importance to the taxon to
which it belongs. As specified in the
DPS policy (61 FR 4722), this
consideration of the population
segment’s significance may include, but
is not limited to, the following:
(1) Persistence of the discrete
population segment in an ecological
setting unusual or unique to the taxon;
(2) Evidence that loss of the discrete
population segment would result in a
significant gap in the range of a taxon;
(3) Evidence that the discrete
population segment represents the only
surviving natural occurrence of a taxon
that may be more abundant elsewhere as
an introduced population outside its
historical range; or
(4) Evidence that the discrete
population segment differs markedly
from other populations of the species in
its genetic characteristics.
A population segment needs to satisfy
only one of these conditions to be
considered significant. Furthermore,
other information may be used as
appropriate to provide evidence for
significance.
Unique Ecological Setting
Water temperature is a key factor
influencing the ecology and physiology
of ectothermic (body temperature
regulated by ambient environmental
conditions) salmonid fishes, and can
dictate reproductive timing, growth and
development, and life-history strategies.
Groundwater temperatures can be
related to air temperatures (Meisner
1990, p. 282), and thus reflect the
49393
regional climatic conditions. Warmer
groundwater influences ecological
factors such as food availability, the
efficiency with which food is converted
into energy for growth and
reproduction, and ultimately growth
rates of aquatic organisms (Allan 1995,
pp. 73–79). Aquifer structure and
groundwater temperature is important
to salmonid fishes because groundwater
can strongly influence stream
temperature, and consequently egg
incubation and fry growth rates, which
are strongly temperature-dependent
(Coutant 1999, pp. 32–52; Quinn 2005,
pp. 143–150).
Missouri River Arctic grayling occur
within the 4 to 7 °C (39 to 45 °F) ground
water isotherm (see Heath 1983, p. 71;
an isotherm is a line connecting bands
of similar temperatures on the earth’s
surface), whereas most other North
American Arctic grayling are found in
isotherms less than 4 °C, and much of
the species’ range is found in areas with
discontinuous or continuous permafrost
(Meisner et al. 1988, p. 5; Table 2).
Much of the historical range of Arctic
grayling in the upper Missouri River is
encompassed by mean annual air
temperature isotherms of 5 to 10 °C (41
to 50 °F) (USGS 2009), with the colder
areas being in the headwaters of the
Madison River in Yellowstone National
Park. In contrast, Arctic grayling in
Canada, Alaska, and Asia are located in
regions encompassed by air temperature
isotherms 5 °C and colder (41 °F and
colder), with much of the species
distributed within the 0 to ¥10 °C
isolines (32 to 14 °F). This difference is
significant because Arctic grayling in
the Missouri River basin have evolved
in isolation for millennia in a generally
warmer climate than other populations.
The potential for thermal adaptations
makes Missouri River Arctic grayling a
significant biological resource for the
species under expected climate change
scenarios.
TABLE 2—DIFFERENCES BETWEEN THE ECOLOGICAL SETTING OF THE UPPER MISSOURI RIVER AND ELSEWHERE IN THE
SPECIES’ RANGE OF ARCTIC GRAYLING
Missouri River
Rest of taxon
Bailey’s Ecoregion ....................
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Ecological setting variable
Dry Domain: Temperate
Steppe.
5 to 10 °C (41 to 50 °F) ..........
4 to 7 °C (39 to 45 °F) ............
Polar Domain: Tundra & Subarctic Humid Temperate: Marine, Prairie, Warm
Continental Mountains.
¥15 to 5 °C (5 to 41 °F).
Less than 4 °C (Less than 39 °F).
Air temperature (isotherm) ........
Groundwater temperature (isotherm).
Arctic grayling in the upper Missouri
River basin occur in a temperate
ecoregion distinct from all other Arctic
grayling populations worldwide, which
occur in Arctic or sub-Arctic ecoregions
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dominated by Arctic flora and fauna. An
ecoregion is a continuous geographic
area within which there are associations
of interacting biotic and abiotic features
(Bailey 2005, pp. S14, S23). These
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ecoregions delimit large areas within
which local ecosystems recur more or
less in a predictable fashion on similar
sites (Bailey 2005, p. S14). Ecoregional
classification is hierarchical, and based
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on the study of spatial coincidences,
patterning, and relationships of climate,
vegetation, soil, and landform (Bailey
2005, p. S23). The largest ecoregion
categories are domains, which represent
subcontinental areas of similar climate
(e.g., polar, humid temperate, dry, and
humid tropical) (Bailey 1994; 2005, p.
S17). Domains are divided into
divisions that contain areas of similar
vegetation and regional climates. Arctic
grayling in the upper Missouri River
basin are the only example of the
species naturally occurring in a dry
domain (temperate steppe division;
Table 2). The vast majority of the
species’ range is found in the polar
domain (all of Asia, most of North
America), with small portions of the
range occurring in the humid temperate
domain (northern British Columbia and
southeast Alaska). Occupancy of
Missouri River Arctic grayling in a
temperate ecoregion is significant for
two primary reasons. First, an ecoregion
represents a suite of factors (climate,
vegetation, landform) influencing, or
potentially influencing, the evolution of
species within that ecoregion. Since
Missouri River Arctic grayling have
existed for thousands of years in an
ecoregion quite different from the
majority of the taxon, they have likely
developed adaptations during these
evolutionary timescales that distinguish
them from the rest of the taxon, even if
we have yet to conduct the proper
studies to measure these adaptations.
Second, the occurrence of Missouri
River Arctic grayling in a unique
ecoregion helps reduce the risk of
species-level extinction, as the different
regions may respond differently to
environmental change.
Arctic grayling in the upper Missouri
River basin have existed for at least
10,000 years in an ecological setting
quite different from that experienced by
Arctic grayling elsewhere in the species’
range. The most salient aspects of this
different setting relate to temperature
and climate, which can strongly and
directly influence the biology of
ectothermic species (like Arctic
grayling). Arctic grayling in the upper
Missouri River have experienced
warmer temperatures than most other
populations. Physiological and lifehistory adaptation to local temperature
regimes are regularly documented in
salmonid fishes (Taylor 1991, pp. 191–
193), but experimental evidence for
adaptations to temperature, such as
unusually high temperature tolerance or
lower tolerance to colder temperatures,
is lacking for Missouri River Arctic
grayling because the appropriate studies
have not been conducted. Lohr et al.
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(1996, p. 934) studied the upper thermal
tolerances of Arctic grayling from the
Big Hole River, but their research design
did not include other populations from
different thermal regimes, so it was not
possible to make between-population
contrasts under a common set of
conditions. Arctic grayling from the
upper Missouri River demonstrate very
high growth rates relative to other
populations (Northcote 1995, p. 157).
Experimental evidence obtained by
growing fish from populations under
similar conditions would be needed to
measure the relative influence of
genetics (local adaptation) versus
environment.
We conclude that the occurrence of
Arctic grayling in the upper Missouri
River is biogeographically important to
the species, that grayling there have
occupied a warmer and more temperate
setting that is distinctly different from
the ecological settings relative to the rest
of the species (see Table 2, above), and
that they have been on a different
evolutionary trajectory for at least
10,000 years. We conclude that these
differences are significant because they
may provide the species with additional
evolutionary resiliency in the future in
light of the changing climate.
Consequently, we believe that Arctic
grayling in the upper Missouri River
occupy a unique ecological setting for
the species.
Gap in the Range
Arctic grayling in Montana (southern
extent is approximately 44°36′23″ N
latitude) represent the southern-most
extant population of the species’
distribution since the Pleistocene
glaciation. The next-closest native
Arctic grayling population outside the
Missouri River basin is found in the
Pembina River (approximately
52°55′6.77″ N latitude) in central
Alberta, Canada, west of Edmonton
(Blackburn and Johnson 2004, pp. ii, 17;
ASRD 2005, p. 6). The Pembina River
drains into Hudson Bay and is thus
disconnected from the Missouri River
basin. Loss of the native Arctic grayling
of the upper Missouri River would shift
the southern distribution of Arctic
grayling by more than 8° latitude (about
500 miles). Such a dramatic range
constriction would constitute a
significant geographic gap in the
species’ range and would eliminate a
genetically distinct group of Arctic
grayling, which may limit the species’
ability to cope with future
environmental change.
Marginal populations, defined as
those on the periphery of the species’
range, are believed to have high
conservation significance (Mitikka et al.
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2008; Gibson et al. 2009, entire; Haak et
al. 2010, entire; Osborne et al. 2012).
Peripheral populations may occur in
suboptimal habitats and thus be
subjected to very strong selective
pressures (Fraser 2000, p. 50).
Consequently, individuals from these
populations may contain adaptations
that may be important to the taxon in
the future. Lomolino and Channell
(1998, p. 482) hypothesize that because
peripheral populations should be
adapted to a greater variety of
environmental conditions, then they
may be better suited to deal with
anthropogenic (human-caused)
disturbances than populations in the
central part of a species’ range. Arctic
grayling in the upper Missouri River
have, for millennia, existed in a climate
warmer than that experienced by the
rest of the taxon. If this selective
pressure has resulted in adaptations to
cope with increased water temperatures,
then the population segment may
contain genetic resources important to
the taxon. For example, if northern
populations of Arctic grayling are less
suited to cope with increased water
temperatures expected under climate
warming, then Missouri River Arctic
grayling might represent an important
population for reintroduction in those
northern regions. We believe that Arctic
grayling’s occurrence at the
southernmost extreme of the range in
the upper Missouri River contributes to
the resilience of the overall taxon
because these peripheral populations
may possess increased adaptability
relative to the rest of the taxon.
Only Surviving Natural Occurrence of
the Taxon That May Be More Abundant
Elsewhere as an Introduced Population
Outside of Its Historical Range
This criterion does not directly apply
to the Arctic grayling in the upper
Missouri River because it is not the only
surviving natural occurrence of the
taxon; there are native Arctic grayling
populations in Canada, Alaska, and
Asia.
Differs Markedly in Its Genetic
Characteristics
Differences in genetic characteristics
can be measured at the molecular,
genetic, or phenotypic level. Three
different types of molecular markers
(allozymes, mtDNA, and microsatellites)
demonstrate that Arctic grayling from
the upper Missouri River are genetically
different from those in Canada, Alaska,
and Asia (Everett 1986, pp. 79–80;
Redenbach and Taylor 1999, p. 23;
Stamford and Taylor 2004, p. 1538;
Peterson and Ardren 2009, pp. 1764–
1766; USFWS, unpublished data). These
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data confirm the reproductive isolation
among populations that establishes the
discreteness of Missouri River Arctic
grayling under the DPS policy. Here, we
speak to whether these data also
establish significance.
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Allozymes
Using allozyme data, Everett (1986,
entire) found marked genetic differences
among Arctic grayling collected from
the Chena River in Alaska; those
descended from fish native to the
Athabasca River drainage in the
Northwest Territories, Canada; and
native upper Missouri River drainage
populations or populations descended
from them (see Leary 2005, pp. 1–2).
The Canadian population had a high
frequency of two unique alleles (forms
of a gene), which strongly differentiated
them from all the other samples (Everett
1986, p. 44). With the exception of one
introduced population in an irrigation
canal (Sunnyslope canal) in Montana
that is believed to have experienced
extreme genetic bottlenecks, the Chena
River (Alaskan) fish were highly
divergent from all the other samples as
they possessed an unusually low
frequency of a specific allele (Everett
1986, p. 60; Leary 2005, p. 1), and
contained a unique variant of another
allele (Leary 2005, p. 1). Overall, each
of the four native Missouri River
populations examined (Big Hole, Miner,
Mussigbrod, and Centennial Valley)
exhibited statistically significant
differences in allele frequencies relative
to both the Chena River (Alaska) and
Athabasca River (Canada) populations
(Everett 1986, pp. 15, 67).
Combining the data of Everett (1986,
entire), Hop and Gharrett (1989, entire),
and Leary (1990, entire) provides
information from 21 allozyme loci
(genes) from five native upper Missouri
River drainage populations, five native
populations in the Yukon River
drainage in Alaska, and the one
population descended from the
Athabasca River drainage in Canada
(Leary 2005, pp. 1–2). Examination of
the genetic variation in these samples
indicated that most of the genetic
divergence is due to differences among
drainages (29 percent) and
comparatively little (5 percent) results
from differences among populations
within a drainage (Leary 2005, p. 1).
Mitochondrial DNA
Analysis using mtDNA indicates that
Arctic grayling in North America
represent at least three evolutionary
lineages that are associated with distinct
glacial refugia (Redenbach and Taylor
1999, entire; Stamford and Taylor 2004,
entire). Arctic grayling in the upper
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Missouri River basin belong to the socalled North Beringia lineage
(Redenbach and Taylor 1999, pp. 27–28;
Samford and Taylor 2004, pp. 1538–
1540) because they possess a form of
mtDNA that was generally absent from
populations collected from other
locations within the species’ range in
North America (Redenbach and Taylor
1999, pp. 27–28; Stamford and Taylor
2004, p. 1538). The notable exceptions
were that some fish from the lower
Peace River drainage in British
Columbia, Canada, and all sampled
individuals from the Saskatchewan
River drainage Saskatchewan, Canada,
also possessed this form of mtDNA
(Stamford and Taylor 2004, p. 1538).
A form of mtDNA common in upper
Missouri River Arctic grayling, which
occurs at lower frequencies in other
populations, indicates that Arctic
grayling native to the upper Missouri
River drainage probably originated from
a glacial refuge in the drainage and
subsequently migrated northwards
when the Missouri River temporarily
flowed into the Saskatchewan River and
was linked to an Arctic drainage (Cross
et al. 1986, pp. 374–375; Pielou 1991, p.
195). When the Missouri River began to
flow southwards because of the advance
of the Laurentide ice sheet (Cross et al.
1986, p. 375; Pileou 1991, p. 10), the
Arctic grayling in the drainage became
physically and reproductively isolated
from the rest of the species’ range (Leary
2005, p. 2; Campton 2004, p. 6), which
would have included those populations
in Saskatchewan. Alternatively, the
Missouri River Arctic grayling could
have potentially colonized
Saskatchewan or the Lower Peace River
(in British Columbia) or both postglacially (Stamford 2001, p. 49) via a
gap in the Cordilleran and Laurentide
ice sheets (Pielou 1991, pp. 10–11),
which also might explain the low
frequency ’Missouri River’’ mtDNA in
Arctic grayling in the Lower Peace River
and Upper Yukon River.
We do not interpret the observation
that Arctic grayling in Montana and
Saskatchewan, and to lesser extent those
from the Lower Peace and Upper Yukon
River systems, share a mtDNA
haplotype to mean that these groups of
fish are genetically identical. Rather, we
interpret it to mean that these fish
shared a common ancestor tens to
hundreds of thousands of years ago.
Microsatellite DNA
Recent analysis of microsatellite DNA
(highly variable portions of nuclear
DNA) showed substantial divergence
between Arctic grayling in Missouri
River and Saskatchewan populations
(Peterson and Ardren 2009, entire). This
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49395
divergence between populations was
measured in terms of allele frequencies,
using a metric called Fst (Allendorf and
Luikart 2007, pp. 52–54, 198–199). An
analogous metric, named Rst, also
measures genetic differentiation
between populations based on
microsatellite DNA, but differs from Fst
in that it also considers the size
differences between alleles (Hardy et al.
2003, p. 1468). An Fst or Rst of 0
indicates that populations are the same
genetically, whereas a value of 1
indicates the populations share no
genetic material at the markers being
surveyed. Fst values range from 0.13 to
0.31 (average 0.18) between Missouri
River and Saskatchewan populations
(Peterson and Ardren 2009, pp. 1758,
1764–1765), whereas Rst values range
from 0.47 to 0.71 (average 0.54) for the
same comparisons (Peterson and Ardren
2009, pp. 1758, 1764–1765). These
values indicate that the two populations
differ significantly in allele frequency
and also in the size of those alleles. This
outcome indicates that the observed
genetic differences are due to
mutational differences, which suggests
the groups may have been separated for
millennia (Peterson and Ardren 2009,
pp. 1767–1768).
Analysis of Arctic grayling
populations from Alaska, Canada, and
the Missouri River basin using nine of
the same microsatellite loci as Peterson
and Ardren (2009, entire) further
supports the distinction of Missouri
River Arctic grayling relative to
populations elsewhere in North
America (USFWS, unpublished data).
This analysis clearly separated sample
fish from 21 populations into two
clusters: One cluster representing
populations from the upper Missouri
River basin, and another cluster
representing populations from across
Canada and Alaska (USFWS,
unpublished data). Divergence in size
among these alleles further supports the
distinction between Missouri River
Arctic grayling and those in Canada and
Alaska (USFWS, unpublished data). The
interpretation of these data is that the
Missouri River populations and the
Canada/Alaska populations are highly
genetically distinct at the microsatellite
loci considered.
Phenotypic Characteristics Influenced
by Genetics—Meristics
Phenotypic variation can be evaluated
by counts of body parts (i.e., meristic
counts of the number of gill rakers, fin
rays, and vertebrae characteristics of a
population) that can vary within and
among species. These meristic traits are
influenced by both genetics and the
environment (Allendorf and Luikart
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2007, pp. 258–259). When the traits are
controlled primarily by genetic factors,
then meristic characteristics can
indicate significant genetic differences
among groups. Arctic grayling north of
the Brooks Range in Alaska and in
northern Canada had lower lateral line
scale counts than those in southern
Alaska and Canada (McCart and Pepper
1971, entire). These two scale-size
phenotypes are thought to correspond to
fish from the North and South Beringia
glacial refuges, respectively (Stamford
and Taylor 2004, p. 1545). Arctic
grayling from the Centennial Valley had
a phenotype intermediate to the largeand small-scale types (McCart and
Pepper 1971, pp. 749, 754). Arctic
grayling populations from the Missouri
River (and one each from Canada and
Alaska) could be correctly assigned to
their group 60 percent of the time using
a suite of seven meristic traits (Everett
1986, pp. 32–35). Those native Missouri
River populations that had high genetic
similarity also tended to have similar
meristic characteristics (Everett 1986,
pp. 80, 83).
Arctic grayling from the Big Hole
River showed marked differences in
meristic characteristics relative to two
populations from Siberia, and were
correctly assigned to their population of
origin 100 percent of the time (Weiss et
al. 2006, pp. 512, 515–516, 518). The
populations that were significantly
different in terms of their meristic
characteristics also exhibited differences
in molecular genetic markers (Weiss et
al. 2006, p. 518).
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Inference Concerning Genetic
Differences in Arctic Grayling of the
Missouri River Relative to Other
Examples of the Taxon
We believe the differences between
Arctic grayling in the Missouri River
and sample populations from Alaska
and Canada measured using allozymes
(Everett 1986, entire; Leary 2005,
entire), mitochondrial DNA (Redenbach
and Taylor 1999, entire; Stamford and
Taylor 2004, entire), and microsatellite
DNA markers (Peterson and Ardren
2009, pp. 1764–1766; USFWS,
unpublished data) represent ‘‘marked
genetic differences’’ in terms of the
extent of differentiation (e.g., Fst, Rst)
and the importance of that genetic
legacy to the rest of the taxon. The
presence of morphological
characteristics separating Missouri River
Arctic grayling from other populations
also likely indicates genetic differences,
although this conclusion is based on a
limited number of populations (Everett
1986, pp. 32–35; Weiss et al. 2006,
entire), and we cannot entirely rule out
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the influence of environmental
variation.
The intent of the DPS policy and the
Act is to preserve important elements of
biological and genetic diversity, not
necessarily to preserve the occurrence of
unique alleles in particular populations.
In Arctic grayling of the Missouri River,
the microsatellite DNA data indicate
that the group is evolving
independently from the rest of the
species. The extirpation of this group
would mean the loss of the genetic
variation in one of the two most distinct
groups identified in the microsatellite
DNA analysis, and the loss of the future
evolutionary potential that goes with it.
Thus, the genetic data support the
conclusion that Arctic grayling of the
upper Missouri River represent a unique
and irreplaceable biological resource of
the type the Act was intended to
preserve. Thus, we conclude that
Missouri River Arctic grayling differ
markedly in their genetic characteristics
relative to the rest of the taxon.
Upper Missouri River Arctic grayling
satisfy the significance criteria outlined
in the Services’ DPS policy because they
occur in a unique ecological setting, are
separated from other Arctic grayling
populations by a large gap in their
range, and differ markedly in their
genetic characteristics relative to other
Arctic grayling populations. Therefore,
we consider the Arctic grayling in the
upper Missouri River basin significant
to the taxon to which it belongs under
the Service’s DPS policy.
Determination of Distinct Population
Segment
We find that a population segment
that includes all native ecotypes of
Arctic grayling in the upper Missouri
River basin satisfies the discreteness
standard of the DPS policy. The segment
is physically isolated, and genetic data
indicate that Arctic grayling in the
Missouri River basin have been
separated from other populations for
thousands of years. The population
segment occurs in an isolated
geographic area far south of all other
Arctic grayling populations worldwide,
and we find that loss of this population
segment would create a significant gap
in the species’ range. Molecular genetic
data clearly differentiate Missouri River
Arctic grayling from other Arctic
grayling populations, including those in
Canada and Alaska.
Based on the best scientific and
commercial information available, as
described above, we find that, under the
Service’s DPS policy, upper Missouri
River Arctic grayling are discrete and
are significant to the taxon to which
they belong. Because the upper Missouri
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River population of Arctic grayling is
both discrete and significant, it qualifies
as a DPS under the Act.
As we described above, we are
including introduced Arctic grayling
populations that occur in lakes in the
upper Missouri River basin as part of
the DPS. The Service has interpreted the
Act to provide a statutory directive to
conserve species in their native
ecosystems (49 FR 33885; August 27,
1984) and to conserve genetic resources
and biodiversity over a representative
portion of a taxon’s historical
occurrence (61 FR 4722; February 7,
1996). The introduced Arctic grayling
populations occur within the
boundaries of the upper Missouri River
basin and represent moderate to high
levels of genetic diversity from within
the basin. The future adaptive
capabilities represented by this genetic
diversity have conservation value,
particularly given a changing climate.
We define the historical range of this
population segment to include the major
streams, lakes, and tributary streams of
the upper Missouri River (mainstem
Missouri, Smith, Sun, Beaverhead,
Jefferson, Big Hole, and Madison Rivers,
as well as their key tributaries, as well
as a few small lakes where Arctic
grayling are or were believed to be
native (Elk Lake, Red Rock Lakes in the
Centennial Valley, Miner Lake, and
Mussigbrod Lake, all in Beaverhead
County, Montana)). We define the
current range of the DPS to consist of
extant native populations in the Big
Hole River, Miner Lake, Mussigbrod
Lake, Madison River-Ennis Reservoir,
and Centennial Valley, as well as all
known introduced populations within
the upper Missouri River basin. We refer
to this entity as the Upper Missouri
River DPS of Arctic grayling. The
remainder of this finding will thus focus
on the population status of and
potential threats to this entity.
Population Status and Trends of
Populations in the Upper Missouri River
DPS
The Upper Missouri River DPS of
Arctic grayling is comprised of 20
populations, including 2 fluvial
populations and 16 adfluvial
populations. Two other populations
(Centennial Valley and Madison River/
Ennis Reservoir) appear to exhibit both
fluvial and adfluvial components (Table
3). Arctic grayling from the Centennial
Valley (Long Creek) and Ennis
Reservoir/Madison River (mainstem
Madison River) have been documented
well past the spawning period through
autumn. These occurrences are more
prevalent in Long Creek in the
Centennial Valley than in the Madison
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River population and do not appear to
be linked to individual Arctic grayling
seeking thermal refugia during summer
(Montana Arctic Grayling Workgroup
(AGW) 1995; p. 1; Cayer 2014a, pers.
comm.; MFISH 2014b, unpublished
data). These occurrences include
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multiple age classes (Age-1 to Age-3) of
Arctic grayling in both Long Creek and
the Madison River and are located in
stream reaches that are considerable
distances (up to 15 miles in the Madison
River) from adfluvial habitats (Cayer
2014a, pers. comm.; MFISH 2014b,
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unpublished data). Eighteen of the 20
populations occur solely on Federal or
majority Federal land; the remaining
two (Big Hole River and Ennis
Reservoir/Madison River) occur on
primarily private land.
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the two fluvial populations varies from
about one hundred to several thousand
E:\FR\FM\20AUP2.SGM
Estimated abundance of
reproductively mature individuals in
PO 00000
EP20AU14.001
Agnes Lake
Odell Lake
Bobcat Lake
Schwinegar Lake
Pintlar Lake
Deer Lake
Emerald Lake
Grayling Lake
Hyalite Lake
Big Hole
Big Hole
Big Hole
Big Hole
Big Hole
Gallatin
Gallatin
Gallatin
Gallatin
Federal
Federal
Federal
Federal
Federal
Federal
Federal
Federal
Federal
44
13
2
2
16
5
6
1
64
A
A
A
A
A
A
A
A
A
577 (222 - 00)
252 (114- oo)
-
-
-24,000
2,481-8251
1084- 3604
1972
2001-2003
2001-2003
Common
Common
800- 1,100
-
499''* (5-
1989-2002
Abundant
Rare
-
1998-2012
Stable
Stable
Stable
Stable
Stable
Stable
Stable
Stable
Stable
1341)
Diversion Lake
Gibson Reservoir
Lake Levale
Park Lake
Grebe Lake
Sun
Sun
Sun
Missouri
Madison
Federal
Federal
Federal
Federal
Federal
30
521
5
13
59
A
A
A
A
A
Rare
Abundant
Common
Infinite'
-27,000
1999-2003 (Ne)
1954 (census)
Stable
Stable
Stable
•Habitat extent is the amount of habitat currently being used by Arctic grayling for some portion of their life history. It does not mean the amount specified is occupied continuously.
~. denotes effective population size; a theoretical size of a population that would result in the same level of inbreeding or genetic drift as that of the population under study. For more information, see
discussion of effective population size below in Factor E.
"Nb denotes the number of breeding adults that contributed genetics to a sample of offspring from a given population.
dPopulation size of reproductively mature individuals (not to be confused with total annual census population size which includes adults and juveniles)estimated from N. assuming Ne IN .07 (minimum
estimate) and .23 (maximum estimate). These two values represent the range of median N. IN ratios for salmonids cited in Palstra and Fraser 2012.
•Qualitative descriptors are from Montana Fish, Wildlife, and Parks MFISH database and are based on biomass estimates where available, or biologist observationsand professional biological judgment.
rApproximate date to which theN., Nb, or annual census population size refers. Biological dates for Ne or Nb estimates refer to the generation of breeders that produced the sample of offspring that were
genotyped.
gPopulation trends are derived from genetic data or population monitoring data or a combination of these two data types, if present.
'Point estimate for Grebe Lake N. was negative, indicating no evidence for any disequilibrium caused by genetic drift due to finite number of parents (Perterson and Arden 2009, p. 1767).
"The Nb estimate for the Big Hole River in 2012 is reported as a range because of uncertainty in the frequency rate of rare alleles in the analysis.
'"The Nb estimate for Hyalite Lake is reported as the mean number (and range) of adult spawning individuals observed in the spawning run in Hyalite Creek from 1998-2012.
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Table 3. Characteristics of populations within the Upper Missouri River DPS of Arctic grayling. Rows highlighted in gray indicate populations occurring in native habitat. For ecotype,
F =fluvial, A= adfluvial, and F/A both fluvial and adfluvial characteristics present.
Extent"
Nb(95% Cl or
Qualitative
Stream miles
Federal Register / Vol. 79, No. 161 / Wednesday, August 20, 2014 / Proposed Rules
Arctic grayling (Table 3). Where
quantitative data are available,
estimated abundance of mature
individuals in adfluvial populations
(including the two populations
exhibiting both life histories) varies
from a few hundred to around 25,000
Arctic grayling. Most populations are
currently stable or increasing in
abundance, with the exception of the
Ennis Reservoir/Madison River
population (Table 3).
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Distinct Population Segment Five-Factor
Analysis
Since the Arctic grayling in the upper
Missouri River basin qualifies as a DPS,
we will now evaluate its status with
regard to its potential for listing as
endangered or threatened based on the
five factors enumerated in section 4(a)
of the Act. Our evaluation of the Upper
Missouri River DPS of Arctic grayling
follows.
Summary of Information Pertaining to
the Five Factors
Section 4 of the Act (16 U.S.C. 1533)
and implementing regulations (50 CFR
424) set forth procedures for adding
species to, removing species from, or
reclassifying species on the Federal
Lists of Endangered and Threatened
Wildlife and Plants. Under section
4(a)(1) of the Act, a species may be
determined to be endangered or
threatened based on any of the
following 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.
In making this finding, information
pertaining to the Upper Missouri River
DPS of Arctic grayling in relation to the
five factors provided in section 4(a)(1) of
the Act is discussed below. In
considering what factors might
constitute threats, we must look beyond
the mere exposure of the species to the
factor to determine whether the species
responds to the factor in a way that
causes actual impacts to the species. If
there is exposure to a factor, but no
response, or only a positive response,
that factor is not a threat. If there is
exposure and the species responds
negatively, the factor may be a threat
and we then attempt to determine how
significant a threat it is. If the threat is
significant, it may drive or contribute to
the risk of extinction of the species such
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that the species warrants listing as
endangered or threatened as those terms
are defined by the Act. This does not
necessarily require empirical proof of a
threat. The combination of exposure and
some corroborating evidence of how the
species is likely impacted could suffice.
The mere identification of factors that
could impact a species negatively is not
sufficient to compel a finding that
listing is appropriate; we require
evidence that these factors are operative
threats that act on the species to the
point that the species meets the
definition of an endangered or
threatened species under the Act.
In making our revised 12-month
finding on the petition, we consider and
evaluate the best available scientific and
commercial information. This
evaluation includes all factors we
previously considered in the 2010
finding and, at the end of this analysis,
explains how the Services’ conclusions
differ now.
Factor A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range
Curtailment of Range and Distribution
The range and distribution of fluvial
Arctic grayling in the upper Missouri
River basin was reduced over the past
100 years (Kaya 1992, p. 51), primarily
due to historical habitat fragmentation
by dams and irrigation diversions and
by habitat degradation or modification
from unregulated land use (Vincent
1962, pp. 97–121). Fluvial Arctic
grayling typically need large expanses of
connected habitat to fulfill their lifehistory stages (Armstrong 1986, p. 8).
For example, fluvial Arctic grayling in
the Big Hole River have been
documented migrating over 60 miles (97
km) between overwintering, spawning,
and foraging habitats (Shepard and
Oswald 1989, pp. 18–21, 27). These past
reductions in range and distribution
reproductively isolated fluvial Arctic
grayling populations within the basin
(Peterson and Ardren 2009, p. 1770).
Although the range and distribution
of fluvial Arctic grayling has contracted
from historical levels, expression of the
fluvial life history is represented, at
least in part, in four Arctic grayling
populations within the Upper Missouri
River DPS. Whether strictly fluvial (e.g.,
Big Hole and Ruby River) or partially
fluvial (e.g., Centennial Valley (Long
Creek) and Ennis Reservoir/Madison
River (mainstem Madison River)), these
populations occur in four watersheds
where large reaches of connected habitat
remain and still permit the expression
of the fluvial life history, despite the
presence of mainstem dams in three of
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four watersheds (Kaya 1992, entire; see
Figure 1). Thus, despite historical
curtailment of range, the amount of
connected habitat in some systems is
adequate to permit the expression of the
fluvial life history.
Of the four Arctic grayling
populations still expressing a fluvial life
history, three of four populations (Big
Hole River, Centennial Valley, and Ruby
River) are currently increasing in
abundance (see Table 3). In each of
these populations, as abundance
increases, there is a corresponding
increase in distribution. Natural
reproduction is occurring in all three of
these populations. In the Big Hole River
and the Centennial Valley, remote site
incubators (RSIs) have been used as a
conservation tool to help facilitate
increased abundance and distribution of
Arctic grayling. Thus, observed
increases in abundance and distribution
may be partially attributable to the use
of RSIs (for more in-depth discussion on
RSI use, see ‘‘Native Arctic Grayling
Genetic Reserves and Translocation,’’
below). Given the above information, it
appears that three of four fluvial, or
partly fluvial, populations are viable
and have the necessary configuration
and amount of habitat to fulfill their
life-history needs. Thus, effects of past
range curtailment on the fluvial
component of Arctic grayling in the
upper Missouri River basin are present,
but there appears to be sufficient
adequate habitat remaining to support
expression of the fluvial life history.
Adfluvial Arctic grayling populations
in the upper Missouri River basin are
present in all lakes originally thought to
have had native populations historically
(Miner, Mussigbrod, Upper Red Rock,
and Elk Lakes (present but not included
in Table 3, above, because of uncertain
viability)). Thus, there has been no
contraction of the range of adfluvial
populations. Given the above
information, curtailment of range and
distribution is not precluding the
expression of either fluvial or adfluvial
life history. Although curtailment of
range and distribution occurred
historically, Arctic grayling populations
are still present in 7 of 10 historically
occupied watersheds in the upper
Missouri River basin (see ‘‘Drainage’’
column in Table 3). Accordingly, we
have no evidence that curtailment of
range and distribution is a current threat
to the DPS. In addition, we have no
information suggesting curtailment of
range and distribution will be a threat
in the future.
Dams on Mainstem Rivers
Much of the historical range of the
Upper Missouri River DPS of Arctic
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Federal Register / Vol. 79, No. 161 / Wednesday, August 20, 2014 / Proposed Rules
grayling has been altered by the
construction of dams and reservoirs
(Kaya 1990, pp. 51–52; Kaya 1992, p.
57). The construction of large dams on
mainstem river habitats throughout the
upper Missouri River system fragmented
river corridors necessary for the
expression of Arctic grayling migratory
life histories in some systems.
Construction of dams that obstructed
fish passage on the mainstem Missouri
River (Hauser, Holter, Canyon Ferry,
and Toston dams), Madison River
(Madison-Ennis, Hebgen dams),
Beaverhead River and its tributary Red
Rock River (Clark Canyon, Lima dams),
Ruby River (Ruby dam), and Sun River
(Gibson dam) all likely contributed to
the historical decline of fluvial Arctic
grayling in the DPS (Vincent 1962, pp.
127–128; Kaya 1992, p. 57). Lack of fish
passage at these dams contributed to the
extirpation of fluvial Arctic grayling
from some waters by blocking migratory
corridors (Vincent 1962, p. 128),
curtailing access to important spawning
and rearing habitats, and impounding
water over former spawning locations
(Vincent 1962, p. 128). Most dams
within the upper Missouri River basin
were constructed between 1905 and
1960 (Kaya 1990, entire).
Despite the construction of multiple
dams throughout the historical range of
Arctic grayling, multiple populations, or
portions of populations, of the fluvial
ecotype are still represented in the DPS.
These populations reside in areas where
sufficient quantity and quality of habitat
exist and permit the expression of this
life history. In some cases, dams may be
providing a benefit, because currently
many of the dams that historically
affected fluvial Arctic grayling
populations are now precluding
invasion by nonnative fish from
downstream sources. For example, Lima
Dam in the Centennial Valley is
currently precluding brown trout
invasion from downstream sources
(Mogen 2014, pers. comm). Currently,
there are five Arctic grayling
populations within the DPS that occur
above mainstem dams (Centennial
Valley, Ruby River, Hyalite Lake,
Diversion Lake, and Gibson Reservoir)
with at least one nonnative fish species
occurring downstream of these dams
(MFISH 2014d, unpublished data).
Some reservoirs created by dams are
currently being used by Arctic grayling
as overwintering, rearing and foraging
areas. Both adult and juvenile Arctic
grayling use Ennis Reservoir for
overwintering, rearing, and foraging
(Byorth and Shepard 1990, entire). In
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the Centennial Valley, Arctic grayling
have recently been detected in Lima
Reservoir (MFISH 2014e, unpublished
data). The movements of Arctic grayling
within and out of Lima Reservoir are
unknown; however, Lima Reservoir is a
large reservoir and, as such, is likely
used for overwintering purposes.
Arctic grayling have been
documented in stream and river reaches
below some dams, most likely
indicating downstream passage of fish
over or through dams. These fish are
essentially ‘‘lost’’ to the population
residing above the dam, because none of
the mainstem river dams in the upper
Missouri River basin provides upstream
fish passage. Substantial losses from a
population resulting from downstream
entrainment of fish through dams could
cause declines in reproductive potential
and abundance in the reservoir
population above the dam (Kimmerer
2008, entire). However, it is unknown
what entrainment rates currently are in
populations residing near dams. Rate of
entrainment is likely dependent on a
number of factors, including dam
operations, season, water conditions in
the reservoir, initial population size
above the dam, etc. Recent monitoring
data and angler reports of Arctic
grayling observed downstream of
reservoirs supporting Arctic grayling
populations are sporadic (Horton 2014c,
pers. comm.; SSA 2014); thus it appears
the threat of mainstem dams is likely
affecting some individuals, but not
affecting populations or the DPS as a
whole.
Historically, operational practices at
Madison Dam have likely affected the
Arctic grayling population in Ennis
Reservoir/Madison River. A population
decline in Arctic grayling appeared to
coincide with a reservoir drawdown in
the winter of 1982–1983 (Byorth and
Shepard 1990, pp. 52–53). This
drawdown likely affected the forage
base, rearing habitat, and spawning
cycle of Arctic grayling in the reservoir.
However, under a new licensing
agreement dated September 27, 2000,
between the Federal Energy Regulatory
Commission and Ennis Dam operators,
such substantial drawdowns in
elevation of Ennis Reservoir are no
longer permitted (Clancey 2014, pers.
comm.).
Given the above information,
mainstem dams were a historical threat
to Arctic grayling populations in the
upper Missouri River basin. Dams still
impact individuals, because some Arctic
grayling are currently being entrained
and lost from their source population. In
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Ennis Reservoir, the new licensing
agreement is expected to reduce the
effects of dam operations on the Arctic
grayling population. Most Arctic
grayling populations residing above
dams are stable or increasing; thus, it
does not appear this impact is acting at
the population or DPS level. We have no
information to conclude that mainstem
dams will be a threat in the future at the
population or DPS level.
Water Management in the Upper
Missouri River Basin
The predominant use of private lands
in the upper Missouri River basin is
irrigated agriculture and ranching.
These activities have historically had
significant effects on aquatic habitats,
primarily changes in water availability
and alteration of the structure and
function of aquatic habitats. Changes in
water availability can affect Arctic
grayling reproduction, survival, and
movements among habitat types (Kaya
1990, entire).
In contrast to most of the Arctic
grayling populations in the Upper
Missouri River DPS that occur on
Federal land, the fluvial population of
Arctic grayling in the Big Hole River
occurs on primarily (∼90 percent)
private land. Thus, any conservation
efforts conducted in the Big Hole River
Valley need support from involved
agencies and private landowners. In
2006, a candidate conservation
agreement with assurances (CCAA;
Montana Fish, Wildlife, and Parks et al.
2006, entire) was developed for Arctic
grayling in the Big Hole River. The
conservation goal of this CCAA is to
secure and enhance the fluvial
population of Arctic grayling in the
upper Big Hole River drainage.
Conservation projects conducted under
the CCAA are prioritized and guided by
the Big Hole Arctic Grayling Strategic
Habitat Conservation Plan (SHCP) (for
more specific information, see
‘‘Conservation Efforts to Reduce Habitat
Destruction, Modification, or
Curtailment of Its Range,’’ below).
Since 2006, many conservation and
restoration projects have been
completed in the upper Big Hole River
under the direction of the CCAA and
SHCP (Table 4). Below, we describe and
evaluate the implementation and
effectiveness of these projects relative to
the potential stressors analyzed under
Factor A for the Big Hole River
population. We also analyze the effects
of potential stressors under Factor A for
the other Arctic grayling populations in
the DPS.
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TABLE 4—CONSERVATION PROJECTS AND RESULTS, AND ARCTIC GRAYLING RESPONSE IN THE BIG HOLE RIVER SINCE
IMPLEMENTATION OF THE BIG HOLE CCAA IN 2006
[All information on conservation projects and conservation results cited from the Big Hole Arctic Grayling Strategic Habitat Conservation Plan]
A ...................
Conservation projects a
Conservation result
Arctic grayling response
Dams/habitat fragmentation.
Fish ladders: 41 ............
Bridges: 7 .....................
Grade control structures: 2.
PODs: 343 of 504 with
signed SSPs.
Irrigation improvements:
88.
Water measuring devices: 67.
Stock water systems:
63.
Stream restoration: 26
miles.
Rock Creek restoration
Stream miles (%) accessible to
grayling b:
• Tier
I82(98%;
preCCAA=87%)..
• Tier
II61(67%;
preCCAA=27%)..
• Tier
III32(20%:
preCCAA=6%)..
• Achievement of instream flow
goals increased from 50%
(pre-CCAA) to 78% (postCCAA).
• Landowner contributions to
streamflow increasing as # of
PODs with signed SSPs increase [landowner contribution to instream flows in Big
Hole River (pre-2006 = 0 cfs;
2013 = 250 cfs)].
• Temperature reductions in tributaries (see Rock Creek example
below).
Pre-restoration (2007): .....................
• 36 days max. temp >70 °F ...
• 16 days max. temp >77 °F ...
Post-restoration (2013): ...................
• 0 days max temp. >70 °F .....
• Number of breeding adults has increased from ∼100 (2007–2011)
to 500–900 c (2013) (Leary 2014,
unpublished data).
Dewatering/Thermal
stress.
Threat factor
Stressor
Fish screens: 2 .............
Prioritized monitoring
protocol.
• No entrainment
documented
since 2010.
• Observed low
entrainment rates
in unscreened
ditches (73 Arctic
grayling/138 ditch
miles).
Riparian habitat loss.
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Entrainment ..
Stream restoration: 26
miles.
Riparian fencing: 108
miles.
Stock water systems:
63.
Grazing mgmt. plans:
21 landowners
(85,000 ac.).
Noxious weed management.
Willow planting (72,200
planted).
• Arctic grayling abundance d (catch
per unit effort) increased from 0.2
fish/mile (2008) to 1.4 fish/mile
(2012) in the CCAA monitoring
reaches of the mainstem Big Hole
River (MFWP 2013a, unpublished
data).
• Arctic grayling abundance d (catch
per unit effort) increased from 2.9
fish/mile (2008) to 7.4 fish/mile
(2012) in the CCAA monitoring
tributaries (MFWP 2013a, unpublished data).
• Arctic grayling distribution has increased 4 miles in Rock Creek
(young-of-year and Age 1+) and 2
miles in Big Lake Creek (Age 1+)
since 2006 (SHCP 2013, p. 12).
• 110 miles (65%) of riparian habitat on enrolled lands improving.
• 15% increase in
sustainable riparian areas from
32% (2006) to
47% (2013).
• Adaptive management in place
to address nonimproving areas.
a PODs = Points of Diversion, SSPs = Site-specific plans; b Tier I is core spawning, rearing and adult habitat that is currently occupied by Arctic
grayling, Tier II is periphery habitat intermittently used by Arctic grayling, Tier III is suitable, but currently unoccupied historical habitat; c The estimate of number of breeding adults in the Big Hole River in 2013 is reported as a range because of uncertainty in the frequency rate of rare
alleles in the analysis; d Abundance estimates from 2013 were lower than those reported for 2012 likely due to unusually high flows (3X normal)
concurrent with fall sampling that likely decreased capture efficiency, resulting in lower abundance estimates in 2013.
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Habitat Fragmentation/Smaller Seasonal
Barriers
Big Hole River: Smaller dams or
diversions associated with irrigation
structures historically posed a threat to
Arctic grayling migratory behavior,
especially in the Big Hole River
drainage. In the Big Hole River,
numerous diversion structures have
been identified as putative fish
migration barriers (Petersen and
Lamothe 2006, pp. 8, 12–13, 29) that
may limit the ability of Arctic grayling
to migrate to spawning, rearing, or
sheltering habitats under certain
conditions. As with the larger dams,
these smaller fish passage barriers can
reduce reproduction (access to
spawning habitat is blocked), reduce
growth (access to feeding habitat is
blocked), and increase mortality (access
to refuge habitat is blocked).
Historically, these types of barriers were
numerous and widespread across the
Big Hole River drainage.
Currently, habitat fragmentation due
to irrigation diversion structures in the
Big Hole is being systematically reduced
under the CCAA for Fluvial Arctic
Grayling in the upper Big Hole River
(hereafter, Big Hole CCAA or CCAA; for
more specific information, see
‘‘Conservation Efforts to Reduce Habitat
Destruction, Modification, or
Curtailment of Its Range’’) and Big Hole
Arctic Grayling SHCP. Since 2006, 41
fish ladders have been installed in the
mainstem Big Hole River and tributaries
(Table 4). Multiple culverts have been
replaced with bridges and several grade
control structures have been installed
(Table 4). As a result, no fish barriers
now exist in the mainstem upper Big
Hole River. Almost all (98 percent) of
tier I habitat and the majority (68
percent) of tier II habitat is connected
and accessible to Arctic grayling (Table
4): 67 miles of stream have been
reconnected in the Big Hole River
system since 2006 (MFWP 2014a,
unpublished data).
Other populations: Smaller fish
passage barriers also have been noted to
affect Arctic grayling in the Centennial
Valley (Unthank 1989, p. 9).
Historically, spawning Arctic grayling
migrated from the Jefferson River
system, through the Beaverhead River
and Red Rock River through the Red
Rock Lakes and into the upper drainage,
and then returned downstream after
spawning (Henshall 1907, p. 5). The
construction of a water control structure
(sill) at the outlet of Lower Red Rock
Lake in 1930 (and reconstruction in
1957 (USFWS 2009, p. 74)) created an
upstream migration barrier that blocked
these migrations (Unthank 1989, p. 10;
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Gillin 2001, p. 4–4). However, recent
changes in water management at the
Red Rock Lakes National Wildlife
Refuge (NWR) have resulted in yearround fish passage through the control
structure at the outlet of Lower Red
Rock Lake (West 2013, pers. comm.).
In Mussigbrod Lake, Arctic grayling
occasionally pass downstream over a
diversion structure at the lake outlet,
and become trapped in an isolated pool
(Olsen 2014, pers. comm.). During highsnowpack years, Arctic grayling likely
can swim back up to the lake from the
pool, but in low snowpack years, some
Arctic grayling perish when the isolated
pool dries up (Olsen 2014, pers. comm.).
However, this phenomenon has
occurred periodically in recent history
and has had no discernible impacts on
Arctic grayling abundance in
Mussigbrod Lake (Olsen 2014, pers.
comm.).
All 16 adfluvial Arctic grayling
populations in the upper Missouri River
basin occur on Federal land (U.S. Forest
Service) and are not influenced by
irrigation structures because none are
present. The effect of a barrier at the
outlet of Mussigbrod Lake is likely
impacting individuals, but not the
population because of the robust
population size in Mussigbrod Lake and
historical stability of that population
since the outlet structure was created.
Based on this information, we conclude
that the threats from habitat
fragmentation have been sufficiently
mitigated or minimized and are no
longer are acting as a stressor at the
population or DPS level.
Degradation of Riparian Habitat
Riparian corridors are important for
maintaining habitat for Arctic grayling
in the upper Missouri River basin, and
in general are critical for the ecological
function of aquatic systems (Gregory et
al. 1991, entire). Riparian zones are
important for Arctic grayling because of
their effect on water quality and water
temperature, and their role in
maintaining natural ecological process
responsible for creating and maintaining
necessary physical habitat features (i.e.,
pools, riffles, and scour areas) used by
the species to meet its life-history
requirements.
Big Hole: Arctic grayling abundance
in the upper Big Hole River is positively
related to the presence of overhanging
vegetation, primarily willows (Salix
spp.), that is associated with pool
habitat (Lamothe and Magee 2004, pp.
21–22). Removal of willows and
riparian clearing concurrent with
livestock and water management along
the upper Big Hole River has led to a
shift in channel form (i.e., braided
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channels becoming a single wide
channel), increased erosion rates,
reduced cover, increased water
temperatures, and reduced recruitment
of large wood debris into the active
stream channel (Confluence Consulting
et al. 2003, pp. 24–26). These factors
combine to reduce the suitability of the
habitat for species like Arctic grayling
(Hubert 1985, entire).
Currently, restoration of riparian areas
in the upper Big Hole River system is a
priority under the CCAA (for more
specific information, see ‘‘Conservation
Efforts to Reduce Habitat Destruction,
Modification, or Curtailment of Its
Range,’’ below). Since 2006, efforts to
restore and conserve riparian habitats
have been numerous and multi-faceted
(see Table 4). About 170 miles (274 km)
of riparian habitat are currently enrolled
in the Big Hole CCAA, out of a total of
about 340 miles (547 km) of total
riparian habitat in the CCAA
Management Area. Of the enrolled
riparian habitat, 65 percent (110 miles
(177 km)) is improving in condition, as
rated by a standardized riparian
protocol (NRCS 2004, entire). Further,
47 percent of enrolled riparian habitat
(80 miles (129 km)) is functioning at a
sustainable level, which is a 15 percent
increase in 5 years (MTFWP et al. 2006,
p. 92; see Table 4). A sustainable rating
indicates that the stream can access its
flood plain, transport its sediment load,
build banks, store water, and dissipate
flood energy in conjunction with a
healthy riparian zone (NRCS 2004, p. 7).
Riparian habitats are reassessed every 5
years and are scored on 10 stability and
sustainability metrics (for example,
stream incisement), with any reach
scoring at 80 percent or above rated as
sustainable (NRCS 2004, entire). In
addition, adaptive management within
the CCAA framework will allow for
reevaluation of conservation measures
being implemented in non-improving
habitat.
Other populations: In the Centennial
Valley, historical livestock grazing both
within the Red Rock Lakes NWR and on
adjacent private lands negatively
affected the condition of riparian
habitats on tributaries to the Red Rock
Lakes (Mogen 1996, pp. 75–77; Gillin
2001, pp. 3–12, 3–14). In general,
degraded riparian habitat limits the
creation and maintenance of aquatic
habitats, especially pools, which are
preferred habitats for adult Arctic
grayling (Lamothe and Magee 2004, pp.
21–22; Hughes 1992, entire), although
many spawning adult Arctic grayling in
Red Rock Creek outmigrate soon after
spawning and likely do not use
available pool habitat (Jordan 2014,
pers. comm.). Loss of riparian vegetation
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increases bank erosion, which can lead
to siltation of spawning gravels, which
may in turn harm Arctic grayling by
reducing the extent of suitable spawning
habitat and reducing survival of Arctic
grayling embryos already present in the
stream gravels.
Recently, the Red Rock Lakes NWR
acquired land on Red Rock Creek,
upstream of the refuge boundary (West
2014a, pers. comm.). Much of this
parcel was riparian habitat that was
historically heavily grazed; thus, the
refuge implemented a rest-rotation
grazing system where more durable
lands are grazed while more sensitive
lands (e.g., riparian areas) are rested for
up to 4 years. On average, grazing
intensities on the refuge have decreased
from 20,000 Animal Unit Months
(AUMs, number of cow/calf pairs
multiplied by the number of months
grazed) to about 5,000 AUMs. As a
result of these changes, riparian habitat
within the refuge has dramatically
improved (West 2014b, pers. comm.)
and is expected to continue improving
under the new grazing regime.
Concurrent with riparian improvement
within Red Rock Lakes NWR, the
number of adult Arctic grayling
migrating up Red Rock Creek to spawn
has increased from fewer than 500 to
more than 2,000 (Patterson 2014,
unpublished data). Given the riparian
improvements within Red Rock Lakes
NWR, and that the refuge represents the
vast majority of current Arctic grayling
habitat in the Centennial Valley, the
effects of degraded riparian habitat do
not appear to be acting on the core of
the Centennial Valley population at the
individual or population level.
Most of the riparian habitat
surrounding high-elevation lakes on
Federal land where the remaining
populations are found is intact and of
high quality (MFISH 2014a,
unpublished data; MFWP 2014e,
unpublished data; USFS 2014, p. 2),
because these habitats are in remote
locations or wilderness areas with little
anthropogenic disturbance. Given that
riparian degradation is being
systematically addressed in the Big Hole
River and Centennial Valley on the
National Refuge land where the majority
of Arctic grayling reside, we conclude
that riparian degradation is not a current
threat to the DPS. Riparian habitat is
expected to remain intact on Federal
land because of existing regulatory
mechanisms (see in Factor D discussion,
below). Riparian habitat in the Big Hole
River is expected to continue improving
because of the proven track record of
conservation evidenced by the current
upward trend in riparian habitat quality.
As more site-specific plans are signed
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under the Big Hole CCAA, more riparian
improvement is expected because
conservation measures will be similar
between currently implemented and
future site-specific plans. Given that
riparian habitat is intact or improving
for populations of Arctic grayling
occurring on Federal land and the Big
Hole population, and these populations
account for 19 of 20 populations in the
DPS, we conclude riparian habitat
degradation is not a current rangewide
threat and is not expected to become a
threat in the future.
Dewatering From Irrigation and
Consequent Increased Water
Temperatures
Demand for irrigation water in the
semi-arid upper Missouri River basin
historically dewatered many rivers
formerly or currently occupied by Arctic
grayling. The primary effects of this
dewatering were: (1) Increased water
temperatures, and (2) reduced habitat
capacity. In ectothermic species like
salmonid fishes, water temperature sets
basic constraints on species’
distribution and physiological
performance, such as activity and
growth (Coutant 1999, pp. 32–52).
Increased water temperatures can
reduce the growth and survival of Arctic
grayling (physiological stressor).
Reduced habitat capacity can
concentrate fishes and thereby increase
competition and predation (ecological
stressor). Below we discuss the potential
effects of increased water temperature
on the Upper Missouri River DPS of
Arctic grayling. For discussion of the
potential effects of reduced habitat
capacity, see Cumulative Effects from
Factors A through E, below.
Big Hole: In the Big Hole River
system, surface-water (flood) irrigation
has altered the natural hydrologic
function of the river (Shepard and
Oswald 1989, p. 29; Byorth 1993, p. 14;
1995, pp. 8–10; Magee et al. 2005, pp.
13–15). An inverse relationship between
flow volume and water temperature
(i.e., lower flows can lead to higher
water temperatures) is apparent in the
Big Hole River (Flynn et al. 2008, pp.
44, 46, but see Sladek 2013, p. 31).
Summer water temperatures exceeding
21 °C (70 °F) are considered to be
physiologically stressful for cold-water
fish species, such as Arctic grayling
(Hubert et al. 1985, pp. 7, 9). Summer
water temperatures consistently exceed
21 °C (70 °F) in the mainstem of Big
Hole River (Cayer and McCullough
2012, p. 7; (Cayer and McCullough
2013, p. 6) and have exceeded the upper
incipient lethal temperature (UILT; the
temperature that is survivable for
periods longer than 1 week by 50
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49403
percent of a ‘‘test population’’ in an
experimental setting) for Arctic grayling
(e.g., 25 °C or 77 °F) (Lohr et al. 1996).
As a result, thermal fish kills have been
documented in the Big Hole River (Lohr
et al. 1996, p. 934) in the past. The most
recent fish kill in the Big Hole River that
we are aware of occurred in 1994, and
included eight fish species, including
Arctic grayling (Lohr et al. 1996, p. 934).
Arctic grayling in the Big Hole River
use tributaries as a thermal refuge when
summer water temperatures in the
mainstem become stressful (Vatland et
al. 2009, p. 11). Summer water
temperatures within most tributaries are
cooler than those observed in some
reaches of the mainstem Big Hole River
(Vatland et al. 2009, entire; MFWP
2014b, unpublished data).
Since 2006, water conservation and
restoration projects associated with the
Big Hole Arctic grayling CCAA (for
more specific information, see
‘‘Conservation Efforts to Reduce Habitat
Destruction, Modification, or
Curtailment of Its Range,’’ below) have
been implemented to increase instream
flows and reduce water temperatures in
the Big Hole River and tributaries.
Varying flow targets for different
management segments of the Big Hole
River were outlined in the CCAA, based
on the wetted perimeter method, a
biologically based method for
determining instream flow requirements
to provide necessary resources for all
life stages of Arctic grayling. Over 300
irrigation diversions are operated under
flow agreements within finalized sitespecific plans (Table 4). The 10
remaining site-specific plans
representing the remainder of points of
diversion are expected to be signed in
August 2014. Although we are aware of
the future potential of more points of
diversion being managed under signed
site plans to contribute to Arctic
grayling conservation, we do not
consider these anticipated future efforts
to contribute to Arctic grayling
conservation currently, and have not
considered them as part of this status
review or our listing determination for
this DPS. Multiple other projects
designed to decrease dewatering and
thermal stress have been implemented
since 2006 (Table 4). The collective
result of these efforts are increasing
streamflows, increased access to coldwater refugia via fish ladders, and
marked temperature reductions,
particularly in some tributaries (Table
4).
Specific flow targets were developed
for the different Management Segments
in the CCAA Management Area (see
MFWP et al. 2006, pp. 7, 9, 13, for more
information on CCAA Management
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Segments). The goal for increasing
instream flow was to achieve flow
targets 75 percent of days in each
Management Segment during years of
average or greater snowpack. This goal
was based on a comparison between
minimum flow targets and historical
streamflows recorded in Management
Segments C and D. Achieving flow
targets 75 percent of days in each
Management Segment was intended to
be a general goal because many other
factors influence instream flows in the
Big Hole River that are outside the
control of landowners (e.g., snowpack,
precipitation). Before implementation of
the CCAA (2000–2005), average flow
targets were met among all Management
Segments 50 percent of the time, and
since implementation of the CCAA
(2006–2012), they have been met 78
percent of the time (SHCP 2013, p. 12).
Thus, the targets are being met.
Consistently since 2006, one
management area, known as
Management Segment C, has exhibited
the lowest instream flows among all
Management Segments. In part,
instream flows in Management Segment
C are influenced by several large
diversions immediately upstream of the
flow measuring device at the
downstream boundary of Management
Segment C (Robert 2014, pers. comm.).
Some of this diverted water is returned
to the Big Hole River downstream of the
flow measuring device (Robert 2014,
pers. comm.). As such, instream flows
in Management Segment C represent the
‘‘worst case’’ scenario among all
Management Segments. The Montana
Department of Natural Resources and
Conservation conducted an analysis of
this ‘‘worst case’’ scenario, to explore
how instream flows in Management
Segment C have changed since the
inception of the Big Hole CCAA. Given
that natural factors such as summer
precipitation and annual snowpack
influence instream flows in the Big Hole
River, the analysis of instream flows in
Management Segment C included
comparisons among several years of
similar (but below average) snow pack
and similar summer precipitation, both
before and after CCAA implementation
(Table 5).
TABLE 5—COMPARISON OF NUMBER OF DAYS VARYING FLOW TARGETS WERE ACHIEVED AMONG SIMILAR YEARS OF
BELOW AVERAGE SNOWPACK IN THE BIG HOLE RIVER CCAA MANAGEMENT SEGMENT C, PRE- AND POST CCAA.
ALL INFORMATION IN THIS TABLE CITED FROM ROBERTS 2014, UNPUBLISHED DATA
Pre-CCAA
1988
2003
Post-CCAA
2012
2013
Peak snowpack (percent of average) ..............................................................................................................
May–Aug. precipitation (in.) .............................................................................................................................
July–Aug. temps (degrees F; departure from normal) ....................................................................................
Signed SSPs ....................................................................................................................................................
Landowner contributions (cfs) .........................................................................................................................
Days <160 b cfs ................................................................................................................................................
Days <60 b cfs ..................................................................................................................................................
Days <20 cfs ....................................................................................................................................................
Days <10 cfs ....................................................................................................................................................
Mean discharge (cfs; July–Sept.) ....................................................................................................................
Mean discharge (cfs; Aug.) .............................................................................................................................
a73
4.14
¥1.3
0
0
50
123
79
65
8.4
1.1
108
3.85
8.0
0
0
8
123
68
7
19.7
14.2
81
4.74
1.4
12
252
11
87
0
0
45
33.7
a75
5.14
1.9
15
260
40
69
28
1
39
21
Total Days 2010
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(Roberts 2014, unpublished data).
Landowner contributions to instream
flow from reducing irrigation
withdrawals appears to be the primary
factor increasing instream flows in the
Big Hole River in late summer (Table 5),
a critical thermal period for Arctic
grayling.
Despite Management Segment C
exhibiting the lowest rate of instream
flow achievement relative to the other
Management Segments, we note that the
proportion of Tier I habitat
encompassed by Management Segment
C is 12 percent; the remainder of Tier
I habitat (88 percent) is located in
Management Segments D and E (MFWP
2014c, unpublished data). Since the
initiation of the Big Hole Arctic grayling
CCAA in 2006, average achievement
rate of instream flow goals in
Management Segments D and E during
the spring is 96 percent and 99 percent,
respectively. Average achievement rate
of instream flow goals in Management
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Segments D and E during the summer/
fall is 84 percent and 76 percent,
respectively. Thus, flow targets are
being met. We conclude the Big Hole
CCAA has ameliorated dewatering as a
stressor in the Big Hole River.
Other populations: Increased water
temperatures also are present in the
Madison River and Centennial Valley.
Mean and maximum summer water
temperatures can exceed 21 °C (70 °F)
in the Madison River below Ennis
Reservoir (U.S. Geological Survey
(USGS) 2010), and have exceeded 22 °C
(72 °F) in the reservoir, and 24 °C (75
°F) in the reservoir inlet (Clancey and
Lohrenz 2005, p. 34). However, Arctic
grayling in these systems appear to be
able to cope with these temperatures by
using cooler tributaries and spring
sources as thermal refugia (Jaeger 2014b,
pers. comm.). For example, the presence
of Arctic grayling in the lower 100 m
(328 ft) of East Shambow Creek in 1994
was attributed to fish seeking refuge
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from high water temperatures in the
lake (Mogen 1996, p. 44). The
Centennial Valley, in particular, appears
to have many cool spring-fed tributaries
that are accessible to Arctic grayling and
are used intermittently (Mogen 1996, p.
44). Mean summer water temperatures
in Red Rock Creek can occasionally
exceed 20 °C (68 °F) during drought
conditions (Mogen 1996, pp. 19, 45);
however, on average, these are much
cooler than summer water temperatures
observed in the Big Hole River.
Increased water temperatures do not
appear to be prevalent in most other
adfluvial populations, likely due to the
high elevation of these habitats and the
intact nature of riparian areas bordering
inlet tributaries. Given the presence of
cooler tributaries and spring sources
used by Arctic grayling in the
Centennial Valley and Ennis Reservoir/
Madison River, it does not appear
thermal stress is a current threat to these
populations. Although water
temperatures will likely increase with
climate change in the future, the springfed sources of cool water will likely
remain intact and within a temperature
range suitable for Arctic grayling
occupancy. Thus, thermal stress is not
expected to be a future threat.
Entrainment
Entrainment can permanently remove
individual fish from a natural
population and strand them in a habitat
that lacks the required characteristics
for reproduction and survival. Irrigation
ditches may dry completely when
irrigation headgates are closed, resulting
in mortality of entrained Arctic grayling.
Big Hole: Entrainment of individual
Arctic grayling in irrigation ditches
historically occurred and currently
occurs in the Big Hole River (Skarr
1989, p. 19; Streu 1990, pp. 24–25;
MFWP et al. 2006, p. 49; Lamothe 2008,
p. 22; MFWP 2013b, unpublished data).
Over 1,000 unscreened diversion
structures occur in the upper Big Hole
River watershed, and more than 300 of
these are located in or near occupied
Arctic grayling habitat (MFWP et al.
2006, pp. 48–49).
However, recent entrainment surveys
in irrigation ditches along the mainstem
Big Hole River and tributaries indicate
low levels of Arctic grayling
entrainment. Since 2006, 138 ditch
miles have been sampled using
electrofishing to estimate entrainment,
resulting in the capture of 73 Arctic
grayling, most of which were young-ofyear (MFWP 2013b, unpublished data).
This number is very low relative to the
size of the population. All documented
entrainment has occurred in 4 irrigation
ditches, one of which recently had a fish
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screen installed (MFWP 2013b,
unpublished data). No entrainment of
Arctic grayling has been documented in
any irrigation ditch since 2010
(including the 4 previously mentioned
where entrainment of Arctic grayling
had occurred), despite intensive
sampling by Montana Fish, Wildlife,
and Parks (Cayer 2014b, pers. comm.).
We do note that sampling typically does
not occur during the larval stage for
Arctic grayling. Larval losses into
irrigation ditches could be substantial
and go undetected under the current
sampling protocol. However,
observations of young of year Arctic
grayling in the Big Hole River indicate
that many, but not likely all, newly
emerged fry stay relatively close to the
area where they were born (Skaar 1989,
p. 51; Streu 1990, p. 28; McMichael
1990, p. 38), thus reducing the risk of
entrainment because of minimal
instream movements during their first
summer.
Irrigation ditches are prioritized and
systematically monitored based on the
ditch location relative to known Arctic
grayling distribution, additive
maximum flow rate, and distance from
the mainstem Big Hole River (MFWP et
al. 2006, p. 116). In addition,
electrofishing efficiency in simple
habitats (such as irrigation ditches) is
high (Kruse et al. 1998, pp. 942–943);
thus, we have high confidence that
these surveys have been accurate and
that entrainment in the Big Hole River
system is currently low and likely not
a threat at the population level.
Other populations: Entrainment was
likely a historical threat for Arctic
grayling at some locations within the
Centennial Valley (Unthank 1989, p. 10;
Gillin 2001, pp. 2–4, 3–18, 3–25),
particularly outside of the Red Rock
Lakes NWR (Boltz 2010, pers. comm.).
Currently, one irrigation ditch is present
near the core of the Centennial Valley
population within the Red Rock Lakes
NWR. This ditch conveys water from
Red Rock Creek to a waterfowl slough
for a portion of the year; however, it is
not operated by the Refuge when Arctic
grayling fry are expected to be in Red
Rock Creek (Bill West 2014a, pers.
comm.). Other irrigation ditches are
present upstream and downstream of
the NWR boundary; however, Arctic
grayling densities in these areas are low,
and any mortality associated with
entrainment in these areas is expected
to be negligible at the population level.
Entrainment of Arctic grayling does
not appear to be a threat in the Big Hole
River and Centennial Valley
populations. Habitats occupied by all 16
adfluvial Arctic grayling populations in
the upper Missouri River basin are not
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subjected to irrigation withdrawals; thus
entrainment is not a threat to these
populations. We expect irrigation
withdrawal volume to remain similar to
current levels, particularly in the Big
Hole River, in the future as more flow
agreements are signed under the CCAA.
Thus, we conclude entrainment will
likely not be a future threat.
Sedimentation
Sedimentation has been proposed as a
mechanism behind the decline of Arctic
grayling and its habitat in the
Centennial Valley (Unthank 1989, p. 10;
Mogen 1996, p. 76), which includes
Upper and Lower Red Rock Lakes.
Historically, livestock grazing upstream
likely led to accelerated sediment
transport in tributary streams, and
deposition of silt in both stream and
lakes, thus modifying and reducing fish
habitat by filling in pools, covering
spawning gravels, and reducing water
depth in Odell and Red Rock Creeks,
where Arctic grayling spawn (MFWP
1981, p. 105; Mogen 1996, pp. 73–76).
Sedimentation in the Upper and Lower
Red Rock Lakes is believed to affect
Arctic grayling in winter by reducing
habitat volume (e.g., lakes freezing to
the bottom) and promoting hypoxia (low
oxygen), which generally concentrates
fish in specific locations, thus
increasing the probability of
competition and predation. In summer,
reduced habitat volume could
contribute to increased warming.
Dissolved oxygen levels in Upper Red
Rock Lake during winter can drop
below levels typically considered lethal
for Arctic grayling (Gangloff 1996, pp.
41–42, 72). As a result, winter kill of
invertebrates and fishes (e.g., suckers
(Catostomus spp.)) has been recorded in
Upper Red Rock Lake (Gangloff 1996,
pp. 39–40); however, no Arctic grayling
kills have been documented. Gangloff
(1996, pp. 71, 79) hypothesized that
Arctic grayling in Upper Red Rock Lake
exhibit behavioral mechanisms or
physiological adaptations that permit
them to survive otherwise lethally low
oxygen levels. Arctic grayling under
winter ice seek areas of higher oxygen
concentration (oxygen refugia) within
the lake or near inlet streams of Upper
Red Rock Lake (Gangloff 1996, pp. 78–
79).
It has been reported that depths in the
Red Rock Lakes have decreased
significantly, with a decline in
maximum depth from 7.6 to 5.0 m (25
to 16.4 ft) to less than 2 m (6.5 ft) noted
in Upper Red Rock Lake over the past
century (Mogen 1996, p. 76). This
conclusion is prevalent among historical
accounts of the Centennial Valley.
However, a more recent analysis of
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sedimentation entering Upper Red Rock
Lake indicated modest rates of sediment
accumulation in Upper Red Rock Lake
over the last century and that the rate of
infilling in Upper Red Rock Lake has
been relatively constant, based on lead
and cesium analysis in lake bottom
cores (Allison 1996, unpublished data).
Thus, it appears historical accounts of
rapid infilling of Upper Red Rock Lake
were invalid and that sedimentation in
Upper Red Rock Lake is not a stressor.
Sedimentation in tributary streams
due to unregulated land use may have
contributed to historical Arctic grayling
declines in the Centennial Valley
(Vincent 1962, p. 114). Now, land use is
regulated, particularly on Federal land,
which comprises the majority of
ownership in the Centennial Valley.
However, some of the tributary streams
in the Centennial Valley are still
affected by sediment, even some spring
source streams. The effect of these levels
of sediment on Arctic grayling in the
Centennial Valley is unclear. However,
spawning conditions in Red Rock Creek
are currently supporting 40-year highs
in hybrid cutthroat and Arctic grayling
abundance (MFWP 2013c, unpublished
data).
The effects of erosion and
sedimentation on spawning gravels in
Red Rock Creek and reduction of habitat
volume in Upper and Lower Red Rock
Lakes do not appear to be current
threats because improved grazing
practices appear to be reducing erosion
rates upstream of Red Rock Lakes NWR
(USFWS 2009, pp. 75–76; Korb 2010,
pers. comm.; West 2014; pers. comm.).
Natural infilling of Upper Red Rock
Lake is occurring (Allison 1996,
unpublished data), but is not occurring
at a rate or scale that constitutes a threat
to Arctic grayling.
Climate Change
Our analyses under the Endangered
Species Act include consideration of
ongoing and projected changes in
climate. The terms ‘‘climate’’ and
‘‘climate change’’ are defined by the
Intergovernmental Panel on Climate
Change (IPCC). ‘‘Climate’’ refers to the
mean and variability of different types
of weather conditions over time, with 30
years being a typical period for such
measurements, although shorter or
longer periods also may be used (IPCC
2007, p. 78). The term ‘‘climate change’’
thus refers to a change in the mean or
variability of one or more measures of
climate (e.g., temperature or
precipitation) that persists for an
extended period, typically decades or
longer, whether the change is due to
natural variability, human activity, or
both (IPCC 2007, p. 78). Various types
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of changes in climate can have direct or
indirect effects on species. These effects
may be positive, neutral, or negative and
they may change over time, depending
on the species and other relevant
considerations, such as the effects of
interactions of climate with other
variables (e.g., habitat fragmentation)
(IPCC 2007, pp. 8–14, 18–19). In our
analyses, we use our expert judgment to
weigh relevant information, including
uncertainty, in our consideration of
various aspects of climate change.
Water temperature and hydrology
(stream flow) are sensitive to climate
change, and influence many of the basic
physical and biological processes in
aquatic systems. For ectothermic
organisms like fish, temperature sets
basic constraints on species’
distribution and physiological
performance, such as activity and
growth (Coutant 1999, pp. 32–52).
Stream hydrology not only affects the
structure of aquatic systems across
space and time, but influences the life
history and phenology (timing of lifecycle events) of aquatic organisms such
as fishes. For example, the timing of
snowmelt runoff can be an
environmental cue that triggers
spawning migrations in salmonid fishes
(Brenkman et al. 2001, pp. 981, 984),
and the timing of floods relative to
spawning and emergence can strongly
affect population establishment and
persistence (Fausch et al. 2001, pp.
1438, 1450). Significant trends in water
temperature and stream flow have been
observed in the western United States
(Kaushal et al. 2010, entire; Isaak et al.
2012, entire; Null et al. 2013, entire, and
climatic forcing (the energy difference
between incoming solar radiation and
outgoing radiation from Earth) caused
by increased air temperatures and
changes in precipitation are partially
responsible.
Observations on flow timing in the
Big Hole River, upper Madison River,
and Red Rock Creek in the Centennial
Valley indicate a tendency toward
earlier snowmelt runoff (Wenger et al.
2011, entire; Towler et al. 2013, entire;
De Haan et al. 2014, p. 41). These
hydrologic alterations may be
biologically significant for Arctic
grayling in the Missouri River basin
because they typically spawn prior to
the peak of snowmelt runoff (Shepard
and Oswald 1989, p. 7; Mogen 1996, pp.
22–23; Rens and Magee 2007, pp. 6–7).
A trend toward earlier snowmelt runoff
has resulted in earlier spawning in
European grayling in Switzerland
(Wedekind and Kung 2010, pp. 1419–
1420). The effects of altered timing of
spawn on Arctic grayling demographics
are unknown. However, it has been
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hypothesized that the timing of fry
emergence in salmonids is synchronized
with when food resources are available
(Crozier et al. 2008). Given that many
ecological processes in aquatic
environments are water temperature
dependent (Durance and Ormerod 2007,
entire), it is likely that any alterations in
timing of salmonid fry emergence would
be synchronous with alterations in the
timing of emergence and availability of
prey species.
Recent climate analyses in the Big
Hole River Valley and Centennial Valley
indicate rising air temperatures (1.8–3.2
°F (1.0–1.8 °C)/decade) from the 1980s
to mid-2000s (De Haan et al. 2014, p.
29). During this time, number of
breeding Arctic grayling in the Big Hole
River declined while the number of
breeding Arctic grayling in the
Centennial Valley increased (DeHaan et
al. 2014, p. 17), despite a coherent
climate signal between both drainages.
This may be partially attributable to
cool-water springs helping ameliorate
increasing air temperatures in the
Centennial Valley. Since the late 2000s,
the number of breeding Arctic grayling
has increased in the Big Hole River
(Leary 2014, unpublished data), and the
number of Arctic grayling in the Red
Rock Creek spawning run has increased
in the Centennial Valley (Patterson
2014, unpublished data). Thus, we have
no information to conclude that
increasing air temperatures have had a
significant effect on number of breeding
Arctic grayling in these systems in
recent years.
The effect of warming water from
increased air temperatures would be
similar to that described for increased
temperatures associated with stream
dewatering (see discussion under Factor
A), namely there has been an increased
frequency of high water temperatures
that have the potential to affect survival
or optimal growth for Arctic grayling,
which is considered a cold-water
(stenothermic) species (Selong et al.
2001, p. 1032). However, the transfer of
heat from air to water (i.e., convection)
is a relatively small proportion of the
energy exchange that occurs (Johnson
2003, p. 497). The more important factor
influencing water temperature is likely
to be solar radiation input (Johnson
2003, p. 497; Cassie 2006, p. 1393).
Thus, the changes in ambient air
temperature predicted to occur as the
climate changes are not likely to have as
large an effect on water temperatures as
solar radiation. Changes in channel
morphology (reducing width-to-depth
ratios) and riparian vegetation (shading)
resulting from the conservation actions
being implemented for Arctic grayling
are expected to reduce water
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temperatures by blocking some solar
radiation and reducing surface area that
solar radiation can interact with. In the
Big Hole River, where riparian areas are
improving and braided channels are
being restored, substantial reductions in
water temperature have been observed
(see Table 4, Rock Creek restoration for
example). We expect the restoration of
riparian areas and concomitant channel
morphology changes that have occurred
to help mitigate the effects of climate
change. In the Centennial Valley, intact
riparian areas are expected to mitigate
the effects of climate change, through
similar processes as in the Big Hole
River.
Warming patterns in the western
United States are not limited to streams.
In California and Nevada, lake water
surface temperatures have increased by
an average of 0.11 °C (0.2 °F) per year
since 1992, and at a rate twice that of
the average minimum air surface
temperature (Schneider et al. 2009, p.
L22402). This suggests lake habitats are
not immune to the predicted effects of
climate change. Shallow lakes with a
large surface area, such as Upper Red
Rock Lake and Ennis Reservoir, would
be expected to warm faster than deeper
lakes. However, all 16 strictly adfluvial
Arctic grayling populations in the upper
Missouri River drainage occur in lake
habitats that are expected to have
thermal regimes well below upper
thermal tolerances for Arctic grayling
because of high elevation, bathymetry
(underwater topography), and cool
inputs from shaded inlet streams.
The land area of the upper Missouri
River basin is predicted to warm
through the end of the century (Ray et
al. 2010, p. 23), although currently
occupied Arctic grayling habitat tends
to be in colder areas of moderate-to-high
elevation. Most of the Arctic grayling
populations are at approximately 1,775
to 2,125 m (5,860 to 9,000 ft) elevation
(Peterson and Ardren 2009, p. 1761;
MFISH 2014a, unpublished data).
Alterations to instream flow and timing
of runoff are already documented.
However, Arctic grayling are likely to
persist in the upper Missouri River
drainage because of what appears to be
an inherent ability possessed by Arctic
grayling to adjust spawn timing with
changing water temperature regimes
(Wedekind and Kung 2010, pp. 1419–
1420). In addition, it has been
demonstrated in the Big Hole River and
Centennial Valley that Arctic grayling
are capable of increasing in abundance
and distribution, despite a warming
climate (Dehaan et al. 2014, p. 17; Leary
2014, unpublished data). It appears
Arctic grayling within the upper
Missouri River basin are responding
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favorably to increasing quality of habitat
based on increasing abundance and
distribution in systems with large-scale,
ongoing habitat improvements (Big Hole
River and Centennial Valley). Riparian
restoration, particularly in the Big Hole
River and Centennial Valley, is expected
to minimize the effects of increasing
water temperatures due to climate
change. Sixteen other adfluvial
populations are currently in habitats
that will likely not be affected
significantly by climate change due to
their high elevation, intact riparian
areas, and cool inputs of tributary water.
Thus, we do not consider climate
change to be a current threat to Arctic
grayling in the upper Missouri River.
Further, observed water temperature
reductions from riparian restoration
projects indicate that intact riparian
areas can mitigate for many of the
anticipated effects of climate change in
the future. Therefore, we conclude
climate change is not a future threat to
the Upper Missouri River DPS of Arctic
grayling.
Conservation Efforts To Reduce Habitat
Destruction, Modification, or
Curtailment of Its Range
Big Hole River CCAA
In 2006, a CCAA was developed for
Arctic grayling in the Big Hole River.
The conservation goal of this CCAA is
to secure and enhance the fluvial
population of Arctic grayling in the
upper Big Hole River drainage. The
CCAA Management Area encompasses
about 382,000 acres and is divided into
five management segments to make the
conservation guidelines more spatially
meaningful to property owners enrolled
in the CCAA and to allow the involved
agencies to track the progress of the
conservation measures both temporally
and spatially.
Site-specific plans are developed with
each enrolled landowner; these plans
identify conservation actions needed (or
already completed) to meet the
conservation goals of the CCAA. The
conservation guidelines of the CCAA are
met by implementing conservation
measures that:
(1) Remove barriers to Arctic grayling
migration;
(2) Improve streamflows;
(3) Identify and reduce or eliminate
entrainment threats for Arctic grayling;
and
(4) Improve and protect the function
of riparian habitats.
Currently, 31 landowners have
enrolled 158,000 acres (∼52 percent total
enrollable land) in the Big Hole CCAA
Management Area. Of the 31
landowners, 21 have signed (finalized)
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site-specific plans. The remaining 10
landowners have site-specific plans in
various stages of completion. All of
these unfinished site plans are
scheduled to be finalized by August
2014, and will include measures
identified previously in the CCAA that
have a successful track record; however,
we have not considered future
anticipated conservation actions
outlined in unsigned site plans as part
of this status review or our listing
determination for this DPS.
Restoration and conservation efforts
outlined in site-specific plans are
guided by the Big Hole SHCP, a sciencebased framework for making
management decisions and prioritizing
where and how to deliver conservation
efficiently to achieve specific biological
outcomes for Arctic grayling. The SHCP
delineates four spatial ‘‘tiers’’ that help
prioritize where conservation will most
benefit Arctic grayling:
(1) Tier I is 84 miles of core spawning,
rearing and adult habitat that is
currently occupied by Arctic grayling;
(2) Tier II is 91 miles of periphery
habitat intermittently used by Arctic
grayling;
(3) Tier III is 161 miles of suitable, but
currently unoccupied, historical habitat;
and
(4) Tier IV is 33 miles of potentially
suitable habitat with unknown
historical occupancy.
For reference, lands currently
enrolled in the CCAA include 86
percent of tier I, 73 percent of tier II, 42
percent of tier III, and 24 percent of tier
IV habitats. Given that the conservation
measures outlined in the CCAA directly
address known threats to Arctic grayling
and their habitat in the Big Hole River,
and that all conservation actions are
strategically prioritized through the
SHCP, the Service is encouraged by the
positive habitat and Arctic grayling
response to the conservation actions in
the Big Hole River.
Conservation Efforts by Landowners Not
Enrolled in the CCAA
Since 2006, twelve landowners in the
Big Hole Valley who are not enrolled in
the Big Hole CCAA have implemented
voluntary conservation measures to
benefit Arctic grayling. These
conservation measures are similar to the
conservation measures outlined in the
SSP’s of landowners enrolled in the
CCAA, including irrigation withdrawal
reductions, installation of fish passage
ladders, riparian fencing, stream
restoration, and installation of
stockwater tanks (MFWP 2014f,
unpublished data). In addition, several
of these landowners have informal flow
agreements where the landowners have
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agreed to not utilize water returned to
the stream by upstream enrollees in the
Big Hole CCAA (MFWP 2014f,
unpublished data). Although the
majority of conservation projects in the
Big Hole are completed through the
CCAA, the Service is very encouraged
by the participation of non-enrolled
landowners to further grayling
conservation in the Big Hole River and
its tributaries.
Big Hole River Drought Management
Plan
The purpose of the Drought
Management Plan (DMP) is to mitigate
the effects of low stream flows and
lethal water temperatures for fisheries
(particularly fluvial Arctic grayling)
through a voluntary effort among
participants including agriculture,
municipalities, business, conservation
groups, anglers, and affected
government agencies (Big Hole
Watershed Committee 2014, p.1). The
DMP outlines flow triggers that, when
met, initiate specific voluntary actions
to conserve water. The flow triggers in
the DMP are the same as the flow targets
outlined in the Big Hole CCAA. The
DMP has been in effect since 1999.
One key difference between the DMP
and the CCAA is that the DMP is in
effect for the entire Big Hole River, not
just the upper Big Hole River like the
CCAA. Arctic grayling occur outside of
the CCAA Management Area; thus, any
conservation efforts occurring in these
areas still likely benefit Arctic grayling,
although Arctic grayling densities
outside the CCAA Management Area are
low and represent a small fraction of the
total population inside the CCAA
Management Area (MFWP 2013d,
unpublished data). Another key
difference is that the DMP is structured
to disseminate flow and water
temperature information to all users of
the Big Hole River, not just private
landowners in the CCAA. This
structuring allows for near real-time
information sharing that helps inform
users when voluntary conservation
actions are needed. Such actions
include reductions in irrigation
withdrawal (for downstream users not
in the CCAA); reductions in municipal,
industrial, and personal water use; and
reductions in recreation (e.g., angling).
The extent and magnitude of
beneficial effects to Arctic grayling from
the voluntary conservation measures
recommended in the DMP are unclear.
However, the DMP appears to have
broad-based support. Most participants
reduce irrigation withdrawals in
response to observed low flows on
nearby USGS gauges, before phone calls
are made to request irrigation reductions
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(Downing 2014, pers. comm.). Increases
in instream flow attributable to efforts
under the DMP have been observed as
‘‘bumps’’ in the hydrograph in the
middle and lower reaches of the Big
Hole River (Downing 2014, pers.
comm.). Although difficult to quantify,
these ‘‘bumps’’ typically result in
instream flows rising above low flow
triggers (Downing 2014, pers. comm.). In
addition, the inherent value of
information sharing among diverse
stakeholder groups about the potential
effects of dewatering and thermal stress
on the Big Hole fishery is likely
significant. An increased understanding
of conservation efforts needed to benefit
Arctic grayling, and aquatic habitat in
general, has been demonstrated to be a
necessary precursor for more formalized
conservation actions, such as the
creation and implementation of the Big
Hole CCAA.
Native Arctic Grayling Genetic Reserves
and Translocation
Given concern over the status of
native Arctic grayling, the Montana
Arctic Grayling Recovery Program
(AGRP) was formed in 1987, to address
conservation concerns for primarily the
fluvial ecotype inhabiting the Big Hole
River, and to a lesser extent the native
adfluvial population in the Centennial
Valley (Memorandum of Understanding
(MOU) 2007, p. 2). The Arctic Grayling
Workgroup (AGW) was established as
an ad hoc technical workgroup of the
AGRP. In 1995, the AGW finalized a
restoration plan that outlined an agenda
of restoration tasks and research,
including management actions to secure
the Big Hole River population, brood
stock development, and a program to reestablish four additional fluvial
populations (Montana AGW 1995, pp.
7–17).
Consequently, Montana Fish, Wildlife
and Parks established genetic reserves of
Big Hole River and Centennial Valley
Arctic grayling (Leary 1991, entire).
Currently, brood (genetic) reserves of
Big Hole River Arctic grayling are held
in two closed-basin lakes in southcentral Montana (Rens and Magee 2007,
p. 22). These fish are manually spawned
to provide gametes for translocation
efforts in Montana (e.g., Ruby River
population) (Rens and Magee 2007, p.
22). A brood reserve of Centennial
Valley Arctic grayling has recently been
established in Elk Lake; however, no
natural reproduction has been
documented since the brood reserve was
established. Instream flows in the sole
spawning tributary (Narrows Creek) to
Elk Lake have been low in recent years,
likely as a result of low snowpack in
some years and seismic activity that
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altered the hydrology of Narrows Creek
(Jaeger 2014c, pers. comm.). Future
conservation actions on Narrows Creek
include securing a more consistent
water supply during the Arctic grayling
spawning season through a water rights
exchange; however, at this time, these
conservation actions and the future
viability of the Elk Lake population are
too uncertain to warrant consideration
in this finding.
A reintroduction effort in the upper
Ruby River, where Arctic grayling were
previously extirpated, using Big Hole
River genetic reserves recently
concluded. Arctic grayling eggs from the
Big Hole River reserves were hatched
on-site in incubators, and fry were
allowed to drift into the reintroduction
area. Supplementation of the Ruby River
population concluded in 2008. For the
last 5 years since then, natural
reproduction has been documented in
the upper Ruby River (Cayer and
McCullough 2013, p. 21). Recent genetic
analyses of the Ruby River population
indicate high levels of genetic
heterozygosity and allelic richness,
albeit low estimate of effective number
of breeders (Leary 2014, unpublished
data). It has been hypothesized that the
population is likely still expanding.
Encouragingly, the number of breeding
adults has trended upward over the past
3 years (Leary 2014, unpublished data).
Most experts participating in the SSA
workshop indicated that the Ruby River
population was viable, given the
evidence of natural reproduction
occurring over the last 5 years at rates
sufficient to increase the number of
breeding adults over the past 3 years.
Thus, we conclude the Ruby River
population is viable.
Another recent conservation effort
using Big Hole River genetic reserves
involves an assisted recolonization
effort of Arctic grayling in Rock Creek,
a historically occupied tributary of the
upper Big Hole River. Since 2010,
incubators placed directly on location
have been used to reintroduce young-ofyear Arctic grayling into Rock Creek.
Recolonization efforts are scheduled to
be implemented in Rock Creek through
2015. Encouragingly, young-of-year and
older Arctic grayling have been
documented in 4 miles of Rock Creek
over the past several years. This
increase of Arctic grayling abundance
and distribution in Rock Creek is likely
due, at least in part, to the introduction
of thousands of fry via the onsite
incubators. Habitat improvement
projects on Rock Creek have occurred
simultaneously with fry reintroduction,
so it is difficult to distinguish the
relative effects of fry reintroduction and
habitat improvement on the resulting
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increase in distribution and abundance
of young Arctic grayling. Likely, both
factors have played a role in
reestablishing Arctic grayling in Rock
Creek. Regardless, both conservation
actions are having their intended effect:
Increasing Arctic grayling abundance
and distribution in historically occupied
habitat.
In 2013, an increase in the number of
breeding Arctic grayling was observed
in the Big Hole River (Leary 2014,
unpublished data). Given that fry were
being reintroduced into the Big Hole
River (and Rock Creek) beginning in
2010, there was initial uncertainty about
the relative contribution of RSIproduced fish to the observed increase
in breeding adults. Genetic analysis of a
sample of young Arctic grayling
obtained in 2013 indicated a low level
of relatedness (<10 percent of sample
were half- or full siblings) among
individuals within the sample (Leary
2014, unpublished data). These results
indicate that RSI-produced fish in 2010
contributed very little to the increase in
breeding adults in 2013, as we would
have expected a high degree of
relatedness within the 2013 sample due
to a small number of grayling spawned
to produce eggs for the RSI
reintroduction effort. Thus, these data
suggest that factors other than the
influence of RSIs were responsible for
increasing abundance of adult spawners
in the Big Hole River in 2013.
Similar reintroductions to the Rock
Creek effort are also underway in
several other tributaries and lakes
within the upper Big Hole drainage and
elsewhere, including the Wise River,
Trail Creek, Twin Lakes and the
Madison River. This suite of
reintroductions is scheduled to occur
for 5 years and conclude in 2018.
However, the effectiveness of these
reintroductions has not yet been
assessed.
In the Centennial Valley, RSIs have
been used fairly extensively to try to
establish spawning runs of adult Arctic
grayling in multiple tributaries to Upper
Red Rock Lake (Boltz and Kaeding 2002,
entire; Jaeger 2014d, pers. comm.). Thus
far, these attempts have failed, although
it is possible that fry produced by these
efforts spawned in Red Rock Creek,
instead of returning to the stream in
which they were hatched (Mogen 2014,
pers. comm). Recently, RSIs were used
in Red Rock Creek in 2010, as part of a
mitigation strategy to offset the removal
of thousands of Arctic grayling eggs
being taken to develop a brood reserve
in Elk Lake. Similar to the Rock Creek
example in the Big Hole, there was
initial uncertainty whether recent
increases in spawner abundance of
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Arctic grayling in Red Rock Creek were
a direct result of the introduction of
thousands of fry directly into Red Rock
Creek via RSIs. However, fry produced
in 2010 would not have contributed to
the increases in spawning adult Arctic
grayling observed in Red Rock Creek in
2010. Additionally, lengths of all adult
Arctic grayling handled during the 2012
spawning run showed minimal overlap
(<5 percent) with the length range of 2year-old Arctic grayling from Upper Red
Rock Lake, indicating fry produced in
2010 had little potential to contribute to
the observed increase in adult spawners
in Red Rock Creek in 2012 (Patterson
2014, unpublished data). The effect of
RSI-produced Arctic grayling fry on
abundance of spawners in Red Rock
Creek after 2012 is unknown. The
Service hopes that using RSIs as a
conservation tool will result in RSIproduced fish recruiting to the
Centennial Valley population.
Another Arctic grayling
reintroduction project is currently being
planned in Grayling Creek within
Yellowstone National Park.
Approximately 30 miles of historically
occupied habitat are proposed for the
reintroduction. Recently, a fish barrier
was installed at the downstream extent
of this habitat, and removal of all fish
currently above the barrier commenced.
Another round of fish removal is
scheduled for the summer of 2014, to
ensure complete removal of all existing
nonnative fishes. Arctic grayling are
scheduled to be reintroduced as early as
2015. Although the Service is
encouraged by the potential for the
reintroduction in 30 miles of historical
habitat in Grayling Creek, it is unclear
at this time if funding will be available
to complete the project. Thus, although
we are aware of the future potential of
this project to contribute to Arctic
grayling conservation, we do not
consider this project to contribute to
Arctic grayling conservation currently,
and have not considered it as part of
this status review or our listing
determination for this DPS.
Summary of Factor A
Based on the best available
information, we find that the historical
range of the Missouri River DPS of
Arctic grayling has been reduced
particularly by large-scale habitat
fragmentation by dams. However,
despite fragmentation, sufficient habitat
remains intact and is currently
supporting multiple, viable, fluvial and
adfluvial Arctic grayling populations.
Historical threats to habitat quantity and
quality in the Big Hole River are
systematically being eliminated or
minimized by the CCAA and SHCP
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through conservation projects designed
to expressly address the four
conservation criteria outlined in the
CCAA. Large-scale habitat
improvements are occurring; quality of
riparian areas has improved in both the
Big Hole River and Centennial Valley
through riparian restoration projects,
and these projects are expected to
minimize effects of climate change
through blocking of some solar radiation
and channel morphology changes. In
addition, Arctic grayling populations
are responding favorably to habitat
improvements in both the Big Hole
River and Centennial Valley. In the
future, we do not expect habitat to
decline in the Big Hole River because of
the proven track record of CCAA
projects. In the Centennial Valley,
protections provided by the NWR have
sufficiently minimized past threats to
habitat. These protections are expected
to persist into the future and maintain
the integrity of the habitat. Most of the
other adfluvial populations of Arctic
grayling reside in high-quality habitats
on Federal land where mechanisms
exist to conserve that habitat. Thus, we
have no evidence that past threats under
Factor A are acting currently on the
Upper Missouri River DPS of Arctic
grayling at the population or DPS level
and no expectation that those impacts
will pose a threat to the DPS in the
future.
B. Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Arctic grayling of the upper Missouri
River are handled for recreational
angling and for scientific, population
monitoring, and restoration purposes.
Recreational Angling
Arctic grayling are highly susceptible
to capture by angling (ASRD 2005, pp.
19–20), and intense angling pressure
can reduce densities and influence the
demography of exploited populations
(Northcote 1995, pp. 171–172).
Historically, overfishing likely
contributed to the rangewide decline of
the DPS in the upper Missouri River
system (Vincent 1962, pp. 49–52, 55;
Kaya 1992, pp. 54–55). In 1994, concern
over the effects of angling on fluvial
Arctic grayling led the State of Montana
to implement catch-and-release
regulations for Arctic grayling captured
in streams and rivers within its native
range, and those regulations remain in
effect today (MFWP 2014d, p. 51).
Catch-and-release regulations also are in
effect for Ennis Reservoir on the
Madison River (MFWP 2014d, p. 59).
Angling is not permitted in either of the
Red Rock Lakes in the Centennial Valley
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to protect breeding waterfowl and
trumpeter swans (Cygnus buccinator)
(USFWS 2009, p. 147), and catch-andrelease regulations remain in effect for
any Arctic grayling captured in streams
(e.g., Odell Creek or Red Rock Creek) in
the Centennial Valley (MFWP 2014d, p.
67). Additionally, angling is closed in
Red Rock Creek during the Arctic
grayling spawning period (May 15 to
June 14; MFWP 2014d, p. 67). However,
we do note that Red Rock Creek is open
to angling for other species (e.g., hybrid
cutthroat trout) the remainder of the
year and that Arctic grayling are caught
during this time, particularly before the
May 15 closure.
In all other populations, anglers can
keep up to 5 Arctic grayling per day and
have up to 10 in possession, in
accordance with standard daily and
possession limits for that angling
management district (MFWP 2014d,
p. 51). The population trends of Arctic
grayling in many of the lakes (see Table
3, above) suggest that present angling
exploitation rates are not a threat to
those populations, even though harvest
is allowed on most of these populations.
Limited data preclude population
estimates and trend inferences for some
adfluvial populations (see Table 3).
Repeated catch-and-release angling
may harm individual fish, causing
physiological stress and injury (i.e.,
hooking wounds). Catch-and-release
angling also can result in mortality at a
rate dependent on hooking location,
hooking duration, fish size, water
quality, and water temperature
(Faragher et al. 2004, entire;
Bartholomew and Bohnsack 2005,
p. 140; Boyd et al. 2010). Repeated
hooking (up to five times) of Arctic
grayling in Alaska did not result in
significant additional mortality (rates 0
to 1.4 percent; Clark 1991, pp. 1, 25–26).
In Michigan, hooking mortality of Arctic
grayling in lakes averaged 1.7 percent
per capture event, based on 355
individuals captured with artificial flies
and lures (Nuhfer 1992, pp. 11, 29).
Higher mortality rates (5 percent) have
been reported for Arctic grayling
populations in the Great Slave Lake
area, Canada (Falk and Gillman 1975,
cited in Casselman 2005, p. 23).
Comparatively high catch rates for
Arctic grayling have been observed in
the Big Hole River, Montana (Byorth
1993, pp. 26–27, 36), and average
hooking wound rates ranged from 15 to
30 percent among study sections
(Byorth 1993, p. 28). However, overall
hooking mortality from single capture
events was low (1.4 percent), which led
Byorth to conclude that the Big Hole
River population was not limited by
angling (Byorth 1994b, entire).
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Compared to the average catch-andrelease mortality rates of 4.2 to 4.5
percent in salmonids as reported by
Schill and Scarpella (1997, p. 873), and
the mean and median catch-and-release
mortality rates of 18 percent and 11
percent from a meta-analysis of 274
studies (Bartholomew and Bohnsack
2005, pp. 136–137), the catch-andrelease mortality rates for Arctic
grayling are comparatively low (Clark
1991, pp. 1, 25–26; Nuhfer 1992, pp. 11,
29; Byorth 1994b, entire). We are
uncertain whether these lower observed
rates reflect an innate resistance to
effects of catch-and-release angling in
Arctic grayling or whether they reflect
differences among particular
populations or study designs used to
estimate mortality. Even if catch-andrelease angling mortality is low (e.g., 1.4
percent as reported in Byorth 1994b,
entire), the high catchability of Arctic
grayling (ASRD 2005, pp. 19–20) raises
some concern about the cumulative
mortality of repeated catch-and-release
captures. For example, based on the
Arctic grayling catch rates and angler
pressure reported by Byorth (1993, pp.
25–26) and the population estimate for
the Big Hole River reported in Byorth
(1994a, p. ii), a simple calculation
suggests that age 1 and older Arctic
grayling susceptible to recreational
angling may be captured and released
3 to 6 times per year.
In conclusion, angling harvest may
have significantly reduced the
abundance and distribution of the
Upper Missouri River DPS of Arctic
grayling during the past 50 to 100 years,
but current catch-and-release fishing
regulations (or angling closures) in most
waters occupied by extant populations
have likely ameliorated the past threat
of overharvest. Although we do note the
potential for cumulative mortality
caused by repeated catch-and-release of
individual Arctic grayling in the Big
Hole River, we have no evidence
indicating that repeated capture of
Arctic grayling under catch-and-release
regulations is currently limiting that
population or the DPS. Moreover,
fishing is restricted in the Big Hole
River, an important recreational fishing
destination in southwestern Montana,
when streamflow and temperature
conditions are likely to increase stress to
captured Arctic grayling. Anglers can
still capture and keep Arctic grayling in
most lake populations in accordance
with State fishing regulations, but we
have no evidence that current levels of
angling are affecting these populations.
Thus, the best available evidence does
not indicate that recreational angling
represents a current threat to the DPS.
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We have no information at this time to
indicate that future fishing regulations
are likely to change in a way that would
be detrimental to Arctic grayling. Thus,
we do not believe that recreational
angling will represent a threat in the
future.
Monitoring and Scientific Study
Montana Fish, Wildlife and Parks
consistently monitors the Arctic
grayling population in the Big Hole
River and its tributaries, and to a lesser
extent those populations in the Madison
River and Centennial Valley (Cayer and
McCullough 2013, entire). Electrofishing
(use of electrical current to temporarily
and non-lethally immobilize a fish for
capture) is a primary sampling method
to monitor Arctic grayling in these
populations (Rens and Magee 2007, pp.
13, 17, 20). A number of studies have
investigated the effects of electrofishing
on various life stages of Arctic grayling.
Dwyer and White (1997, p. 174) found
that electrofishing reduced the growth
of juvenile Arctic grayling and
concluded that long-term, sublethal
effects of electrofishing were possible.
Hughes (1998, pp. 1072, 1074–1075)
found evidence that electrofishing and
tagging affected the growth rate and
movement behavior of Arctic grayling in
the Chena River, Alaska. Roach (1999, p.
923) studied the effects of electrofishing
on fertilized Arctic grayling eggs and
found that while electrofishing could
result in egg mortality, the populationlevel effects of such mortality were not
likely to be significant. Lamothe and
Magee (2003, pp. 16, 18–19) noted
mortality of Arctic grayling in the Big
Hole River during a radio-telemetry
study, and concluded that handling
stress or predation were possible causes
of mortality. However, population
monitoring activities in the Big Hole
River are curtailed when environmental
conditions become unsuitable (Big Hole
Watershed Committee 1997, entire), and
recent monitoring reports (Cayer and
McCullough 2012, 2013, entire) provide
no evidence that electrofishing is
harming the Arctic grayling population
in the Big Hole River.
Traps, electrofishing, and radio
telemetry have been used to monitor
and study Arctic grayling in the
Centennial Valley (Gangloff 1996, pp.
13–14; Mogen 1996, pp. 10–13, 15;
Kaeding and Boltz 1999, p. 4; Rens and
Magee 2007, p. 17); however, there are
no data to indicate these monitoring
activities reduce the growth and
survival of individual Arctic grayling or
otherwise constitute a current or future
threat to the population.
The Arctic grayling population in the
Madison River–Ennis Reservoir is not
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monitored as intensively as the Big Hole
River population (Rens and Magee 2007,
pp. 20–21). When electrofishing surveys
targeting Arctic grayling in the Madison
River occur, they are conducted during
the spawning run for that population
(Clancey 1996, p. 6). Capture and
handling during spawning migrations or
during actual spawning could affect the
reproductive success of individual
Arctic grayling. However, under recent
monitoring frequencies, any populationlevel effect of these activities is likely
negligible, and we have no data to
indicate these monitoring activities
reduce the growth and survival of
individual Arctic grayling or otherwise
constitute a current or future threat to
the Madison River population.
Most of the adfluvial populations of
Arctic grayling are infrequently
monitored (MFISH 2014a, unpublished
data). Because monitoring of these
populations has been minimal, we do
not believe that monitoring or scientific
study constitutes a current or future
threat to these particular populations.
The intensity of monitoring and
scientific investigation varies among the
different populations in the DPS, but we
have no evidence suggesting that
monitoring or scientific study has
influenced the decline of Arctic grayling
in the Missouri River basin. We also
have no evidence indicating these
activities constitute a current threat to
the DPS that would result in
measurable, population-level effects. We
expect similar levels of population
monitoring and scientific study in the
future, and we conclude that these
activities will not represent a threat in
the future.
Reintroduction Efforts
Attempts to restore or re-establish
native populations of both fluvial and
adfluvial Arctic grayling may result in
the mortality of some embryos and
young fish. Currently, gametes (eggs and
sperm) used to re-establish the fluvial
ecotype come from captive brood
reserves of Big Hole River Arctic
grayling maintained in Axolotl and
Green Hollow II Lakes (Rens and Magee
2007, pp. 22–24). Removal of gametes
from the wild Big Hole River population
was necessary to establish this brood
reserve (Leary 1991, entire) and will
likely continue intermittently in the
future to ensure the genetic
representation of the brood reserve. The
previous removal of gametes for
conservation purposes could have
hypothetically reduced temporarily the
abundance of the wild population if the
population was unable to compensate
for this effective mortality by increased
survival of remaining individuals.
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However, the establishment of a brood
reserve provides a conservation benefit
from the standpoint that gametes from
the reserve can be harvested to use for
translocation efforts to benefit the
species. Ultimately, we conclude that
past gamete collection from the Big Hole
River population has not harmed the
wild population or that collection in the
future will harm the population.
Consequently, we conclude that gamete
collection from the Big Hole River
Arctic grayling population does not
constitute a current or future threat to
the population.
Efforts to re-establish native,
genetically representative populations
of adfluvial Arctic grayling in the
Centennial Valley and to maintain a
brood reserve for that population have
resulted in the direct collection of eggs
from Arctic grayling spawning runs in
Red Rock Creek. During 2000–2002, an
estimated 315,000 Arctic grayling eggs
were collected from females captured in
Red Rock Creek (Boltz and Kaeding
2002, pp. v, 8). The Service placed over
180,000 of these eggs in remote site
incubators in streams within the Red
Rock Lakes NWR that historically
supported Arctic grayling spawning
runs (Boltz and Kaeding 2002, pp. v,
10).
Montana Fish, Wildlife and Parks and
the Service are currently collaborating
on an effort to re-establish an Arctic
grayling spawning run in Elk Springs
Creek and a replicate of the Centennial
population in Elk Lake (West 2014a,
pers. comm., Jaeger 2014e, pers. comm.).
These actions required the collection of
gametes (approximately 370,000 eggs)
from Arctic grayling captured in Red
Rock Creek (Jaeger 2014f, pers. comm.).
Approximately 10 percent of these eggs
were returned to Red Rock Creek and
incubated in that stream (using a
method resulting in high survivorship of
embryos) (Kaeding and Boltz 2004,
entire) to mitigate for collection of
gametes from the wild spawning
population. We infer that past gamete
collection in Red Rock Creek has not
significantly influenced recruitment in
Red Rock Creek, as abundance of
returning spawners to Red Rock Creek
was robust in 2013 and 2014.
Overall, we conclude that collection
of gametes from the wild populations in
the Big Hole River and Centennial
Valley systems has not contributed to
population-level declines in those
populations, or that the previous
collections represent overexploitation.
Future plans to collect gametes from
Arctic grayling in the Big Hole River
should be evaluated in light of the status
of those populations at the anticipated
time of the collections. We encourage
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the agencies involved to coordinate
their efforts and develop a strategy for
broodstock development and
conservation efforts that minimizes any
potential impacts to wild native
populations. However, at present, we do
not have any data indicating collection
of gametes for conservation purposes
represents a current threat to the Big
Hole River and Centennial Valley
populations. We have no evidence to
indicate that gamete collection will
increase in the future, so we conclude
that this does not represent a future
threat.
Conservation Efforts To Reduce
Recreational Overutilization
The MFWP closes recreational angling
in specific reaches of the Big Hole River
when environmental conditions are
considered stressful. Specific
streamflow and temperature thresholds
initiate mandatory closure of the fishery
(Big Hole Watershed Committee 1997,
entire). Such closures have been
implemented as recently as 2013;
however, changes to closure types and
criteria in past years preclude any
meaningful comparisons between
different time periods (Horton 2014b,
pers. comm.).
Summary of Factor B
Based on the best information
available, we conclude that
overexploitation by angling may have
contributed to the historical decline of
the Upper Missouri River DPS of Arctic
grayling, but we have no evidence to
indicate that current or future levels of
recreational angling, population
monitoring, scientific study, or
conservation actions constitute
overexploitation; therefore, we do not
consider them a threat. We expect
similar or decreased levels of these
activities to continue in the future, and
we do not believe they are likely to
become a threat in the future.
Factor C. Disease or Predation
Disease
Arctic grayling are resistant to
whirling disease, which is responsible
for population-level declines of other
stream salmonids (Hedrick et al. 1999,
pp. 330, 333). However, Arctic grayling
are susceptible to bacterial kidney
disease (BKD), a bacterial disease
causing reduced immune response and
mortality in some fish species (Meyers
et al. 1993, p. 181). Some wild
populations in pristine habitats test
positive for BKD (Meyers et al. 1993, pp.
186–187), but clinical effects of the
disease are more likely to be evident in
captive populations (Meyers et al. 1993,
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native populations. Arctic grayling in
captive brood reserves (e.g., Axolotl
Lake, Green Hollow Lake) have all
recently tested negative for infectious
hematopoietic necrosis virus (IHNV),
infectious pancreatic necrosis virus
(IPNV), Myxobolus cerebralis (the
pathogen that causes whirling disease),
Renibacterium salmoninarum (the
pathogen that causes BKD), and
Aeromonas salmonicida (the pathogen
that causes furunculosis) (USFWS
2010b). Consequently, the best available
evidence at this time does not indicate
that disease threatens native Arctic
grayling of the upper Missouri River.
We have no basis to conclude that
disease will become a future threat, so
we conclude that disease does not
constitute a threat in the future.
Ecological interactions (predation and
competition) with the brook trout,
brown trout, and rainbow trout are
among the long-standing hypotheses to
explain the historical decline of Arctic
grayling in the upper Missouri River
system and the extirpation of some
populations from specific waters
(Nelson 1954, p. 327; Vincent 1962, pp.
81–96; Kaya 1992, pp. 55–56). Strength
of competition and predation can be
very difficult to measure in wild trout
populations (Fausch 1988, pp. 2238,
2243; 1998, pp. 220, 227). Predation on
Arctic grayling eggs and fry by brook
trout has been observed in both the Big
Hole River and the Centennial Valley
(Nelson 1954, entire; Streu 1990, p. 17;
Katzman 1998, pp. 35, 47, 114), but
such observations have not been
definitively linked to population
declines of Arctic grayling. To our
knowledge, no studies have investigated
or attempted to measure predation by
brown trout or rainbow trout on Arctic
grayling in Montana. Brook trout do not
appear to negatively affect habitat use or
growth of juvenile, hatchery-reared
Arctic grayling (Byorth and Magee 1998,
p. 921), but further studies are necessary
to determine whether competition or
predation occur at other life stages or
with brown or rainbow trout (Byorth
and Magee 1998, p. 929). Predation
represents direct mortality that can limit
populations, and young-of-year Arctic
grayling may be particularly susceptible
to predation by other fishes because
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Predation By and Competition With
Nonnative Trout
Brook trout (Salvelinus fontinalis),
brown trout (Salmo trutta), and rainbow
trout are widely distributed and
abundant in the western United States,
including the upper Missouri River
system (Schade and Bonar 2005, p.
1386; Table 6). One or more of these
nonnative trout species co-occur with
11 of the 20 Arctic grayling population
in the basin. The remaining nine Arctic
grayling populations occur with other
native species or no other fish species
(Table 6).
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entire; Peterson 1997, entire). To
preclude transmission of BKD between
Arctic grayling during brood reserve,
hatchery, and wild Arctic grayling
translocation efforts, MFWP tests kidney
tissue and ovarian fluid for the
causative agent for BKD as well as other
pathogens in brood populations (Rens
and Magee 2007, pp. 22–24).
Information on the prevalence of BKD
or other diseases in native Arctic
grayling populations in Montana is
generally lacking. One reason for this
lack of information is that some disease
assays are invasive or require the
sacrifice of individual fish (e.g., removal
of kidney tissue to test for BKD
pathogen), so they are not done often on
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they are smaller and weaker swimmers
than trout fry (Kaya 1990, pp. 52–53).
The evidence for predation and
competition by nonnative trout on
Arctic grayling in the upper Missouri
River basin is largely circumstantial,
and inferred from the reduced historical
abundance and distribution of Arctic
grayling following encroachment by
nonnative trout (Kaya 1990, pp. 52–54;
Kaya 1992, p. 56; Magee and Byorth
1995, p. 54). In addition, the historical
difficulty in establishing Arctic grayling
populations in waters already occupied
by nonnative trout, especially brown
trout (Kaya 2000, pp. 14–15) may
suggest competition and predation play
a role. However, the often-cited case
histories where nonnative trout were
implicated in the decline of Arctic
grayling also involved prior or
concurrent habitat modification or
degradation, thus confounding the two
factors (Kaya 1990, pp. 52–54; Kaya
1992, p. 56; Magee and Byorth 1995, p.
54) and making it difficult to pinpoint
the cause of the decline. Where past
habitat degradation has not been a factor
(e.g., many of the high-elevation
adfluvial populations), successful
coexistence between brook trout and
rainbow trout and Arctic grayling has
occurred over long durations, greater
than 100 years in some populations
(Jaeger 2014, unpublished data; MFISH
2014a, unpublished data). Despite past
habitat degradation in the Big Hole
River, Arctic grayling have coexisted
with brook, rainbow and brown trout for
at least 60 years (Liknes 1981, p. 34).
In the Big Hole River, brook trout,
rainbow trout, and brown trout are more
abundant than Arctic grayling (Rens and
Magee 2007, p. 42). In general, brook
trout is the most abundant nonnative
trout species in the Big Hole River
upstream from Wisdom, Montana (Rens
and Magee 2007, pp. 7, 42; Lamothe et
al. 2007, pp. 35–38), whereas rainbow
trout and brown trout are comparatively
more abundant in the downstream
reaches (Kaya 1992, p. 56; Oswald 2005,
pp. 22–29; Lamothe et al. 2007, pp. 35–
38; Rens and Magee 2007, p. 10).
Recently, brown trout abundance has
increased in the upper Big Hole River
upstream of Wisdom (MFWP 2013e,
unpublished data). In the reach of the
upper Big Hole River where Arctic
grayling densities are highest, nonnative
trout abundances are lower than
upstream or downstream reaches, and
appear to have been stable since at least
2006 (Cayer 2013, unpublished data).
The potential effects of nonnative
trout species (rainbow, brown, brook,
and Yellowstone cutthroat trout) on
Arctic grayling recruitment are largely
unknown. Arctic grayling experts from
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Montana convened to explore such
effects predicted a less than 12 percent
reduction in Arctic grayling recruitment
when nonnative trout densities for any
species were 500 fish/mile or fewer, on
average (SSA 2014, p. 2). Predicted
reduction in Arctic grayling recruitment
when any of the nonnative species were
present at 1,000 fish/mile was higher
and similar among species (20 to 25
percent; SSA 2014, p. 2). These
estimates were derived with the
assumption that habitat was not a
limiting factor.
Currently, densities of nonnative trout
(brook, brown, rainbow) are fewer than
20 fish/mile (per species) in the
mainstem Big Hole River where Arctic
grayling densities are highest (Cayer
2013, unpublished data). Densities of
brown and rainbow trout are fewer than
20 fish/mile in Big Hole River
tributaries, while brook trout density in
tributaries is higher (∼80 fish/mile).
Brook trout density estimates only
include fish greater than 10 inches, thus
it is unknown how many total brook
trout reside in these areas. At current
densities of rainbow and brown trout,
effects on Arctic grayling recruitment
would be expected to be small, based on
the predictions of recruitment reduction
from nonnatives from the expert
meeting.
In the Madison River in and near
Ennis Reservoir, brown trout and
rainbow trout are abundant and are the
foundation of an important recreational
fishery (e.g., Byorth and Shepard 1990,
p. 1). Nonnative rainbow trout and
brown trout densities in the Madison
River near Ennis Reservoir are about
3,500 to 4,000 fish per mile (both
species included). These densities are
substantially higher than those observed
in other systems occupied by Arctic
grayling, and are higher than those
asked of the Arctic grayling experts to
predict effects of on Arctic grayling
recruitment. Arctic grayling abundance
in the Ennis Reservoir/Madison River
population appears to be suppressed
and declining (MFWP 2013f,
unpublished data). The relationship
between the higher densities of
nonnative trout and the low and
declining abundance of Arctic grayling
in this population is unclear. However,
the densities of nonnative trout
observed in the Madison River are not
representative of densities of nonnatives
in any of the other 19 populations of
Arctic grayling in the DPS. Thus, the
effect of nonnatives on Arctic grayling
recruitment is a concern in the Madison
River, but not in the rest of the DPS.
In the Centennial Valley, brook trout
and hybrid cutthroat trout (Yellowstone
cutthroat trout (Oncorhynchus clarkii
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49413
bouvieri) crossed with rainbow trout;
Mogen 1996, p. 42) have wellestablished populations and dominate
the abundance and biomass of the
salmonid community (Katzman 1998,
pp. 2–3; Boltz 2010, pers. comm.). In
Upper Red Rock Lake, hybrid cutthroat
trout and Arctic grayling exhibit some
dietary overlap (Cutting 2012,
unpublished data), although food may
not be a limiting factor in this system,
given the eutrophic, highly productive
nature of Upper Red Rock Lake (Jaeger
2014g, pers. comm.). In addition, hybrid
Yellowstone cutthroat trout in Upper
Red Rock Lake may occupy a similar
ecological niche once occupied by
native westslope cutthroat trout, a
species with which Arctic grayling coevolved. Thus, the adaptations Arctic
grayling developed over thousands of
years to coexist with westslope
cutthroat trout may be equally
advantageous when coexisting with
hybrid Yellowstone cutthroat trout.
Predation of Arctic grayling by brook
trout and hybrid cutthroat trout occurs
in Upper Red Rock Lake (Nelson 1954,
entire; Katzman 1998, pp. 35, 47, 114).
In 2013, the Service initiated a removal
effort to suppress hybrid cutthroat trout
in Red Rock Creek and Upper Red Rock
Lake. This effort will occur for 5 years,
during and after which the biological
response of Arctic grayling will be
documented. Currently, the relationship
between hybrid cutthroat trout and
Arctic grayling abundance in the
Centennial Valley is unclear. However,
a recent peak in hybrid cutthroat trout
abundance was paralleled by a peak in
Arctic grayling abundance, indicating
predation by hybrid cutthroat trout is
likely not a threat to Arctic grayling in
the Centennial Valley. It is plausible
that extensive macrophyte beds present
in Upper Red Rock Lake (Katzman 1998,
p. 81) provide complex hiding and
rearing cover for juvenile Arctic grayling
and minimize interactions between
young Arctic grayling and nonnative
fishes (Almany 2004, entire).
In the upper Missouri River basin, it
appears that the extent and magnitude
of competition and predation between
nonnative trout and Arctic grayling
likely depends on environmental
context (e.g., habitat type and quality,
environmental conditions such as
temperature, etc.) in most populations.
High-quality habitats likely provide
more food resources and complexity
(rearing areas) than lower quality
habitats (MacArthur and Levins 1967,
entire). These features of high-quality
habitats probably lessen competition
(MacArthur and Levins 1967, entire)
and reduce predation (Almany 2004, p.
107) by providing complex rearing areas
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for the vulnerable young life stages of
Arctic grayling. For these reasons
historically, when many of the fluvial
habitats were degraded, competition
and predation likely had a larger effect
of Arctic grayling populations than they
currently do. Certainly, competition and
predation are still occurring in habitats
occupied by both nonnatives and Arctic
grayling. However, the increase in
habitat quality observed in recent years,
particularly in the Big Hole River and
Centennial Valley, appear to have
minimized effects of competition and
predation on respective Arctic grayling
populations. The primary evidence of
this is recent trends showing increasing
numbers of both nonnatives and Arctic
grayling in systems with high-quality
habitat, including increasing brown
trout and Arctic grayling in the Big Hole
River. Other adfluvial populations of
Arctic grayling have coexisted with
brook, rainbow, and Yellowstone
cutthroat trout for extended periods of
time (>60 years) with no observed
declines in abundance.
Predation by Birds and Mammals
In general, the incidence and effect of
predation by birds and mammals on
Arctic grayling is not well understood
because few detailed studies have been
completed (Northcote 1995, p. 163).
Black bear (Ursus americanus), mink
(Neovison vison), and river otter (Lontra
canadensis) are present in southwestern
Montana, but direct evidence of
predatory activity by these species is
often lacking (Kruse 1959, p. 348).
Osprey (Pandion haliaetus) can capture
Arctic grayling during the summer
(Kruse 1959, p. 348). In the Big Hole
River, Byorth and Magee (1998, p. 926)
attributed the loss of Arctic grayling
from artificial enclosures used in a
competition experiment to predation by
minks, belted kingfisher (Ceryle alcyon),
osprey, and great blue heron (Ardea
herodia). In addition, American white
pelican (Pelecanus erythrorhynchos) are
seasonally present in the Big Hole River,
and they also may feed on Arctic
grayling. The aforementioned mammals
and birds can be effective fish predators;
however, Arctic grayling evolved with
these native predator species and have
developed life-history and reproductive
strategies to mitigate for predation
losses. We have no data demonstrating
any of these species historically or
currently consume Arctic grayling at
levels sufficient to exert a measureable,
population-level impact on native
Arctic grayling in the upper Missouri
River system. We expect the current
situation to continue, so we conclude
that predation by birds and mammals
does not constitute a threat to Missouri
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River Arctic grayling now or in the
future.
Summary of Factor C
Based on the information available at
this time, we conclude disease does not
represent a past or current threat to the
Upper Missouri River DPS of Arctic
grayling. We have no basis for
concluding that disease may become a
future threat.
Predation and competition can
influence the distribution, abundance,
and diversity of species in ecological
communities. Predation by and
competition with nonnative species can
negatively affect native species,
particularly those that are stressed or
occurring at low densities due to
unfavorable environmental conditions.
Historically, the impact of predation
and competition from nonnatives was
likely greater because many of the
habitats used by Arctic grayling were
degraded. Thus, predation and
competition likely played a role
historically in decreasing the abundance
and distribution of Arctic grayling.
Currently, habitat conditions have
improved markedly for those Arctic
grayling populations on Federal land
(18 of 20 populations) and for the Big
Hole River population on primarily
private land. Predation and competition
with nonnative species are still
occurring in these systems, although the
extent and magnitude of these effects
appears to be mediated by habitat
quality. Abundance of Arctic grayling
and nonnative brown trout are
increasing in the Big Hole River. Before
suppression efforts began, Yellowstone
cutthroat hybrids and Arctic grayling
spawners were both at 40 year highs in
Red Rock Creek in the Centennial
Valley. We acknowledge nonnative trout
densities are high in the Madison River
and may be contributing to the decline
of that Arctic grayling population;
however, most other adfluvial
populations appear to have stable
abundance of Arctic grayling and
nonnatives. Thus, based on our review
we have no information that predation
or competition represents a threat at the
DPS level on the Upper Missouri River
DPS of Arctic grayling. Further, Arctic
grayling experts project only a small
effect of predicted nonnative trout
densities on Arctic grayling recruitment
in the future. Thus, we have no
information that predation or
competition from nonnative trout
represents a future threat at the
population or species level.
Little is known about the effect of
predation on Arctic grayling by birds
and mammals. Such predation likely
does occur, but we are not aware of any
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situation where an increase in fisheating birds or mammals has coincided
with the decline of Arctic grayling.
Consequently, the available information
does not support a conclusion that
predation by birds or mammals
represents a substantial past, present, or
future threat to native Arctic grayling in
the upper Missouri River.
Factor D. The Inadequacy of Existing
Regulatory Mechanisms
Section 4(b)(1)(A) of the Act requires
the Service to take into account ‘‘those
efforts, if any, being made by any State
or foreign nation, or any political
subdivision of a State or foreign nation,
to protect such species . . .’’ We
consider relevant Federal, State, and
Tribal laws, and regulations when
evaluating the status of the species.
Regulatory mechanisms, if they exist,
may preclude the need for listing if we
determine that such mechanisms
adequately address the threats to the
species such that listing is not
warranted. Only existing ordinances,
regulations, and laws, that have a direct
connection to a law, are enforceable and
permitted are discussed in this section.
All other measures are discussed under
the specific relevant factor.
U.S. Federal Laws and Regulations
No Federal laws in the United States
specifically address the Arctic grayling,
but several, in their implementation,
may affect the species’ habitat.
National Environmental Policy Act
All Federal agencies are required to
adhere to the National Environmental
Policy Act (NEPA) of 1970 (42 U.S.C.
4321 et seq.) for projects they fund,
authorize, or carry out. The Council on
Environmental Quality’s regulations for
implementing NEPA (40 CFR parts
1500–1518) state that, when preparing
environmental impact statements,
agencies shall include a discussion on
the environmental impacts of the
various project alternatives, any adverse
environmental effects which cannot be
avoided, and any irreversible or
irretrievable commitments of resources
involved (40 CFR part 1502). The NEPA
itself is a disclosure law, and does not
require subsequent minimization or
mitigation measures by the Federal
agency involved. Although Federal
agencies may include conservation
measures for Arctic grayling as a result
of the NEPA process, any such measures
are typically voluntary in nature and are
not required by NEPA.
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Federal Land Policy and Management
Act
The Federal Land Policy and
Management Act (FLPMA) of 1976 (43
U.S.C. 1701 et seq.), as amended, states
that the public lands shall be managed
in a manner that will protect the quality
of scientific, scenic, historical,
ecological, environmental, air and
atmospheric, water resource, and
archeological values. This statute
protects lands within the range of the
Arctic grayling managed by the Bureau
of Land Management (BLM).
The BLM considers the fluvial Arctic
grayling a sensitive species requiring
special management consideration for
planning and environmental analysis
(BLM 2009a, entire, BLM 2009b, entire).
The BLM has recently developed a
resource management plan (RMP) for
the Dillon Field Office Area that
provides guidance for the management
of over 900,000 acres of public land
administered by BLM in southwest
Montana (BLM 2006a, p. 2). The Dillon
RMP area thus includes the geographic
area that contains the Big Hole, Miner,
Mussigbrod, Madison River, and
Centennial Valley populations of Arctic
grayling. A RMP planning area
encompasses all private, State, and
Federal lands within a designated
geographic area (BLM 2006a, p. 2), but
the actual implementation of the RMP
focuses on lands administered by the
BLM that typically represent only a
fraction of the total land area within that
planning area (BLM 2006b, entire).
Restoring Arctic grayling habitat and
ensuring the long-term persistence of
both fluvial and adfluvial ecotypes are
among the RMP’s goals (BLM 2006a, pp.
30–31). However, there is little actual
overlap between the specific parcels of
BLM land managed by the Dillon RMP
and the current distribution of Arctic
grayling (BLM 2006b, entire).
The BLM also has a RMP for the Butte
Field Office Area, which includes more
than 300,000 acres in south-central
Montana (BLM 2008, entire), including
portions of the Big Hole River in
Deerlodge and Silver Bow counties
(BLM 2008, p. 8; 2009c, entire). The
Butte RMP considers conservation and
management strategies and agreements
for Arctic grayling in its planning
process and includes a goal to
opportunistically enhance or restore
habitat for Arctic grayling (BLM 2008,
pp. 10, 30, 36). However, the Butte RMP
does not mandate specific actions to
improve habitat for Arctic grayling in
the Big Hole River and little overlap
exists between BLM-managed lands and
Arctic grayling occupancy in this
planning area.
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National Forest Management Act
Under the U.S. Forest Service (USFS)
National Forest Management Act
(NFMA) of 1976, as amended (16 U.S.C.
1600 et seq.), the USFS strives to
provide for a diversity of plant and
animal communities when managing
national forest lands. Individual
national forests may identify species of
concern that are significant to each
forest’s biodiversity. The USFS
considers fluvial Arctic grayling a
sensitive species (USFS 2004, entire) for
which population viability is a concern.
However, this designation provides no
special regulatory protections.
Most of the upper Missouri River
grayling populations occur on National
Forest land; all 16 adfluvial populations
and the fluvial Ruby River population
(majority on National Forest) occur on
USFS-managed lands. These
populations occur across four different
National Forests; consequently the
riparian habitats surrounding the lakes
and tributaries are managed according
to the standards and guidelines outlined
in each National Forest Plan. All Forest
Plans do not contain the same standards
and guidelines; however, each Plan has
standards and guidelines for protecting
riparian areas around perennial water
sources. In the Beaverhead-Deerlodge
and Helena National Forest Plans, the
Inland Native Fish Strategy (INFS)
standards and guidelines have been
incorporated. The INFS, in part, defines
widths of riparian buffer zones adequate
to protect streams and lakes from nonchannelized sediment inputs and
contribute to other riparian functions,
such as stream shading and bank
stability. These protections have been
incorporated into the BeaverheadDeerlodge and Helena National Forest
Plans through amendments and are
currently preserving intact riparian
areas around most, if not all, adfluvial
Arctic grayling habitats. Exceptions to
the riparian protections outlined in
INFS are occasionally granted; however,
these exceptions require an analysis of
potential effects and review by a USFS
fish biologist.
On the Gallatin National Forest,
standards and guidelines in the Forest
Plan include using ‘‘best management
practices (BMPs)’’ to protect water
sources and riparian areas. Similar to
INFS, BMPs outline buffer strips along
watercourses where disturbance and
activity is minimized to protect riparian
areas and water quality. On the Lewis
and Clark National Forest, standards
and guidelines are in place to leave
timbered buffer strips adjacent to
waterbodies to protect riparian areas.
Grayling habitat on the Gallatin and
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Lewis and Clark National Forests
consists of seven high-elevation
mountain lakes.
The NFMA and INFS are adequately
protecting riparian habitat on National
Forest land, given the intact nature of
most riparian areas surrounding the
high-elevation lake populations and the
Ruby River.
National Park Service (NPS) Organic Act
The NPS Organic Act of 1916 (16
U.S.C. 1 et seq.), as amended, states that
the NPS ‘‘shall promote and regulate the
use of the Federal areas known as
national parks, monuments, and
reservations . . . to conserve the
scenery and the national and historic
objects and the wild life therein and to
provide for the enjoyment of the same
in such manner and by such means as
will leave them unimpaired for the
enjoyment of future generations.’’ Arctic
grayling are native to the western part
of Yellowstone National Park and
habitats are managed accordingly for the
species under the Native Species
Management Plan (NPS 2010, entire).
One adfluvial Arctic grayling
population, Grebe Lake, currently
occurs in Yellowstone National Park.
The Grebe Lake population is one of the
larger adfluvial populations (see Table
3, above) in the DPS. The habitat in
Grebe Lake and the tributaries is
managed for conservation (NPS 2010, p.
44). Further, it is expected that these
habitats will be managed for
conservation in the future, based on
provisions in the Organic Act and
guidance outlined in the Native Species
Management Plan.
National Wildlife Refuge System
Improvement Act of 1997
The National Wildlife Refuge Systems
Improvement Act (NWRSIA) of 1997
(Pub. L. 105–57) amends the National
Wildlife Refuge System Administration
Act of 1966 (16 U.S.C. 668dd et seq.).
The NWRSIA directs the Service to
manage the Refuge System’s lands and
waters for conservation. The NWRSIA
also requires monitoring of the status
and trends of refuge fish, wildlife, and
plants. The NWRSIA requires
development of a comprehensive
conservation plan (CCP) for each refuge
and management of each refuge
consistent with its plan.
The Service has developed a final
CCP to provide a foundation for the
management and use of Red Rock Lakes
NWR (USFWS 2009, entire) in the
Centennial Valley. Since the
development of the CCP, Refuge staff
have conducted numerous habitat
conservation/restoration projects to
benefit Arctic grayling, including:
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Removal of an earthen dam whose
reservoir inundated several hundred
meters of historical Arctic grayling
spawning habitat in Elk Springs Creek,
and subsequent reintroductions and
tracking of young-of-year Arctic grayling
in Elk Springs Creek (West 2014a, pers.
comm.). However to date, the
reintroductions in Elk Springs Creek
have not established a spawning run.
Other conservation projects conducted
on the Refuge include the acquisition of
new land and decreases in grazing
intensities from 20,000 AUMs to about
5,000 AUMs. The Refuge has
implemented a rest-rotation grazing
system where more durable lands are
grazed while more sensitive lands (e.g.,
riparian areas) are rested for up to 4
years (West 2014a, pers. comm.). Some
active riparian restoration has also
occurred, including a project to
reconnect Red Rock Creek to a historical
channel and replacement of four
culverts to allow for natural tributary
migration across alluvial fans (West
2014a, pers. comm.). The Refuge is also
actively engaged in supporting ongoing
graduate research efforts to explore
potential limiting factors for Arctic
grayling in the Centennial Valley.
Other conservation projects under the
CCP have been focused on potential
nonnative species effects on Arctic
grayling, namely a 5-year project
removing hybrid cutthroat trout
captured during their upstream
spawning run and a study of dietary
overlap between Arctic grayling and
Yellowstone cutthroat trout (West
2014a, pers. comm.). The Refuge also
operates a sill dam (previous upstream
fish barrier) to provide upstream fish
passage and operates one irrigation
ditch only when snowpack is average or
above and timing is such that young
Arctic grayling are not present near the
diversion (West 2014a, pers. comm.).
The NWRSIA is adequately protecting
habitat for Arctic grayling on the Refuge
because riparian habitats are improving
and the Centennial Valley population is
increasing in both abundance and
distribution. The proven track record of
completed conservation projects on the
refuge and currently expanding Arctic
grayling population indicate that the
continued implementation of the CCP
during the next 15 years (which is the
life of the CCP) will continue to improve
habitat conditions on the refuge.
Federal Power Act (FPA)
The Federal Power Act of 1920 (16
U.S.C. 791 et seq., as amended) provides
the legal authority for the Federal
Energy Regulatory Commission (FERC),
as an independent agency, to regulate
hydropower projects. In deciding
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whether to issue a license, FERC is
required to give equal consideration to
mitigation of damage to, and
enhancement of, fish and wildlife (16
U.S.C. 797(e)). A number of FERClicensed dams exist in the Missouri
River basin in current (i.e., Ennis Dam
on the Madison River) and historical
Arctic grayling habitat (e.g., Hebgen
Dam on the Madison River; Hauser,
Holter, and Toston dams on the
mainstem Missouri River; and Clark
Canyon Dam on the Beaverhead River).
The FERC license expiration dates for
these dams range from 2024 (Toston) to
2059 (Clark Canyon) (FERC 2010,
entire). None of these structures
provides upstream passage of fish, and
such dams are believed to be one of the
primary factors that led to the historical
decline of Arctic grayling in the
Missouri River basin (see discussion
under Factor A, above). However, recent
monitoring data indicate multiple stable
Arctic grayling populations occurring
above mainstem dams, with the
exception of the Ennis Reservoir/
Madison River population. The
drawdowns in reservoir water level
believed to have historically affected the
Ennis Reservoir/Madison River Arctic
grayling population are not permitted
under a new licensing agreement
between the Federal Energy Regulatory
Commission and Madison Dam
operators, as we described previously in
this finding (Clancey 2014, pers.
comm.). This change in water
management in Ennis Reservoir will
ensure adequate rearing and foraging
habitat for this population. The fluvial
ecotype is still represented in the DPS
and both strictly fluvial Arctic grayling
populations appear to be stable or
increasing. Thus, we conclude the
Federal Power Act is currently adequate
to protect the Upper Missouri River DPS
of Arctic grayling at the population and
DPS level.
Clean Water Act
The Clean Water Act (CWA) of 1972
(33 U.S.C. 1251 et seq.) establishes the
basic structure for regulating discharges
of pollutants into the waters of the
United States and regulating quality
standards for surface waters. The CWA’s
general goal is to ‘‘restore and maintain
the chemical, physical, and biological
integrity of the Nation’s waters’’ (33
U.S.C. 1251(a)). The CWA requires
States to adopt standards for the
protection of surface water quality and
establishment of total maximum daily
load (TMDL) guidelines for rivers. The
Big Hole River has approved TMDL
plans for its various reaches (MDEQ
2009a, entire; 2009b, entire); thus,
complete implementation of this plan
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should improve water quality (by
reducing water temperatures, and
reducing sediment and nutrient inputs)
in the Big Hole River in the future. As
of September 2013, there was no
significant TMDL plan development
activity in the Madison River or Red
Rock watershed in the Centennial
Valley (see MDEQ 2014). Currently,
TMDL documents have been approved
for the Ruby River. All planning areas
containing other adfluvial Arctic
grayling populations in the upper
Missouri River basin have approved
TMDLs, including the Gallatin, Lake
Helena, and Sun watersheds (see MDEQ
2014).
Currently, water temperatures in the
Big Hole River exceed levels outlined in
the TMDL. However, reductions in
water temperature within tributaries
have been demonstrated (see discussion
under Factor A and Table 4). Given that
most Arctic grayling populations within
the upper Missouri River basin are
stable or increasing and habitats are
largely being managed in a manner that
benefits the species, we have no
evidence that the CWA is inadequately
protecting Arctic grayling at the
population or DPS level.
State Laws
Montana Environmental Policy Act
The legislature of Montana enacted
the Montana Environmental Policy Act
(MEPA) as a policy statement to
encourage productive and enjoyable
harmony between humans and their
environment, to protect the right to use
and enjoy private property free of undue
government regulation, to promote
efforts that will prevent or eliminate
damage to the environment and
biosphere and stimulate the health and
welfare of humans, to enrich the
understanding of the ecological systems
and natural resources important to the
State, and to establish an environmental
quality council (MCA 75–1–102). Part 1
of the MEPA establishes and declares
Montana’s environmental policy. Part 1
has no legal requirements, but the
policy and purpose provide guidance in
interpreting and applying statutes. Part
2 requires State agencies to carry out the
policies in Part 1 through the use of
systematic, interdisciplinary analysis of
State actions that have an impact on the
human environment. This is
accomplished through the use of a
deliberative, written environmental
review. In practice, MEPA provides a
basis for the adequate review of State
actions in order to ensure that
environmental concerns are fully
considered (MCA 75–1–102). Similar to
NEPA, the MEPA is largely a disclosure
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law and a decision-making tool that
does not specifically require subsequent
minimization or mitigation measures.
Laws Affecting Physical Aquatic
Habitats
A number of Montana State laws have
a permitting process applicable to
projects that may affect stream beds,
river banks, or floodplains. These
include the Montana Stream Protection
Act (SPA), the Streamside Management
Zone Law (SMZL), and the Montana
Natural Streambed and Land
Preservation Act (Montana Department
of Natural Resources (MDNRC) 2001,
pp. 7.1–7.2). The SPA requires that a
permit be obtained for any project that
may affect the natural and existing
shape and form of any stream or its
banks or tributaries (MDNRC 2001, p.
7.1). The Montana Natural Streambed
and Land Preservation Act (i.e.,
MNSLPA or 310 permit) requires
private, nongovernmental entities to
obtain a permit for any activity that
physically alters or modifies the bed or
banks of a perennially flowing stream
(MDNRC 2001, p. 7.1). The SPA and
MNSLPA laws do not mandate any
special recognition for species of
concern, but in practice, biologists that
review projects permitted under these
laws usually stipulate restrictions to
avoid harming such species (Horton
2010, pers. comm.). The SMZL regulates
forest practices near streams (MDNRC
2001, p. 7.2). The Montana Pollutant
Discharge Elimination System (MPDES)
Stormwater Permit applies to all
discharges to surface water or
groundwater, including those related to
construction, dewatering, suction
dredges, and placer mining, as well as
to construction that will disturb more
than 1 acre within 100 ft (30.5 m) of
streams, rivers, or lakes (MDNRC 2001,
p. 7.2).
Review of applications by MFWP,
MTDEQ, or MDNRC is required prior to
issuance of permits under the above
regulatory mechanisms (MDNRC 2001,
pp. 7.1–7.2). These regulatory
mechanisms are expected to limit
impacts to aquatic habitats in general.
Given that most Arctic grayling
populations are stable or increasing in
abundance in the presence of these
regulatory mechanisms, we have no
basis for concluding that these
regulatory mechanisms are inadequate
to protect the Arctic grayling and their
habitat now or in the future.
Montana Water Use Act
The purpose of the Montana Water
Use Act (Title 85: Chapter 2, Montana
Codes Annotated) is to provide water for
existing and future beneficial use and to
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maintain minimum flows and water
quality in Montana’s streams. The
Missouri River system is generally
believed to be overappropriated, and
water for additional consumptive uses is
only available for a few months during
very wet years (MDNRC 1997, p. 12).
However, the upper Missouri River
basin and Madison River basin have
been closed to new water appropriations
because of water availability problems,
overappropriation, and a concern for
protecting existing water rights (MDNRC
2009, p. 45). In addition, recent
compacts (a legal agreement between
Montana, a Federal agency, or an Indian
tribe determining the quantification of
federally or tribally claimed water
rights) have been signed that close
appropriations in specific waters in or
adjacent to Arctic grayling habitats. For
example, the USFWS–Red Rock Lakes–
Montana Compact includes a closure of
appropriations for consumptive use in
the drainage basins upstream of the
most downstream point on the Red Rock
Lakes NWR and the Red Rock Lakes
Wilderness Area (MDNRC 2009, pp. 18,
47). The NPS–Montana Compact
specifies that certain waters will be
closed to new appropriations when the
total appropriations reach a specified
level, and it applies to Big Hole National
Battlefield and adjacent waters (North
Fork of the Big Hole River and its
tributaries including Ruby and Trail
Creeks), and the portion of Yellowstone
National Park that is in Montana
(MDNRC 2009, p. 48).
The State of Montana is currently
engaged in a Statewide effort to
adjudicate (finalize) water rights
claimed before July 1, 1973. The final
product of adjudication in a river basin
is a final decree. To reach completion,
a decree progresses through several
stages: (1) Examination, (2) temporary
preliminary decree, (3) preliminary
decree, (4) public notice, (5) hearings,
and (6) final decree (MDNRC 2009, pp.
9–14). As of February 2014, the
Centennial Valley has a preliminary
decree, and the Big Hole and Madison
Rivers have preliminary temporary
decrees (MDNRC 2014, entire). We
anticipate the final adjudication of all
the river basins in Montana that
currently contain native Arctic grayling
will be completed in the next 5 years,
but we do not know if this process will
eliminate the overallocation of water
rights. We note that the overallocation
of water in some systems within the
upper Missouri river basin is of general
concern to Arctic grayling because of
the species’ need for adequate quantity
and quality of water for all life stages.
However, we have no information
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indicating that overallocation of water
in the upper Missouri River basin is a
current threat at the individual or DPS
level because most populations are
stable or increasing at this time.
Therefore, we conclude that the
Montana Water Use Act is adequate to
protect the Arctic grayling and its
habitat.
Angling Regulations
Arctic grayling is considered a game
fish (MFWP 2010, p. 16), but is subject
to special catch-and-release regulations
in streams and rivers within its native
range, as was described under Factor B,
above (MFWP 2014d, p. 51). Catch-andrelease regulations also are in effect for
Ennis Reservoir on the Madison River
and Red Rock Creek in the Centennial
Valley (MFWP 2014d, p. 63). Arctic
grayling in other adfluvial populations
are subject to more liberal regulations;
anglers can keep up to 5 per day and
have up to 10 in possession in
accordance with standard daily and
possession limits for that angling
management district (MFWP 2014d, p.
51). We have no evidence to indicate
that current fishing regulations are
inadequate to protect native Arctic
grayling in the Missouri River basin (see
discussion under Factor B, above).
Summary of Factor D
Current Federal and State regulatory
mechanisms are adequate to protect
Arctic grayling of the upper Missouri
River. We conclude this because the
majority of populations are on Federal
land where regulatory mechanisms are
in place to preserve intact habitats and
are expected to remain in place. In the
Big Hole River, fluvial Arctic grayling
generally occupy waters adjacent to
private lands (MFWP et al. 2006, p. 13;
Lamothe et al. 2007, p. 4), so Federal
regulations may have limited ability to
protect that population. However, some
Federal regulations (e.g., CWA, FPA,
NMFA, NWRSIA, NPS Organic Act) in
concert with other existing conservation
efforts (e.g., Big Hole CCAA) are
adequate to sustain and improve habitat
conditions for Arctic grayling. Arctic
grayling in the Big Hole River appear to
be responding positively to these
improvements. In addition, we did not
identify other threats to the DPS that
would require regulatory protections.
For the reasons described above, we
conclude that existing regulatory
mechanisms are adequate to protect the
Upper Missouri River DPS of Arctic
grayling. We do not anticipate any
changes to the existing regulatory
mechanisms; thus we conclude that
existing regulatory mechanisms will
remain adequate in the future.
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Factor E. Other Natural or Manmade
Factors Affecting Its Continued
Existence
Drought
Drought is a natural occurrence in the
interior western United States (see
National Drought Mitigation Center
2010). The duration and severity of
drought in Montana appears to have
increased during the last 50 years, and
precipitation has tended to be lower
than average in the last 20 years
(National Climatic Data Center 2010).
Drought can affect fish populations by
reducing stream flow volumes. This
leads to dewatering and high
temperatures that can limit connectivity
among spawning, rearing, and sheltering
habitats. Drought can also reduce the
volume of thermally suitable habitat and
increase the frequency of water
temperatures above the physiological
limits for optimum growth and survival
in Arctic grayling. In addition, drought
can interact with human-caused
stressors (e.g., irrigation withdrawals,
riparian habitat degradation) to further
reduce stream flows and increase water
temperatures.
Reduced stream flows and elevated
water temperatures during drought have
been most apparent in the Big Hole
River system (Magee and Lamothe 2003,
pp. 10–14; Magee et al. 2005, pp. 23–25;
Rens and Magee 2007, pp. 11–12, 14). In
the Big Hole River, evidence for the
detrimental effects of drought on Arctic
grayling populations is primarily
inferential; observed declines in fluvial
Arctic grayling and nonnative trout
abundances in the Big Hole River
coincide with periods of drought (Magee
and Lamothe 2003, pp. 22–23, 28) and
fish kills (Byorth 1995, pp. 10–11, 31).
Although the response of stream and
river habitats to drought is expected to
be most pronounced because of the
strong seasonality of flows in those
habitats, effects in lake environments
can occur. For example, both the Upper
and Lower Red Rock Lakes are very
shallow (Mogen 1996, p. 7). Increased
frequency or duration of drought could
lead to increased warming in shallower
lakes, such as Upper Red Rock Lake.
However, the Centennial Valley has
many springs sources that could, at least
in part, mitigate for increases in water
temperature due to increased drought
frequency and magnitude. Other
potential effects from drought could
include a reduction in overall lake
depth, which could in turn affect
summer or overwintering habitat.
Adfluvial populations in high mountain
lakes would likely not be affected
significantly by drought because air
(and thus water) temperatures in these
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habitats are relatively cool due to the
greater distance from sea level at high
elevations (∼ a 3.6 °F (6.5 °C) decrease
in air temperature for every 3,200 ft. (1
kilometer) above sea level; Physics
2014). In addition, most of these
habitats are relatively large bodies of
water volumetrically, thus are resistant
to warming, given the high specific heat
of water (USGS 2014). Further, intact
riparian areas in these habitats buffer
against water temperature increases in
tributaries by blocking incoming solar
radiation (Sridhar et al. 2004, entire;
Cassie 2006, p. 1393).
Given the climate of the
intermountain West, we conclude that
drought has been and will continue to
be a natural occurrence. We assume that
negative effects of drought on Arctic
grayling populations, such as reduced
connectivity among habitats or
increased water temperatures at or
above physiological thresholds for
growth and survival, are more frequent
in stream and river environments and in
very shallow lakes relative to larger,
deeper lakes. As discussed under Factor
A, the implementation of the Big Hole
Arctic grayling CCAA is likely to
minimize some of the effects of drought
in the Big Hole River, by reducing the
likelihood that human-influenced
actions or outcomes (irrigation
withdrawals, destruction of riparian
habitats, and fish passage barriers) will
interact with the natural effects of
drought (reduced stream flows and
increased water temperatures). We
expect the impact of drought may act at
the individual level, but not at the
population or DPS level because most
grayling populations reside in droughtresistant habitats in high mountain
lakes. Some populations will likely be
affected by drought, but implemented
conservation measures (Big Hole River
population) and natural spring sources
(Centennial Valley) are expected to
minimize the impact. Overall, we
conclude that drought has been a past
threat when many historical habitats
were degraded, but is not a current
threat because of the intact nature of
most habitats occupied by Arctic
grayling in the upper Missouri River
basin. Drought is expected to increase in
both duration and severity in the future;
however, resiliency currently being
incorporated into riparian and aquatic
habitats through conservation projects
will likely buffer the effects of drought.
Thus, drought is not expected to pose a
threat to the DPS in the future.
Stochastic (Random) Threats, Genetic
Diversity and Small Population Size
A principle of conservation biology is
that the presence of larger and more
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productive (resilient) populations can
reduce overall extinction risk. To
minimize extinction risk due to
stochastic (random) threats, life-history
diversity should be maintained,
populations should not all share
common catastrophic risks, and both
widespread and spatially close
populations are needed (Fausch et al.
2006, p. 23; Allendorf et al. 1997,
entire).
The Upper Missouri River DPS of
Arctic grayling exists largely as a
collection of isolated populations
(Peterson and Ardren 2009, entire), with
little to no gene flow among
populations. While the inability of fish
to move between populations limits
genetic exchange and demographic
support (Hilderbrand 2003, p. 257),
large population sizes coupled with
adequate number of breeding
individuals minimize the effects of
isolation. For example, Grebe Lake, a
large population, receives no genetic
infusion from any other population in
the upper Missouri River basin, yet has
a very large estimated effective
population size (see Table 3, above).
Loss of genetic diversity from genetic
drift is not a concern for this
population, despite it being
reproductively isolated.
Abundance among the 20 Arctic
grayling populations varies widely (see
Table 3, above). Individually, small
populations like Ruby River need to
maintain enough adults to minimize
loss of variability through genetic drift
and inbreeding (Rieman and McIntyre
1993, pp. 10–11). The point estimates of
the effective number of breeders
observed in all populations (where data
are available) are above the level at
which inbreeding is an immediate
concern (Leary 2014, pers. comm.). The
Ruby River population exhibits a low
effective number of breeders, but
contains the second highest genetic
diversity among all populations (Leary
2014, unpublished data). Thus,
inbreeding depression is probably not a
concern for this population in the near
future (Leary 2014, pers. comm.).
Effective population size estimates for
other Arctic grayling populations vary
from 162 to 1,497 (see Table 3, above).
There has been considerable debate
about what effective population size is
adequate to conserve genetic diversity
and long-term adaptive potential (see
Jamieson and Allendorf 2012 for review,
p. 579). However, loss of genetic
diversity is typically not an immediate
threat even in isolated populations with
an Ne >100 (Palstra and Ruzzante 2008,
p. 3441), but rather is a symptom of
deterministic processes acting on the
population (Jamieson and Allendorf
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2012, p. 580). In other words, loss of
genetic diversity due to small effective
population size typically does not drive
species to extinction (Jamieson and
Allendorf 2012, entire); other processes,
such as habitat degradation, have a more
immediate and greater impact on
species persistence (Jamieson and
Allendorf 2012). We acknowledge that
loss of genetic diversity can occur in
small populations; however, in this
case, it appears that there are adequate
numbers of breeding adults to minimize
loss of genetic diversity. Thus, we
conclude that loss of genetic diversity is
not a threat at the DPS level.
Conservation of life-history diversity
is important to the persistence of
species confronted by habitat change
and environmental perturbations
(Beechie et al. 2006, entire). Therefore,
the reintroductions of fluvial Arctic
grayling into the upper Ruby River that
have occurred provide redundancy of
the fluvial ecotype. The number of
breeding individuals in the Ruby River
population has increased over the last 3
years (Leary 2014, unpublished data).
Thus, there is now a viable replicate of
the fluvial ecotype.
Populations of Arctic grayling in the
Upper Missouri River DPS are for the
most part widely separated from one
another, occupying 7 of 10 historically
occupied watersheds (see Table 3,
above). Thus, risk of extirpation by a
rare, high-magnitude environmental
disturbance (i.e., catastrophe) is
relatively low. In addition, multiple
spawning locations exist for 11 of the 20
populations in the Upper Missouri River
DPS. The 11 populations with access to
multiple spawning tributaries include
all the largest populations in terms of
abundance, except Mussigbrod Lake
(see Table 3). Abundance and number of
breeding individuals is adequate in
most populations to sustain moderate to
high levels of genetic diversity currently
observed. Based on this information, we
conclude that stochastic processes are
not a threat to the Upper Missouri River
DPS of Arctic grayling and are not
expected to be in the future.
Summary of Factor E
Overall, we conclude that the Upper
Missouri River DPS of Arctic grayling
has faced historical threats from
drought, loss of genetic diversity, and
small population size. However, the
DPS currently exists as multiple,
isolated populations across a
representative portion of its historical
range. While reproductive isolation can
lead to detrimental genetic effects, the
current size of most Arctic grayling
populations, trends in effective
population size, and number of breeders
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suggest these effects will be minimal.
Redundancies within and among
populations are present: Multiple
spawning tributaries, geographic
separation, life-history replication.
Given this information, we conclude the
redundant nature of multiple resilient
populations across a representative
portion of the species’ historical range
minimizes the impacts of drought, low
abundance, reduced genetic diversity,
and lack of a fluvial ecotype replicate.
Thus, these are not current threats, and
are not expected to be threats in the
future.
Cumulative Effects From Factors A
Through E
We limit our discussion of cumulative
effects from Factors A through E to
interactions involving climate change.
Our rationale for this is that climate
change has the highest level of
uncertainty among other factors
considered, and likely has the most
potential to affect Arctic grayling
populations when interacting with other
factors.
Climate Change and Nonnative Species
Interactions
Changes in water temperature due to
climate change may influence the
distribution of nonnative trout species
(Rahel and Olden 2008, p. 524) and the
outcome of competitive interactions
between those species and Arctic
grayling. Brown trout are generally
considered to be more tolerant of warm
water than many salmonid species
common in western North America
(Coutant 1999, pp. 52–53; Selong et al.
2001, p. 1032), and higher water
temperatures may favor brown trout
where they compete against salmonids
with lower thermal tolerances (Rahel
and Olden 2008, p. 524). Recently,
observed increases in the abundance
and distribution of brown trout in the
upper reaches of the Big Hole River
(MFWP 2013, unpublished data) may be
consistent with the hypothesis that
stream warming is facilitating
encroachment. However, the effect of
increased abundance and distribution of
brown trout on Arctic grayling in the
Big Hole River is unknown.
Currently, brown trout are at
relatively low densities (<20 fish/mile)
in the upper Big Hole River, where
Arctic grayling densities are highest
(MFWP 2013e, unpublished data). At
densities of 100 brown trout per mile (a
plausible future scenario), Arctic
grayling experts predicted a 5 percent
reduction in Arctic grayling recruitment
in the Big Hole River, due to
competition and predation (SSA 2014,
p. 2). Given that natural mortality of
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salmonid fry is typically high (≤90
percent) (Kruse 1959, pp. 329, 333;
Bradford 1995, p. 1330), the predicted
reductions in Arctic grayling
recruitment by current and future
densities of brown trout in the Big Hole
River will likely not impact Arctic
grayling at the population level. Thus,
the potential cumulative effect of
climate change and nonnative species
interactions is not a current or future
threat for the Upper Missouri River DPS
of Arctic grayling.
Climate Change and Dewatering
Synergistic interactions are possible
between effects of climate change and
effects of other potential stressors such
as dewatering. Increases in temperature
and changes in precipitation are likely
to affect the availability of water in the
West. However, it is difficult to project
how climate change will affect water
availability because increased air and
water temperatures may be
accompanied and tempered by more
frequent precipitation events.
Uncertainty about how different
temperature and precipitation scenarios
could affect water availability make
projecting possible synergistic effects of
climate change on the Arctic grayling
too speculative at this time.
Summary
Recent genetic analyses have
concluded that many of the introduced
populations of Arctic grayling in the
upper Missouri River basin contain
moderate to high levels of genetic
diversity and that these populations
were created from local sources within
the basin. These introduced populations
currently occur within the confines of
the upper Missouri River basin and
occupy high quality habitats on Federal
land, the same places the Service would
look to for long-term conservation of the
species, if needed. As such, these
populations and their future adaptive
potential have conservation value and
are included in the Upper Missouri
River DPS of Arctic grayling.
Currently, we recognize 20
populations of Arctic grayling in the
Upper Missouri River DPS, 18 of which
occur on Federal land. Adequate
regulatory mechanisms exist to ensure
the conservation of habitat on Federal
land for these populations. Historical
habitat degradation on private land has
affected the Big Hole River population;
however, habitat conditions have been
improving since the implementation of
the Big Hole CCAA in 2006.
Conservation actions associated with
the Big Hole CCAA and SHCP have
reduced water temperatures in
tributaries, increased instream flows in
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tributaries and the mainstem Big Hole
River, connected almost all core habitat
for Arctic grayling, and improved
riparian health. Arctic grayling have
responded favorably to these
improvements because abundance and
distribution have increased throughout
the upper Big Hole River, and number
of breeding adults has increased by a
factor of at least 5 since 2006. The
Service is encouraged by the successful
track record of conservation actions
implemented under the Big Hole CCAA
and SHCP over the past 7 years.
Riparian restoration efforts in the Big
Hole River and Centennial Valley are
ongoing and will continue to be key in
mitigating the anticipated effects of
drought and climate change. Increased
shading of tributaries and decreased
width-to-depth ratios in stream
channels can effectively minimize
effects from increasing air temperatures
and drought. In addition, these changes
to habitat can alter predation and
competition potential where both
nonnative species and Arctic grayling
coexist, as they have for over 100 years
in some populations.
We acknowledge the uncertainty
regarding the current status of the Ennis
Reservoir/Madison River population
and probable declining trend in
abundance. The factors influencing the
current demographics of this population
are unclear. However, we are
encouraged by the recent FERC
relicensing agreement precluding
reservoir drawdowns that likely affected
this population and its habitat in the
past.
In conclusion, we find viable
populations of both ecotypes present in
the DPS, the majority of which occur on
Federal land and are protected by
Federal land management measures.
Numbers of breeding adults are
currently increasing in both strictly
fluvial populations and in the
Centennial Valley. High-quality habitat
is present for most populations or is
improving where it is not optimal (e.g.,
Big Hole River). Health of riparian areas
is trending upward and will be key to
minimizing effects of climate change
and drought. All Arctic grayling
populations are genetically diverse, are
of Montana-origin, and occur in 7 of 10
historically occupied watersheds.
In 2010, we identified multiple
threats as acting on the Upper Missouri
River DPS of Arctic grayling. At that
time, we determined that habitat-related
threats included habitat fragmentation,
dewatering, thermal stress, entrainment,
riparian habitat loss, and effects from
climate change. Since 2010, we have 4
additional years of monitoring data and
have gained new insight. It is now
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apparent that these threats are being
effectively mitigated on private land
(Big Hole River) by conservation actions
under the Big Hole CCAA and do not
appear to be present or acting at a level
to warrant concern on most of the
adfluvial populations. Almost all (98
percent) of Arctic grayling core habitat
in the Big Hole River is now connected.
Recent riparian restoration activities
have appreciably reduced water
temperatures and improved riparian
habitat in tributaries to the Big Hole
River and are expected to buffer the
effects of climate change. Entrainment
of Arctic grayling into irrigation canals
in the Big Hole system is low, with no
documented entrainment occurring
since 2010. Habitats on Federal land are
largely intact and these populations are
not subject to many of the stressors
historically identified for other
populations because no irrigation
diversions are present, habitats are
primarily high-elevation lakes that have
cool water temperatures, and riparian
areas are largely intact.
In 2010, another threat identified as
acting on the Upper Missouri River DPS
of Arctic grayling was the presence of
nonnative trout. We considered
nonnative trout a threat at that time
because we were aware of several
instances where Arctic grayling declines
had occurred following nonnative trout
introductions. Currently, we have a
better understanding of the interactions
between nonnative trout and Arctic
grayling. Our review of these
interactions and case histories suggests
that habitat degradation, concurrent
with nonnative trout introductions,
likely contributed to historical declines
in Arctic grayling in those instances.
Further, it appears the effect of
nonnative trout on Arctic grayling are
likely habitat-mediated; nonnative trout
affect Arctic grayling disproportionately
when habitat conditions are degraded,
but both Arctic grayling and nonnatives
can coexist at viable levels when habitat
conditions are improved. The primary
evidence supporting this assertion is the
increasing abundance and distribution
of both Arctic grayling and nonnatives
in the Big Hole River (brown trout) and
Centennial Valley (Yellowstone
cutthroat trout before suppression
began). Another line of evidence to
support this assertion is observed
spatial segregation between nonnatives
and Arctic grayling in the core Arctic
grayling areas in the Big Hole River,
especially spawning and rearing areas
(SSA 2014). In addition, Arctic grayling
in adfluvial habitats have maintained
stable or increasing population levels in
the presence of brook, rainbow, and
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Yellowstone cutthroat trout for over 100
years in many instances in the upper
Missouri River basin, where habitat
degradation has not occurred or been
extensive.
In 2010, we stated that existing
regulatory mechanisms were inadequate
to protect the Upper Missouri River DPS
of Arctic grayling. The primary reason
for this assertion was that Arctic
grayling populations were reported as
declining; thus existing regulatory
mechanisms were believed to be
inadequate because they had failed to
halt or reverse this decline. Currently,
we have updated information indicating
that 19 of 20 populations of Arctic
grayling are either stable or increasing.
Existing regulatory mechanisms have
precluded riparian habitat destruction
on Federal lands or mandated
restoration of impaired areas and are
expected to provide similar protections
in the future. Given the updated
information, we now believe these
regulatory mechanisms are adequate.
In 2010, we identified reduced genetic
diversity, low abundance, random
events, drought, and lack of a fluvial
replicate as threats to the Upper
Missouri River DPS of Arctic grayling.
Updated genetic information that was
not available in 2010 indicates moderate
to high levels of genetic diversity within
most Arctic grayling populations in the
DPS. Further, abundance estimates
derived from this updated genetic
information indicate higher Arctic
grayling abundances than previously
thought. Adequate redundancy exists
within the DPS to minimize the effects
of random events and drought; lake
habitats occupied by most Arctic
grayling populations are droughtresistant. Lastly, a viable fluvial
replicate now exists (Ruby River), with
5 years of natural reproduction
documented and an increasing number
of breeding adults.
Finding
As required by the Act, we considered
the five factors in assessing whether the
Upper Missouri River DPS of Arctic
grayling is endangered or threatened
throughout all of its range. We
examined the best scientific and
commercial information available
regarding the present and future threats
faced by the Upper Missouri River DPS
of Arctic grayling. We reviewed the
petition, information available in our
files and other available published and
unpublished information, including
information submitted by the public,
and we consulted with recognized
Arctic grayling experts and other
Federal and State agencies. Habitatrelated threats previously identified,
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including habitat fragmentation,
dewatering, thermal stress, entrainment,
riparian habitat loss, and effects from
climate change, have been sufficiently
ameliorated and the information
indicates that 19 of 20 populations of
Arctic grayling are either stable or
increasing. On the basis of the best
scientific and commercial information
available and the analysis provided
above, we find that the magnitude and
imminence of threats do not indicate
that the Upper Missouri River DPS of
Arctic grayling is in danger of extinction
(endangered), or likely to become
endangered within the foreseeable
future (threatened), throughout its
range. Therefore, we find that listing the
Upper Missouri River DPS throughout
its range as a threatened or an
endangered species is not warranted at
this time.
Significant Portion of the Range
Under the Act and our implementing
regulations, a species may warrant
listing if it is an endangered or a
threatened species throughout all or a
significant portion of its range. The Act
defines ‘‘endangered species’’ as any
species which is ‘‘in danger of
extinction throughout all or a significant
portion of its range,’’ and ‘‘threatened
species’’ as any species which is ‘‘likely
to become an endangered species within
the foreseeable future throughout all or
a significant portion of its range.’’ The
term ‘‘species’’ includes ‘‘any
subspecies of fish or wildlife or plants,
and any distinct population segment
[DPS] of any species of vertebrate fish or
wildlife which interbreeds when
mature.’’ On July 1, 2014, we published
a final policy interpreting the phrase
‘‘Significant Portion of its Range’’ (SPR)
(79 FR 37578). The final policy states
that (1) if a species is found to be an
endangered or a threatened species
throughout a significant portion of its
range, the entire species is listed as an
endangered or a threatened species,
respectively, and the Act’s protections
apply to all individuals of the species
wherever found; (2) a portion of the
range of a species is ‘‘significant’’ if the
species is not currently an endangered
or a threatened species throughout all of
its range, but the portion’s contribution
to the viability of the species is so
important that, without the members in
that portion, the species would be in
danger of extinction, or likely to become
so in the foreseeable future, throughout
all of its range; (3) the range of a species
is considered to be the general
geographical area within which that
species can be found at the time FWS
or NMFS makes any particular status
determination; and (4) if a vertebrate
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species is an endangered or a threatened
species throughout an SPR, and the
population in that significant portion is
a valid DPS, we will list the DPS rather
than the entire taxonomic species or
subspecies.
The SPR policy is applied to all status
determinations, including analyses for
the purposes of making listing,
delisting, and reclassification
determinations. The procedure for
analyzing whether any portion is an
SPR is similar, regardless of the type of
status determination we are making.
The first step in our analysis of the
status of a species is to determine its
status throughout all of its range. If we
determine that the species is in danger
of extinction, or likely to become so in
the foreseeable future, throughout all of
its range, we list the species as an
endangered (or threatened) species and
no SPR analysis will be required. If the
species is neither an endangered nor a
threatened species throughout all of its
range, we determine whether the
species is an endangered or a threatened
species throughout a significant portion
of its range. If it is, we list the species
as an endangered or a threatened
species, respectively; if it is not, we
conclude that listing the species is not
warranted.
When we conduct an SPR analysis,
we first identify any portions of the
species’ range that warrant further
consideration. The range of a species
can theoretically be divided into
portions in an infinite number of ways.
However, there is no purpose to
analyzing portions of the range that are
not reasonably likely to be significant
and either an endangered or a
threatened species. To identify only
those portions that warrant further
consideration, we determine whether
there is substantial information
indicating that (1) the portions may be
significant and (2) the species may be in
danger of extinction in those portions or
likely to become so within the
foreseeable future. We emphasize that
answering these questions in the
affirmative is not a determination that
the species is an endangered or a
threatened species throughout a
significant portion of its range—rather,
it is a step in determining whether a
more detailed analysis of the issue is
required. In practice, a key part of this
analysis is whether the threats are
geographically concentrated in some
way. If the threats to the species are
affecting it uniformly throughout its
range, no portion is likely to warrant
further consideration. Moreover, if any
concentration of threats apply only to
portions of the range that clearly do not
meet the biologically based definition of
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‘‘significant’’ (i.e., the loss of that
portion clearly would not be expected to
increase the vulnerability to extinction
of the entire species), those portions
will not warrant further consideration.
If we identify any portions that may
be both (1) significant and (2)
endangered or threatened, we engage in
a more detailed analysis to determine
whether these standards are indeed met.
The identification of an SPR does not
create a presumption, prejudgment, or
other determination as to whether the
species in that identified SPR is an
endangered or a threatened species. We
must go through a separate analysis to
determine whether the species is an
endangered or a threatened species in
the SPR. To determine whether a
species is an endangered or a threatened
species throughout an SPR, we will use
the same standards and methodology
that we use to determine if a species is
an endangered or a threatened species
throughout its range.
Depending on the biology of the
species, its range, and the threats it
faces, it may be more efficient to address
the ‘‘significant’’ question first, or the
status question first. Thus, if we
determine that a portion of the range is
not ‘‘significant,’’ we do not need to
determine whether the species is an
endangered or a threatened species
there; if we determine that the species
is not an endangered or a threatened
species in a portion of its range, we do
not need to determine if that portion is
‘‘significant.’’
We evaluated the current range of the
Upper Missouri River DPS of Arctic
grayling to determine if there is any
apparent geographic concentration of
potential threats. We examined
potential threats from curtailment of
range, dams, habitat fragmentation,
dewatering and thermal stress,
entrainment, riparian habitat loss,
sediment, exploitation, disease and
competition/predation, drought, climate
change, stochastic events, reduced
genetic diversity, low abundance, and
lack of a fluvial ecotype replicate. The
type and magnitude of stressors acting
on the Arctic grayling populations in
the DPS are varied.
Currently, nineteen of the twenty
Arctic grayling populations in the DPS
are stable or increasing in abundance.
Given this trend, we conclude that there
is no concentration of threats acting on
these nineteen populations because
these populations are able to maintain
viability, despite some stressors acting
at the individual level on some of these
populations. However, we acknowledge
the probable declining population trend
in the Ennis Reservoir/Madison River
population. It is unclear what factor or
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combination of factors is contributing to
this decline. Nonnative trout abundance
is highest in the Madison River, relative
to all other systems occupied by
nonnative trout and Arctic grayling, and
this factor may be contributing to the
decline of Arctic grayling in Ennis
Reservoir/Madison River.
Given the probable decline of Arctic
grayling in Ennis Reservoir/Madison
River, we analyzed the potential
significance of this population to the
overall Upper Missouri River DPS of
Arctic grayling. To do this analysis, we
evaluated whether the Ennis Reservoir/
Madison River population’s
contribution to the viability of the DPS
is so important that, without the
members in this portion, the DPS would
be in danger of extinction, or likely to
become so in the foreseeable future,
throughout all of its range. The Ennis
Reservoir/Madison River population
occupies a small portion of the range
within the DPS and represents only 1 of
20 populations in the overall DPS. We
conclude that the DPS would still be
viable if the Ennis Reservoir/Madison
River population were extirpated
because adequate redundancy (3 other
fluvial or partially fluvial and 16 other
adfluvial populations) of Arctic grayling
populations would still exist. In
addition, representation of resilient
populations would remain in the
Madison drainage (Grebe Lake
population) and rangewide in 7 of 10
historically occupied watersheds in the
Upper Missouri River basin. Further,
resiliency of the DPS would not be
compromised by the loss of the Ennis
Reservoir/Madison River population
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because all remaining Arctic grayling
populations are widespread and viable.
Therefore, in the hypothetical absence
of the Ennis Reservoir/Madison River
population, the remainder of the Upper
Missouri River DPS of Arctic grayling
would not meet the definition of
threatened or endangered under the Act.
For the reasons stated above, the Ennis
Reservoir/Madison River population
does not meet the definition of
‘‘significant’’ for the purposes of this
SPR analysis.
In conclusion, we find no
concentration of stressors acting on
nineteen of twenty Arctic grayling
populations in the DPS. The Ennis
Reservoir/Madison River population
does appear to have a stressor or
combination of stressors acting at the
population level. However, further
analysis indicates that the Ennis
Reservoir/Madison River does not meet
the definition of ‘‘significant’’ in our
SPR policy because adequate
redundancy, representation, and
resiliency would still exist within the
DPS if the Ennis Reservoir/Madison
River population were extirpated. Thus,
the remainder of the Upper Missouri
River DPS of Arctic grayling would not
meet the definition of threatened or
endangered. Therefore, we find that
there is not a significant portion of the
range of the Upper Missouri River DPS
of Arctic grayling that warrants listing.
Our review of the best available
scientific and commercial information
indicates that the Upper Missouri River
DPS of Arctic grayling is not in danger
of extinction (endangered), nor likely to
become endangered within the
foreseeable future (threatened),
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throughout all or a significant portion of
its range. Therefore, we find that listing
the Upper Missouri River DPS of Arctic
grayling as an endangered or threatened
species under the Act is not warranted
at this time.
We request that you submit any new
information concerning the status of, or
threats to, the Upper Missouri River
DPS of Arctic grayling to our Montana
Ecological Services Office (see
ADDRESSES) whenever it becomes
available. New information will help us
monitor the Upper Missouri River DPS
of Arctic grayling and encourage its
conservation. If an emergency situation
develops for the Upper Missouri River
DPS of Arctic grayling, we will act to
provide immediate protection.
References Cited
A complete list of references cited is
available on the Internet at https://
www.regulations.gov and upon request
from the Montana Ecological Services
Office (see ADDRESSES).
Authors
The primary authors of this document
are the staff members of the Montana
Ecological Services Office.
Authority
The authority for this section is
section 4 of the Endangered Species Act
of 1973, as amended (16 U.S.C. 1531 et
seq.).
Dated: August 6, 2014.
David Cottingham,
Acting Director, Fish and Wildlife Service.
[FR Doc. 2014–19353 Filed 8–19–14; 4:15 pm]
BILLING CODE 4310–55–P
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[Federal Register Volume 79, Number 161 (Wednesday, August 20, 2014)]
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[FR Doc No: 2014-19353]
[[Page 49383]]
Vol. 79
Wednesday,
No. 161
August 20, 2014
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; Revised 12-Month Finding
on a Petition To List the Upper Missouri River Distinct Population
Segment of Arctic Grayling as an Endangered or Threatened Species;
Proposed Rule
Federal Register / Vol. 79 , No. 161 / Wednesday, August 20, 2014 /
Proposed Rules
[[Page 49384]]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R6-ES-2013-0120; 4500030113]
Endangered and Threatened Wildlife and Plants; Revised 12-Month
Finding on a Petition To List the Upper Missouri River Distinct
Population Segment of Arctic Grayling as an Endangered or Threatened
Species
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of 12-month petition finding.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), announce a
revised 12-month finding on a petition to list the Upper Missouri River
distinct population segment (Upper Missouri River DPS) of Arctic
grayling (Thymallus arcticus) as an endangered or threatened species
under the Endangered Species Act of 1973, as amended (Act). After
review of the best available scientific and commercial information, we
find that listing the Upper Missouri River DPS of Arctic grayling is
not warranted at this time. The best available scientific and
commercial information indicates that habitat-related threats
previously identified, including habitat fragmentation, dewatering,
thermal stress, entrainment, riparian habitat loss, and effects from
climate change, for the Upper Missouri River DPS of Arctic grayling
have been sufficiently ameliorated and that 19 of 20 populations of
Arctic grayling are either stable or increasing. This action removes
the Upper Missouri River DPS of the Arctic grayling from our candidate
list. Although listing is not warranted at this time, we ask the public
to submit to us any new information that becomes available concerning
the threats to the Upper Missouri River DPS of Arctic grayling or its
habitat at any time.
DATES: The finding announced in this document was made on August 20,
2014.
ADDRESSES: This finding is available on the Internet at https://www.regulations.gov at Docket Number FWS-R6-ES-2013-0120. Supporting
documentation we used in preparing this finding is available for public
inspection, by appointment, during normal business hours at the U.S.
Fish and Wildlife Service, Montana Ecological Services Office, 585
Shepard Way, Suite 1, Helena, MT 59601. Please submit any new
information, materials, comments, or questions concerning this finding
to the above street address.
FOR FURTHER INFORMATION CONTACT: Jodi Bush, Field Supervisor, Montana
Ecological Services Office (see ADDRESSES); telephone 406-449-5225. If
you use a telecommunications device for the deaf (TDD), please call the
Federal Information Relay Service (FIRS) at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Background
Section 4(b)(3)(B) of the Act (16 U.S.C. 1531 et seq.) requires
that, for any petition to revise the Federal Lists of Endangered and
Threatened Wildlife and Plants that contains substantial scientific or
commercial information that listing the species may be warranted, we
make a finding within 12 months of the date of receipt of the petition.
In this finding, we will determine that the petitioned action is: (1)
Not warranted, (2) warranted, or (3) warranted, but the immediate
proposal of a regulation implementing the petitioned action is
precluded by other pending proposals to determine whether species are
endangered or threatened, and expeditious progress is being made to add
or remove qualified species from the Federal Lists of Endangered and
Threatened Wildlife and Plants. We must publish these 12-month findings
in the Federal Register.
Previous Federal Actions
We have published a number of documents on Arctic grayling since
1982, and have been involved in litigation over previous findings. We
describe previous federal actions that are relevant to this document
below.
We published our first status review for the Montana Arctic
grayling (Thymallus arcticus montanus), then thought to be a subspecies
of Arctic grayling, in a Federal Register document on December 30, 1982
(47 FR 58454). In that document, we designated the purported
subspecies, Montana Arctic grayling, as a Category 2 species. At that
time, we designated a species as Category 2 if a listing as endangered
or threatened was possibly appropriate, but we did not have sufficient
data to support a proposed rule to list the species.
On October 9, 1991, the Biodiversity Legal Foundation and George
Wuerthner petitioned us to list the fluvial (riverine) populations of
Arctic grayling in the Upper Missouri River basin as an endangered
species throughout its historical range in the coterminous United
States. We published a notice of a 90-day finding in the January 19,
1993, Federal Register (58 FR 4975), concluding the petitioners
presented substantial information indicating that listing the fluvial
Arctic grayling of the Upper Missouri River in Montana and northwestern
Wyoming may be warranted. This finding also noted that taxonomic
recognition of the Montana Arctic grayling (Thymallus arcticus
montanus) as a subspecies (previously designated as a category 2
species) was not widely accepted, and that the scientific community
generally considered this population a geographically isolated member
of the wider species (T. arcticus).
On July 25, 1994, we published notification of a 12-month finding
in the Federal Register (59 FR 37738), concluding that listing the DPS
of fluvial Arctic grayling in the Upper Missouri River was warranted
but precluded by other higher priority listing actions. This DPS
determination predated our DPS policy (61 FR 4722, February 7, 1996),
so the entity did not undergo a DPS analysis as described in the
policy. The 1994 finding placed fluvial Arctic grayling of the Upper
Missouri River on the candidate list and assigned it a listing priority
of 9, indicating that the threats were imminent but of moderate to low
magnitude.
On May 31, 2003, the Center for Biological Diversity and Western
Watersheds Project (Plaintiffs) filed a complaint in U.S. District
Court in Washington, DC, challenging our 1994 ``warranted but
precluded'' determination for the DPS of fluvial Arctic grayling in the
Upper Missouri River basin. On May 4, 2004, we elevated the listing
priority number of the fluvial Arctic grayling to 3 (69 FR 24881),
indicating threats that were imminent and of high magnitude. On July
22, 2004, the Plaintiffs amended their complaint to challenge our
failure to emergency list this population. We settled with the
Plaintiffs in August 2005, and we agreed to submit a revised
determination on whether this population warranted listing as
endangered or threatened to the Federal Register on or before April 16,
2007.
On April 24, 2007, we published a revised 12-month finding on the
petition to list the Upper Missouri River DPS of fluvial Arctic
grayling (72 FR 20305) (``2007 finding''). In this finding, we
determined that fluvial Arctic grayling of the upper Missouri River did
not constitute a species, subspecies, or DPS under the Act. Therefore,
we found that the upper Missouri River
[[Page 49385]]
population of fluvial Arctic grayling was not a listable entity under
the Act, and, as a result, listing was not warranted. With that
document, we withdrew the fluvial Arctic grayling from our candidate
list.
On November 15, 2007, the Center for Biological Diversity,
Federation of Fly Fishers, Western Watersheds Project, George
Wuerthner, and Pat Munday filed a complaint (CV-07-152, in the District
Court of Montana) to challenge our 2007 finding. We settled this
litigation on October 5, 2009. In the stipulated settlement, we agreed
to: (a) Publish, on or before December 31, 2009, a document in the
Federal Register soliciting information on the status of the upper
Missouri River Arctic grayling; and (b) submit, on or before August 30,
2010, a new 12-month finding for the upper Missouri River Arctic
grayling to the Federal Register.
On October 28, 2009, we published in the Federal Register a notice
of intent to conduct a status review of Arctic grayling (Thymallus
arcticus) in the upper Missouri River system (74 FR 55524). To ensure
the status review was based on the best available scientific and
commercial data, we requested information on the taxonomy, biology,
ecology, genetics, and population status of the Arctic grayling of the
upper Missouri River system; information relevant to consideration of
the potential DPS status of Arctic grayling of the upper Missouri River
system; threats to the species; and conservation actions being
implemented to reduce those threats in the upper Missouri River system.
That document further specified that the status review might consider
various DPS designations that include different life histories of
Arctic grayling in the upper Missouri River system and different DPS
configurations, including fluvial, adfluvial (lake populations), or all
life histories of Arctic grayling in the upper Missouri River system.
On September 8, 2010, we published a revised 12-month finding on
the petition to list the Upper Missouri River DPS of Arctic grayling
(75 FR 54708) (``2010 finding''). In this finding, we determined that
fluvial and adfluvial Arctic grayling of the upper Missouri River did
constitute a DPS under the Act. Further, we found that a DPS
configuration including both adfluvial and fluvial life histories was
the most appropriate for the long-term conservation of Arctic grayling
because genetic evidence indicated that fluvial and adfluvial life-
history forms did not represent distinct evolutionary lineages. We
concluded by finding that the Upper Missouri River DPS of Arctic
grayling was warranted for listing under the Act, but precluded by
other higher priority listing actions.
On September 9, 2011, we reached an agreement with plaintiffs in
Endangered Species Act Section 4 Deadline Litig., Misc. Action No. 10-
377 (EGS), MDL Docket No. 2165 (D. D.C.) (known as the ``MDL case'') on
a schedule to publish proposed listing rules or not-warranted findings
for the species on our candidate list. This agreement stipulated that
we would submit for publication in the Federal Register either a
proposed listing rule for the Upper Missouri River DPS of Arctic
grayling, or a not-warranted finding, no later than the end of Fiscal
Year 2014.
On November 26, 2013, we published a document in the Federal
Register (78 FR 70525) notifying the public that we were initiating a
status review of the Upper Missouri River DPS of Arctic grayling to
determine whether the entity meets the definition of an endangered or
threatened species under the Act. That document requested general
information (taxonomy, biology, ecology, genetics, and status) on the
Arctic grayling of the upper Missouri River system, as well as
information on the conservation status of, threats to, planned and
ongoing conservation actions for, habitat selection of, habitat
requirements of, and considerations concerning the possible designation
of critical habitat for the Arctic grayling of the upper Missouri River
system.
This document constitutes a revised 12-month finding (``2014
finding'') on whether to list the Upper Missouri River DPS of Arctic
grayling (Thymallus arcticus) as endangered or threatened under the
Act, and fulfills our commitments under the MDL case.
Species Information
Taxonomy and Species Description
The Arctic grayling (Thymallus arcticus) is a fish belonging to the
family Salmonidae (salmon, trout, charr, whitefishes), subfamily
Thymallinae (graylings), and it is represented by a single genus,
Thymallus. Arctic grayling have elongate, laterally compressed, trout-
like bodies with deeply forked tails, and adults typically average 300-
380 millimeters (mm) (12-15 inches (in.)) in length. Coloration can be
striking, and varies from silvery or iridescent blue and lavender, to
dark blue (Behnke 2002, pp. 327-328). A prominent morphological feature
of Arctic grayling is the sail-like dorsal fin, which is large and
vividly colored with rows of orange to bright green spots, and often
has an orange border (Behnke 2002, pp. 327-328).
For more detail on taxonomy and species description, see the 2010
finding (75 FR 54708).
Distribution
Arctic grayling are native to Arctic Ocean drainages of Alaska and
northwestern Canada, as far east as Hudson's Bay, and westward across
northern Eurasia to the Ural Mountains (Scott and Crossman 1998, pp.
301-302; Froufe et al. 2005, pp. 106-107; Weiss et al. 2006, pp. 511-
512). In North America, they are native to northern Pacific Ocean
drainages as far south as the Stikine River in British Columbia (Nelson
and Paetz 1991, pp. 253-256; Behnke 2002, pp. 327-331).
For a full discussion on the global distribution of Arctic
grayling, see the 2010 finding (75 FR 54709-54710). Here, we focus on
the distribution of Arctic grayling within the conterminous United
States.
Distribution in the Conterminous United States
Two disjunct groups of Arctic grayling were native to the
conterminous United States: One in the upper Missouri River basin in
Montana and Wyoming (currently extant only in Montana); and another in
Michigan that was extirpated in the late 1930s (Hubbs and Lagler 1949,
p. 44), and has not been detected since.
During the status review process, the Service received information
indicating that Arctic grayling may have also been native to areas
outside the Upper Missouri River basin in Montana and Wyoming. This
information included multiple historical newspaper clippings and
several reports from early Army expeditions purporting that Arctic
grayling were captured in the Yellowstone River drainage in Montana and
the Snake River drainage in Idaho (Shea 2014, entire). Some of these
reports even included descriptions of captured fish. However, none of
the descriptions mentions the colorful, sail-like dorsal fin of Arctic
grayling, a prominent feature that clearly distinguishes Arctic
grayling from other salmonids. In addition, a similar species
resembling Arctic grayling (i.e., mountain whitefish) is native to both
the Yellowstone River drainage and Snake River drainage. Mountain
whitefish were sometimes referred to as ``grayling'' in some areas of
the West (Ellis 1914, p. 75). Thus, it is likely that early reports of
Arctic grayling occurring outside the upper Missouri River basin were
mountain whitefish misidentified as Arctic grayling. Therefore, without
information to the contrary, we consider Arctic grayling to
[[Page 49386]]
be native only to the upper Missouri River basin in Montana and Wyoming
and to Michigan.
Native Distribution of Arctic Grayling in the Upper Missouri River
Basin
The first Euro-American ``discovery'' of Arctic grayling in North
America is attributed to members of the Lewis and Clark Expedition, who
encountered the species in the Beaverhead River in August 1805 (Nell
and Taylor 1996, p. 133). Vincent (1962, p. 11) and Kaya (1992, pp. 47-
51) synthesized accounts of Arctic grayling occurrence and abundance
from historical surveys and contemporary monitoring to determine the
historical distribution of the species in the upper Missouri River
system (Figure 1). We base our conclusions on the historical
distribution of Arctic grayling in the upper Missouri River basin on
these two reviews. Arctic grayling were widely but irregularly
distributed in the upper Missouri River system above the Great Falls in
Montana and in northwest Wyoming within the present-day location of
Yellowstone National Park (Vincent 1962, p. 11). They were estimated to
inhabit up to 2,000 kilometers (km) (1,250 miles (mi)) of stream
habitat until the early 20th century (Kaya 1992, pp. 47-51). Arctic
grayling were reported in the mainstem Missouri River, as well as in
the Smith, Sun, Jefferson, Madison, Gallatin, Big Hole, Beaverhead, and
Red Rock Rivers (Vincent 1962, p. 11; Kaya 1992, pp. 47-51; USFWS 2007;
72 FR 20307, April 24, 2007). Anecdotal accounts report that the
species may have been present in the Ruby River, at least seasonally
(Magee 2005, pers. comm.), and were observed there as recently as the
early 1970s (Holton, undated).
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Fluvial Arctic grayling were historically widely distributed in the
upper Missouri River basin, but a few adfluvial populations also were
native to the basin. For example, Arctic grayling are native to Red
Rock Lakes, in the Centennial Valley (Vincent 1962, pp. 112-121; Kaya
1992, p. 47). Vincent (1962, p. 120) stated that Red Rock Lakes were
the only natural lakes in the upper Missouri River basin accessible to
colonization by Arctic grayling, and concluded that Arctic grayling
there were the only native adfluvial population in the basin. However,
Arctic grayling were also native to Elk Lake (in the Centennial Valley;
Kaya 1990, p. 44) and a few small lakes in the upper Big Hole River
drainage, based on recent genetic information (Peterson and Ardren
2009, p. 1768).
The distribution of native Arctic grayling in the upper Missouri
River went through a dramatic reduction in the first 50 years of the
20th century, especially in riverine habitats (Vincent 1962, pp. 86-90,
97-122, 127-129; Kaya 1992, pp. 47-53). The native populations that
formerly resided in the
[[Page 49388]]
Smith, Sun, Jefferson, Beaverhead, Gallatin, and mainstem Missouri
Rivers are considered extirpated, and the only remaining native fluvial
population is found in the Big Hole River and some of its tributaries
(Kaya 1992, pp. 51-53). The fluvial form currently occupies less than
10 percent of its historical range in the Missouri River system (Kaya
1992, p. 51). Other native populations in the upper Missouri River
occur in two small, headwater lakes in the upper Big Hole River system
(Miner and Mussigbrod Lakes); the upper Ruby River (recently
reintroduced from Big Hole River stock); the Madison River upstream
from Ennis Reservoir; Elk Lake in the Centennial Valley (recently
reintroduced from Red Rock Lakes stock); and the Red Rock Lakes in the
Centennial Valley (Everett 1986, p. 7; Kaya 1992, p. 53; Peterson and
Ardren 2009, pp. 1762, 1768; see Figure 1).
Introduced Lake-Dwelling Arctic Grayling in the Upper Missouri River
Basin
From 1898 through the 1960s, an estimated 100 million Arctic
grayling were stocked across Montana and other western States. The
sources of these stockings varied through time as different State,
Federal, and private hatchery operations were created, but the ultimate
source for all hatcheries in Montana appears to be stock from two
Montana populations: Centennial Valley and Madison River (Peterson and
Ardren 2009, p. 1767; Leary 2014, unpublished data; MFISH 2014a).
Arctic grayling derived from these two sources were stocked on top of
every known native Arctic grayling population in the upper Missouri
River basin. In addition, Arctic grayling were stocked in multiple high
elevation lakes, some of which likely were historically fishless.
There are 20 known, introduced Arctic grayling populations that
exist in the upper Missouri River basin. These 20 populations, along
with the 6 populations existing in native habitat, comprise the
listable entity (total of 26 populations) of Arctic grayling in the
upper Missouri River basin. However, six of these introduced
populations are considered to have low conservation value because they
occupy unnatural habitat, are not self-sustaining, or are used as
captive brood reserves. These six populations are Axolotl Lake, Green
Hollow Lake, Sunnyslope Canal, Tunnel Lake, South Fork Sun River, and
Elk Lake. The Axolotl and Green Hollow populations are captive brood
reserves maintained in natural lakes for reintroduction purposes.
Sunnyslope Canal is a fluvial population that occurs in unnatural
habitat (irrigation canal). Tunnel Lake is stocked with ``rescued''
fish from Sunnyslope Canal, but lacks a spawning tributary and is
consequently not self-sustaining (SSA 2014). South Fork Sun River is a
small fluvial population that resides in about \1/4\ mile of stream
during the summer and is not considered self-sustaining (SSA 2014). The
Elk Lake population is a genetic replicate of the Centennial Valley
population, but no documented spawning has occurred to date (Jaeger
2014a, pers. comm.); thus this population is not currently considered
self-sustaining. For these reasons, we primarily focus our analysis on
the populations considered to have high conservation value; those
populations that are self-sustaining, in natural habitats, and wild.
The 14 known remaining introduced, lake-dwelling (adfluvial) Arctic
grayling populations within the upper Missouri River basin are likely
the result of historical stocking (Table 1). In our 2010 finding, we
considered and discussed the conservation value of these populations.
Based on the information available at that time, we considered these
introduced populations to not have conservation value for multiple
reasons. Below, we list each of the reasons for this conclusion as
provided in the 2010 finding, and provide an updated assessment and
conclusion about the potential conservation value of these populations,
based on new information obtained since 2010.
Table 1--Geographic Distribution, Genetic Status, and Source of Introduced Adfluvial Arctic Grayling Populations
in the Upper Missouri River Basin
----------------------------------------------------------------------------------------------------------------
Genetic
Population Drainage analysis Source \a\ Citation
completed?
----------------------------------------------------------------------------------------------------------------
Agnes Lake................... Big Hole....... No............. Madison/ MFISH 2014a.
Centennial.
Odell Lake................... Big Hole....... Yes............ Centennial..... Peterson and Ardren 2009, p.
1766; Leary 2014, unpublished
data.
Bobcat Lake.................. Big Hole....... Yes............ Centennial..... Peterson and Ardren 2009, p.
1766; Leary 2014, unpublished
data.
Schwinegar Lake.............. Big Hole....... No............. Madison/ ..............................
Centennial.\c\.
Pintlar Lake................. Big Hole....... Yes............ Madison/ Leary 2014, unpublished data.
Centennial.
Deer Lake.................... Gallatin....... Yes............ Madison/ Leary 2014, unpublished data.
Centennial.
Emerald Lake................. Gallatin....... Yes............ Madison/ Leary 2014, unpublished data.
Centennial.
Grayling Lake................ Gallatin....... Yes............ Madison/ Leary 2014, unpublished data.
Centennial.
Hyalite Lake................. Gallatin....... Yes............ Madison/ Leary 2014, unpublished data.
Centennial.
Diversion Lake............... Sun............ Yes \b\........ Big Hole....... Horton 2014a, pers. comm.;
Magee 2014, pers. comm.
Gibson Reservoir............. Sun............ Yes \b\........ Big Hole....... Horton 2014a, pers. comm.;
Magee 2014, pers. comm.
Lake Levale.................. Sun............ Yes \b\........ Big Hole....... Horton 2014a, pers. comm.;
Magee 2014, pers. comm.
Park Lake.................... Missouri....... No............. Madison/ ..............................
Centennial.\c\.
Grebe Lake................... Madison........ Yes............ Centennial..... Peterson and Ardren 2009, p.
1766; Varley 1981, p. 11.
----------------------------------------------------------------------------------------------------------------
\a\ Origin of source stock was determined by genetic analysis and through analysis of historical stocking
records and scientific literature, in some cases. Where multiple sources are cited, fish from each population
were known to be stocked, although the genetic contribution of each donor population to the current population
structure is unknown.
\b\ These populations are the result of reintroductions using known sources of Montana origin.
\c\ Schwinegar and Park Lakes Arctic grayling populations are likely from Montana-origin sources due to
proximity to other lakes with known Montana origin; however, definitive evidence is lacking.
1. The Service interprets the Act to provide a statutory directive
to conserve species in their native ecosystems (49 FR 33885, August 27,
1984) and to conserve genetic resources and biodiversity over a
representative portion of a taxon's historical occurrence (61 FR 4722,
February 7, 1996). Since most of the introduced populations of Arctic
grayling were of unknown genetic origin and in lakes that were likely
historically fishless,
[[Page 49389]]
these populations were considered in 2010 to be outside the species'
native range, and we concluded that they did not appear to add
conservation value to the DPS.
Since 2010, new genetic information from 7 of the 14 introduced
populations indicates there are moderate to high levels of genetic
diversity within and among these populations, and indicates these
populations were derived from native sources within the upper Missouri
River basin (Leary 2014, unpublished data; Table 1). In addition,
stocking records show common stocking sources for introduced
populations that were genotyped (as described previously) and the two
populations that were not genotyped (the remaining 3 populations were
reintroductions of known Montana origin sources; Table 1). Thus, it
appears that all 14 introduced Arctic grayling populations contain
moderate to high levels of genetic diversity of Arctic grayling in the
upper Missouri River basin that was not captured within the DPS
designation in the 2010 finding.
The Service's current interpretation of the Act is consistent with
that in the 2010 finding; we believe it is important to conserve
species in their native ecosystems and to conserve genetic resources
and biodiversity over a representative portion of a taxon's historical
occurrence. In light of the new genetics information gained since 2010
(Leary 2014, unpublished data), we also believe it is important to
acknowledge the moderate to high levels of genetic diversity within the
introduced populations in the upper Missouri River basin and the
potential adaptive capabilities represented by this diversity. All
Arctic grayling populations (introduced or not) currently within the
upper Missouri River basin are derived from a common ancestor and have
a distinct evolutionary trajectory relative to the historical founding
populations in Canada and Alaska. Thus, Arctic grayling originating
from and currently within the upper Missouri River basin represent the
southernmost assemblage of the species, facing similar selection
pressures and evolving independent of more northern populations.
The introduced Arctic grayling populations in the upper Missouri
River basin occupy, for the most part, high-elevation habitats that are
high-quality because of intact riparian areas and a consistent supply
of cool water. Given the predicted effects of climate change in the
West (see discussion under ``Climate Change'' in Factor A below), these
types of habitats are the same habitats that the Service would explore
for long-term conservation of Arctic grayling, if needed, because they
may serve as thermal refugia as temperatures rise and provide greater
redundancy in case of catastrophic events.
2. In 2010, the Service concluded there did not appear to be any
formally recognized conservation value for the introduced populations
of Arctic grayling in the upper Missouri River basin because they were
not being used in conservation or restoration programs. This conclusion
was based on an interpretation of a National Marine Fisheries Service
final policy on the consideration of hatchery-origin fish in Endangered
Species Act listing determinations for Pacific salmon and steelhead
(anadromous Oncorhynchus spp.) (NMFS 2005, entire).
Until recently, the genetic structure and source of these
introduced populations were unknown. Populations with a high likelihood
of being Montana origin were used for conservation purposes (e.g.,
reintroductions) as a precautionary approach to Arctic grayling
conservation. Now that the amount of genetic diversity within and among
the introduced Arctic grayling populations and their source(s) are
known, it is probable these introduced populations could be used in
future conservation actions as source stock, if needed.
3. In 2010, the Service indicated there were concerns that
introduced, lake-dwelling Arctic grayling populations could pose
genetic risks to the native fluvial population (i.e., Big Hole
Population) as cited in the Montana Fluvial Arctic Grayling Restoration
Plan (``Restoration Plan,'' 1995, p. 15). In the Restoration Plan,
Arctic grayling populations in Agnes, Schwinegar, Odell, Miner and
Mussigbrod lakes were identified as potential threats to the genetic
integrity of the Big Hole River population because of hydrologic
connectivity between these lakes and the Big Hole River and the
potential for genetic mixing.
Recently, genetic analyses have confirmed reproductive isolation
among extant Arctic grayling populations in the upper Missouri River
basin and within the Big Hole River watershed (Peterson and Ardren
2009, p. 1770; Leary 2014, unpublished data). In addition, multiple
historical stockings have occurred in the Big Hole River from other
sources within the upper Missouri River basin. Recent genetic analysis
found no evidence of a significant genetic contribution from historical
stocking on the current genetic structure of Arctic grayling in the Big
Hole River (Peterson and Ardren 2009, p. 1768). Thus, we now conclude
that the concern that lake-dwelling populations within the Big Hole
River watershed could pose genetic risks to the Big Hole River fluvial
population appears unfounded.
4. In 2010, the Service concluded that introduced populations of
Arctic grayling in the upper Missouri River basin had no conservation
value because these populations apparently had been isolated from their
original source stock for decades without any supplementation from the
wild and were established without any formal genetic consideration to
selecting and mating broodstock.
It is now apparent from our review of historical stocking records
that many of these introduced populations received multiple stockings
from the same source or multiple stockings from several different
sources over a wide range of years (MFISH 2014a, unpublished data).
Additionally, most individual stockings involved a large number of eggs
or fry (up to 1 million for some stockings). Cumulatively, this
information suggests several points. First, stockings that used a large
number of eggs or fry necessitate that gametes from multiple brood fish
were used per stocking, given the physical constraints of number of
eggs per unit body size of female Arctic grayling. Second, stockings in
most of the introduced populations occurred over many years (up to 60
years in some cases). This indicates different cohorts of Arctic
grayling had to be used, since the generation time of Arctic grayling
is approximately 3.5 years in the upper Missouri River basin
(references in Dehaan et al. 2014, p. 10). Lastly, the new genetic
analyses from seven of the introduced Arctic grayling populations
indicate moderate to high levels of genetic diversity within the
populations. This result could likely only be obtained from the
founding of these populations using large numbers of brood fish and
gametes over multiple years. Mutation is unlikely to have accounted for
these levels of genetic diversity over a relatively short time period
of isolation (Freeman and Herron 2001, p. 143).
For perspective, Montana Fish, Wildlife, and Parks has developed
guidelines for the establishment and maintenance of Arctic grayling
broodstock. To adequately capture most of the genetic variation in a
source population, the crossing of a minimum of 25 male and 25 female
Arctic grayling is currently recommended (Leary 1991, p. 2151). It is
likely that the historical stockings used to found the introduced
Arctic grayling populations in the upper Missouri River basin equaled
or exceeded this through stocking large
[[Page 49390]]
numbers of eggs or fry over multiple years.
5. In 2010, the Service concluded that the source populations used
to found the introduced Arctic grayling populations in the upper
Missouri River drainage were not well documented (Peterson and Ardren
2009, p. 1767), so we could not be certain of whether these Arctic
grayling were of local origin.
Since 2010, new genetic information (Leary 2014, unpublished data)
and review of historical stocking records (MFISH 2014a, unpublished
data) indicate the founding populations used for stocking are local and
believed representative of the Upper Missouri River DPS of Arctic
grayling, and contain moderate to high levels of genetic diversity.
6. In 2010, the Service concluded the primary intent of culturing
and introducing Arctic grayling populations within the upper Missouri
River basin was to provide recreational fishing opportunities in high
mountain lakes, and that, therefore, these introduced populations had
no conservation value.
Since 2010, review of the historical literature indicates adfluvial
Arctic grayling populations were presumably stocked both for
recreational fishing and conservation purposes (Brown 1943, pp. 26-27;
Nelson 1954, p. 341; Vincent 1962, p. 151). Following the drought in
the 1930s, conservation stockings of Arctic grayling were advocated
because most rivers and streams were dewatered, prompting fish managers
to introduce Arctic grayling into habitats with a more consistent
supply of cool water (e.g., high-elevation mountain lakes; Brown 1943,
pp. 26-27; Nelson 1954, p. 341; Vincent 1962, p. 151).
In conclusion, introduced populations of Arctic grayling
established within the upper Missouri River basin, whether they were
originally established for recreational fishing or conservation
purposes, captured moderate to high levels of genetic diversity of
upper Missouri River basin Arctic grayling. The potential adaptive
capabilities represented by this genetic diversity have conservation
value, particularly in a changing climate. These populations reside in
high-quality habitat, the same habitat the Service would look to for
long-term conservation, if needed. Thus, the introduced populations of
Arctic grayling within the upper Missouri River basin have conservation
value, and, therefore, we include them in our analysis of a potential
DPS of Arctic grayling.
Origins, Biogeography, and Genetics of Arctic Grayling in North America
North American Arctic grayling are most likely descended from
Eurasian Thymallus that crossed the Bering land bridge during or before
the Pleistocene glacial period (Stamford and Taylor 2004, pp. 1533,
1546). There were multiple opportunities for freshwater faunal exchange
between North America and Asia during the Pleistocene, but genetic
divergence between North American and Eurasian Arctic grayling suggests
that the species could have colonized North America as early as the
mid-late Pliocene (more than 3 million years ago) (Stamford and Taylor
2004, p. 1546). Genetic studies of Arctic grayling using mitochondrial
DNA (mtDNA, maternally inherited DNA located in cellular organelles
called mitochondria) and microsatellite DNA (repeating sequences of
nuclear DNA) have shown that North American Arctic grayling consist of
at least three major lineages that originated in distinct Pleistocene
glacial refugia (Stamford and Taylor 2004, p. 1533). These three groups
include a South Beringia lineage found in western Alaska to northern
British Columbia, Canada; a North Beringia lineage found on the North
Slope of Alaska, the lower Mackenzie River, and to eastern
Saskatchewan; and a Nahanni lineage found in the lower Liard River and
the upper Mackenzie River drainage in northeastern British Columbia and
southeastern Yukon (Stamford and Taylor 2004, pp. 1533, 1540). Arctic
grayling from the upper Missouri River basin were tentatively placed in
the North Beringia lineage because a small sample (three individuals)
of Montana Arctic grayling shared a mtDNA haplotype (form of the mtDNA)
with populations in Saskatchewan and the lower Peace River, British
Columbia (Stamford and Taylor 2004, p. 1538).
The existing mtDNA data suggest that Missouri River Arctic grayling
share a common ancestry with the North Beringia lineage, but other
genetic markers (e.g., allozymes, microsatellites) and biogeographic
history indicate that Missouri River Arctic grayling have been
physically and reproductively isolated from northern populations for
millennia. Pre-glacial colonization of the Missouri River basin by
Arctic grayling was possible because the river flowed to the north and
drained into the Arctic-Hudson Bay prior to the last glacial cycle
(Cross et al. 1986, pp. 374-375; Pielou 1991, pp. 194-195). Low mtDNA
diversity observed in a small number of Montana Arctic grayling samples
and a shared ancestry with Arctic grayling from the North Beringia
lineage suggest a more recent, post-glacial colonization of the upper
Missouri River basin. In contrast, microsatellite DNA show substantial
divergence between Montana and Saskatchewan (i.e., same putative mtDNA
lineage) (Peterson and Ardren 2009, entire). Differences in the
frequency and size distribution of microsatellite alleles between
Montana populations and two Saskatchewan populations indicate that
Montana Arctic grayling have been isolated long enough for mutations
(i.e., evolution) to be responsible for the observed genetic
differences.
Additional comparison of 21 Arctic grayling populations from
Alaska, Canada, and the Missouri River basin using 9 of the same
microsatellite loci as Peterson and Ardren (2009, entire) further
supports the distinction of Missouri River Arctic grayling relative to
populations elsewhere in North America (USFWS, unpublished data).
Analyses of these data using two different methods clearly separates
sample fish from 21 populations into two clusters: One cluster
representing populations from the upper Missouri River basin, and
another cluster representing populations from Canada and Alaska (USFWS,
unpublished data). These new data, although not yet peer reviewed,
support the interpretation that the previous analyses of Stamford and
Taylor (2004, entire) underestimated the distinctiveness of Missouri
River Arctic grayling relative to other sample populations, likely
because of the combined effect of small sample sizes and the lack of
variation observed in the Missouri River for the markers used in that
study (Stamford and Taylor 2004, pp. 1537-1538). Thus, these recent
microsatellite DNA data suggest that Arctic grayling may have colonized
the Missouri River before the onset of Wisconsin glaciation (more than
80,000 years ago).
Genetic relationships among native and introduced populations of
Arctic grayling in Montana have recently been investigated (Peterson
and Ardren 2009, entire). Introduced, lake-dwelling populations of
Arctic grayling trace some of their original ancestry to the Centennial
Valley (Peterson and Ardren 2009, p. 1767), and stocking of hatchery
Arctic grayling did not have a large effect on the genetic composition
of the extant native populations (Peterson and Ardren 2009, p. 1768).
Differences between native populations of the two Arctic grayling
ecotypes (adfluvial, fluvial) are not as large as differences resulting
from geography (i.e., drainage of origin). For example, native
adfluvial Arctic grayling populations from
[[Page 49391]]
different lakes are genetically different (Peterson and Ardren 2009, p.
1766).
Habitat
Arctic grayling generally require clear, cold water, and are
characterized as belonging to a ``coldwater'' group of salmonids, which
also includes bull trout (Salvelinus confluentus) and Arctic char
(Salvelinus alpinus) (Selong et al. 2001, p. 1032). Arctic grayling
optimal thermal habitat is between 7 to 17 [deg]C (45 to 63 [deg]F),
but becomes unsuitable above 20 [deg]C (68 [deg]F) (Hubert et al. 1985,
p. 24). Arctic grayling fry may be more tolerant of high water
temperature than adults (LaPerriere and Carlson 1973, p. 30; Feldmeth
and Eriksen 1978, p. 2041).
Having a broad, nearly circumpolar distribution, Arctic grayling
occupy a variety of habitats including small streams, large rivers,
lakes, and even bogs (Northcote 1995, pp. 152-153; Scott and Crossman
1998, p. 303). They may even enter brackish water (less than or equal
to 4 parts per thousand salt content) when migrating between adjacent
river systems (West et al. 1992, pp. 713-714). Native populations are
found at elevations ranging from near sea level, such as in Bristol
Bay, Alaska, to high-elevation montane valleys (more than 1,830 meters
(m) or 6,000 feet (ft)), such as the Big Hole River and Centennial
Valley in southwestern Montana. Despite this broad distribution, Arctic
grayling have specific habitat requirements that can constrain their
local distributions, especially water temperature and channel gradient.
At the local scale, Arctic grayling prefer cold water and are often
associated with spring-fed habitats in regions with warmer climates
(Vincent 1962, p. 33). Arctic grayling are generally not found in
swift, high-gradient streams, and Vincent (1962, pp. 36-37, 41-43)
characterized typical Arctic grayling habitat in Montana (and Michigan)
as low-to-moderate gradient (less than 4 percent) streams and rivers
with low-to-moderate water velocities (less than 2 feet/sec (60
centimeters/sec)). Juvenile and adult Arctic grayling in streams and
rivers spend much of their time in pool habitat (Kaya 1990 and
references therein, p. 20; Lamothe and Magee 2003, pp. 13-14).
Breeding
Arctic grayling typically spawn in the spring or early summer,
depending on latitude and elevation (Northcote 1995, p. 149). In
Montana, Arctic grayling generally spawn from late April to mid-May by
depositing adhesive eggs over gravel substrate without excavating a
nest (Kaya 1990, p. 13; Northcote 1995, p. 151). In general, the
reproductive ecology of Arctic grayling differs from other salmonid
species (trout and salmon) in that Arctic grayling eggs tend to be
comparatively small; thus, they have higher relative fecundity (females
have more eggs per unit body size). Males establish and defend spawning
territories rather than defending access to females (Northcote 1995,
pp. 146, 150-151). The time required for development of eggs from
embryo until they emerge from stream gravel and become swim-up fry
depends on water temperature (Northcote 1995, p. 151). In the upper
Missouri River basin, development from embryo to fry averages about 3
weeks (Kaya 1990, pp. 16-17). Small, weakly swimming fry (typically 1-
1.5 centimeters (cm) (0.4-0.6 in.) at emergence) prefer low-velocity
stream habitats (Armstrong 1986, p. 6; Kaya 1990, pp. 23-24; Northcote
1995, p. 151).
Arctic grayling of all ages feed primarily on aquatic and
terrestrial invertebrates captured on or near the water surface, but
also will feed opportunistically on fish and fish eggs (Northcote 1995,
pp. 153-154; Behnke 2002, p. 328). Feeding locations for individual
fish are typically established and maintained through size-mediated
dominance hierarchies where larger individuals defend favorable feeding
positions (Hughes 1992, p. 1996).
General Life History Diversity
Migratory behavior is a common life-history trait in salmonid
fishes such as Arctic grayling (Armstrong 1986, pp. 7-8; Northcote
1995, pp. 156-158; 1997, pp. 1029, 1031-1032, 1034). In general,
migratory behavior in Arctic grayling and other salmonids results in
cyclic patterns of movement between refuge, rearing-feeding, and
spawning habitats (Northcote 1997, p. 1029).
Arctic grayling may move to refuge habitat as part of a regular
seasonal migration (e.g., in winter), or in response to episodic
environmental stressors (e.g., high summer water temperatures). In
Alaska, Arctic grayling in rivers typically migrate downstream in the
fall, moving into larger streams or mainstem rivers that do not
completely freeze (Armstrong 1986, p. 7). In Arctic rivers, fish often
seek overwintering habitat influenced by groundwater (Armstrong 1986,
p. 7). In some drainages, individual fish may migrate considerable
distances (greater than 150 km or 90 mi) to overwintering habitats
(Armstrong 1986, p. 7). In the Big Hole River, Montana, similar
downstream and long-distance movement to overwintering habitat has been
observed in Arctic grayling (Shepard and Oswald 1989, pp. 18-21, 27).
In addition, Arctic grayling in the Big Hole River may move downstream
in proximity to colder tributary streams in summer when thermal
conditions in the mainstem river become stressful (Lamothe and Magee
2003, p. 17).
In spring, mature Arctic grayling leave overwintering areas and
migrate to suitable spawning sites. In river systems, this typically
involves an upstream migration to tributary streams or shallow riffles
within the mainstem (Armstrong 1986, p. 8; Shepard and Oswald 1989; p.
18). Arctic grayling in lakes typically migrate to either the inlet or
outlet to spawn (Armstrong 1986, p. 8; Kaya 1989, p. 474; Northcote
1995 p. 148). In some situations, Arctic grayling exhibit natal homing,
whereby individuals spawn in or near the location where they were born
(Northcote 1995 pp. 157-160; Boltz and Kaeding 2002, p. 22); however,
it is unclear what factors may be influencing the extent of this
phenomenon.
Fry from river populations typically seek feeding and rearing
habitats in the vicinity of where they were spawned (Armstrong 1986,
pp. 6-7; Kaya and Jeanes 1995, p. 455; Northcote 1995, p. 156), while
those from lake populations migrate downstream (inlet spawners) or
upstream (outlet spawners) to the adjacent lake. Following spawning,
adults move to appropriate feeding areas if they are not adjacent to
spawning habitat (Armstrong 1986, pp. 7-8; Shepard and Oswald 1989; p.
18). Juvenile Arctic grayling may undertake seasonal migrations between
feeding and overwintering habitats until they reach maturity and add
the spawning migration to this cycle (Northcote 1995, pp. 156-157).
Life History Diversity in Arctic Grayling in the Upper Missouri River
Basin
Two general life-history forms or ecotypes of native Arctic
grayling occur in the upper Missouri River Arctic: Fluvial and
adfluvial. Fluvial fish use river or stream (lotic) habitat for all of
their life cycles and may undergo extensive migrations within river
habitat, up to 50 miles in the Big Hole River in Montana (Shepard and
Oswald 1989, p. 18). Adfluvial fish live in lakes and migrate to
tributary streams to spawn. These same life-history forms also are
expressed by Arctic grayling elsewhere in North America (Northcote
1997, p. 1030). Historically, the fluvial life-history form
predominated in the Missouri River basin above the Great Falls, perhaps
because there were only a few lakes accessible to natural colonization
of Arctic grayling that would permit expression of the
[[Page 49392]]
adfluvial ecotype (Kaya 1992, p. 47). The fluvial and adfluvial life-
history forms of Arctic grayling in the upper Missouri River do not
appear to represent distinct evolutionary lineages. Instead, they
appear to represent an example of adaptive radiation (Schluter 2000, p.
1), whereby the forms differentiated from a common ancestor and
developed traits that allowed them to exploit different habitats. The
primary evidence for this conclusion is genetic data that indicate that
within the Missouri River basin the two ecotypes are more closely
related to each other than they are to the same ecotype elsewhere in
North America (Redenbach and Taylor 1999, pp. 27-28; Stamford and
Taylor 2004, p. 1538; Peterson and Ardren 2009, p. 1766). Historically,
there may have been some genetic exchange between the two life-history
forms as individuals strayed or dispersed into different populations
(Peterson and Ardren 2009, p. 1770), but the genetic structure of
current populations in the upper Missouri River basin is consistent
with reproductive isolation.
The fluvial and adfluvial forms of Arctic grayling appear to differ
in their genetic characteristics, but there appears to be some
plasticity in behavior where individuals from a population can exhibit
a range of behaviors. Arctic grayling fry in Montana can exhibit
heritable, genetically-based differences in swimming behavior between
fluvial and adfluvial ecotypes (Kaya 1991, pp. 53, 56-58; Kaya and
Jeanes 1995, pp. 454, 456). Progeny of Arctic grayling from the fluvial
ecotype exhibited a greater tendency to hold their position in flowing
water relative to progeny from adfluvial ecotypes (Kaya 1991, pp. 53,
56-58; Kaya and Jeanes 1995, pp. 454, 456). Similarly, young Arctic
grayling from inlet and outlet spawning adfluvial ecotypes exhibited an
innate tendency to move downstream and upstream, respectively (Kaya
1989, pp. 478-480). All three studies (Kaya 1989, entire; 1991, entire;
Kaya and Jeanes 1995, entire) demonstrate that the response of fry to
flowing water depended strongly on the life-history form (ecotype) of
the source population, and that this behavior has a genetic basis.
However, behavioral responses also were mediated by environmental
conditions (light--Kaya 1991, pp. 56-57; light and water temperature--
Kaya 1989, pp. 477-479), and some progeny of each ecotype exhibited
behavior characteristic of the other; for example some individuals from
the fluvial ecotype moved downstream rather than holding position, and
some individuals from an inlet-spawning adfluvial ecotype held position
or moved upstream (Kaya 1991, p. 58). These observations indicate that
some plasticity for behavior exists, at least for very young Arctic
grayling.
The ability of the fluvial ecotype to give rise to a functional
population of the adfluvial ecotype has been demonstrated. Most extant
adfluvial Arctic grayling populations in the Upper Missouri River
originated from fluvial-dominated sources (see Table 1; Kaya 1992, p.
53; Jeanes 1996, pp. 54). However, the ability of the adfluvial ecotype
to give rise to a functional population of fluvial ecotype is less
certain. Circumstantial support for reduced plasticity in adfluvial
Arctic grayling comes from observations that adfluvial fish stocked in
river habitats almost never establish populations (Kaya 1990, pp. 31-
34). However, we note that adfluvial Arctic grayling retain some life-
history flexibility--at least in lake environments--as naturalized
populations derived from inlet-spawning stocks have established outlet-
spawning demes (a deme is a local populations that shares a distinct
gene pool) in Montana and in Yellowstone National Park (Kruse 1959, p.
318; Kaya 1989, p. 480). In addition, a small percentage of young
adfluvial Arctic grayling exposed to flow exhibited fluvial-like
characteristics (e.g., station-holding or upstream movement) in a
laboratory experiment designed to assess movement tendencies of
adfluvial and fluvial Arctic grayling in flowing water (Kaya 1991, p.
56). These results indicate some plasticity exists in adfluvial Arctic
grayling that may allow some progeny of adfluvial individuals to
express a fluvial life history. Nonetheless, the frequent failure of
introductions of adfluvial Arctic grayling into fluvial habitats
suggest a cautionary approach to the loss of particular life-history
forms is warranted.
Age and Growth
Age at maturity and longevity in Arctic grayling varies regionally
and is probably related to growth rate, with populations in colder,
northern latitudes maturing at later ages and having a greater lifespan
(Kruse 1959, pp. 340-341; Northcote 1995 and references therein, pp.
155-157). Arctic grayling in the upper Missouri River typically mature
at age 2 (males) or age 3 (females), and individuals greater than age 6
are rare (Kaya 1990, p. 18; Magee and Lamothe 2003, pp. 16-17). The
majority of the Arctic grayling spawning in two tributaries in the
Centennial Valley, Montana, were age 3, and the oldest individuals aged
from a larger sample were age 6 (Nelson 1954, pp. 333-334). Arctic
grayling spawning in Red Rock Creek were mostly ages 2 to 5, but some
individuals were age 7 (Mogen 1996, pp. 32-34).
Generally, growth rates of Arctic grayling are greatest during the
first years of life then slow dramatically after maturity. Within that
general pattern, there is substantial variation among populations from
different regions. Arctic grayling populations in Montana (Big Hole
River and Red Rock Lakes) have very high growth rates relative to those
from British Columbia, Asia, and the interior and North Slope of Alaska
(Carl et al. 1992, p. 240; Northcote 1995, pp. 155-157; Neyme 2005, p.
28).
Distinct Vertebrate Population Segment
Under the Service's Policy Regarding the Recognition of Distinct
Vertebrate Population Segments Under the Endangered Species Act (61 FR
4722; February 7, 1996), three elements are considered in the decision
concerning the establishment and classification of a possible DPS.
These are applied similarly for additions to or removal from the
Federal List of Endangered and Threatened Wildlife. These elements
include:
(1) The discreteness of a population in relation to the remainder
of the species to which it belongs;
(2) The significance of the population segment to the species to
which it belongs; and
(3) The population segment's conservation status in relation to the
Act's standards for listing, delisting, or reclassification (i.e., is
the population segment endangered or threatened).
Discreteness
Under the DPS policy, a population segment of a vertebrate taxon
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.
(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.
Arctic grayling native to the upper Missouri River are isolated
from all other populations of the species, which inhabit the Arctic
Ocean, Hudson Bay, and north Pacific Ocean drainages in
[[Page 49393]]
Asia and North America. Arctic grayling native to the upper Missouri
River occur as a disjunct group of populations approximately 800 km
(500 mi) to the south of the next-nearest Arctic grayling population in
central Alberta, Canada. Missouri River Arctic grayling have been
isolated from other populations for at least 10,000 years based on
historical reconstruction of river flows at or near the end of the
Pleistocene (Cross et al. 1986, p. 375; Pileou 1991, pp. 10-11).
Genetic data confirm Arctic grayling in the Missouri River basin have
been reproductively isolated from populations to the north for
millennia (Everett 1986, pp. 79-80; Redenbach and Taylor 1999, p. 23;
Stamford and Taylor 2004, p. 1538; Peterson and Ardren 2009, pp. 1764-
1766; USFWS, unpublished data). Consequently, we conclude that Arctic
grayling native to the upper Missouri River are markedly separated from
other native populations of the taxon as a result of physical factors
(isolation), and therefore meet the first criterion of discreteness
under the DPS policy. As a result, Arctic grayling native to the upper
Missouri River are considered a discrete population according to the
DPS policy. Because the entity meets the first criterion (markedly
separated), an evaluation with respect to the second criterion
(international boundaries) is not needed.
Significance
If a population segment is considered discrete under one or more of
the conditions described in the Service's DPS policy, its biological
and ecological significance will be considered in light of
Congressional guidance that the authority to list DPSs be used
``sparingly'' while encouraging the conservation of genetic diversity.
In making this determination, we consider available scientific evidence
of the discrete population segment's importance to the taxon to which
it belongs. Since precise circumstances are likely to vary considerably
from case to case, the DPS policy does not describe all the classes of
information that might be used in determining the biological and
ecological importance of a discrete population. However, the DPS policy
describes four possible classes of information that provide evidence of
a population segment's biological and ecological importance to the
taxon to which it belongs. As specified in the DPS policy (61 FR 4722),
this consideration of the population segment's significance may
include, but is not limited to, the following:
(1) Persistence of the discrete population segment in an ecological
setting unusual or unique to the taxon;
(2) Evidence that loss of the discrete population segment would
result in a significant gap in the range of a taxon;
(3) Evidence that the discrete population segment represents the
only surviving natural occurrence of a taxon that may be more abundant
elsewhere as an introduced population outside its historical range; or
(4) Evidence that the discrete population segment differs markedly
from other populations of the species in its genetic characteristics.
A population segment needs to satisfy only one of these conditions
to be considered significant. Furthermore, other information may be
used as appropriate to provide evidence for significance.
Unique Ecological Setting
Water temperature is a key factor influencing the ecology and
physiology of ectothermic (body temperature regulated by ambient
environmental conditions) salmonid fishes, and can dictate reproductive
timing, growth and development, and life-history strategies.
Groundwater temperatures can be related to air temperatures (Meisner
1990, p. 282), and thus reflect the regional climatic conditions.
Warmer groundwater influences ecological factors such as food
availability, the efficiency with which food is converted into energy
for growth and reproduction, and ultimately growth rates of aquatic
organisms (Allan 1995, pp. 73-79). Aquifer structure and groundwater
temperature is important to salmonid fishes because groundwater can
strongly influence stream temperature, and consequently egg incubation
and fry growth rates, which are strongly temperature-dependent (Coutant
1999, pp. 32-52; Quinn 2005, pp. 143-150).
Missouri River Arctic grayling occur within the 4 to 7 [deg]C (39
to 45 [deg]F) ground water isotherm (see Heath 1983, p. 71; an isotherm
is a line connecting bands of similar temperatures on the earth's
surface), whereas most other North American Arctic grayling are found
in isotherms less than 4 [deg]C, and much of the species' range is
found in areas with discontinuous or continuous permafrost (Meisner et
al. 1988, p. 5; Table 2). Much of the historical range of Arctic
grayling in the upper Missouri River is encompassed by mean annual air
temperature isotherms of 5 to 10 [deg]C (41 to 50 [deg]F) (USGS 2009),
with the colder areas being in the headwaters of the Madison River in
Yellowstone National Park. In contrast, Arctic grayling in Canada,
Alaska, and Asia are located in regions encompassed by air temperature
isotherms 5 [deg]C and colder (41 [deg]F and colder), with much of the
species distributed within the 0 to -10 [deg]C isolines (32 to 14
[deg]F). This difference is significant because Arctic grayling in the
Missouri River basin have evolved in isolation for millennia in a
generally warmer climate than other populations. The potential for
thermal adaptations makes Missouri River Arctic grayling a significant
biological resource for the species under expected climate change
scenarios.
Table 2--Differences Between the Ecological Setting of the Upper
Missouri River and Elsewhere in the Species' Range of Arctic Grayling
------------------------------------------------------------------------
Ecological setting variable Missouri River Rest of taxon
------------------------------------------------------------------------
Bailey's Ecoregion.......... Dry Domain: Polar Domain: Tundra &
Temperate Subarctic Humid
Steppe. Temperate: Marine,
Prairie, Warm
Continental Mountains.
Air temperature (isotherm).. 5 to 10 [deg]C -15 to 5 [deg]C (5 to 41
(41 to 50 [deg]F).
[deg]F).
Groundwater temperature 4 to 7 [deg]C Less than 4 [deg]C (Less
(isotherm). (39 to 45 than 39 [deg]F).
[deg]F).
------------------------------------------------------------------------
Arctic grayling in the upper Missouri River basin occur in a
temperate ecoregion distinct from all other Arctic grayling populations
worldwide, which occur in Arctic or sub-Arctic ecoregions dominated by
Arctic flora and fauna. An ecoregion is a continuous geographic area
within which there are associations of interacting biotic and abiotic
features (Bailey 2005, pp. S14, S23). These ecoregions delimit large
areas within which local ecosystems recur more or less in a predictable
fashion on similar sites (Bailey 2005, p. S14). Ecoregional
classification is hierarchical, and based
[[Page 49394]]
on the study of spatial coincidences, patterning, and relationships of
climate, vegetation, soil, and landform (Bailey 2005, p. S23). The
largest ecoregion categories are domains, which represent
subcontinental areas of similar climate (e.g., polar, humid temperate,
dry, and humid tropical) (Bailey 1994; 2005, p. S17). Domains are
divided into divisions that contain areas of similar vegetation and
regional climates. Arctic grayling in the upper Missouri River basin
are the only example of the species naturally occurring in a dry domain
(temperate steppe division; Table 2). The vast majority of the species'
range is found in the polar domain (all of Asia, most of North
America), with small portions of the range occurring in the humid
temperate domain (northern British Columbia and southeast Alaska).
Occupancy of Missouri River Arctic grayling in a temperate ecoregion is
significant for two primary reasons. First, an ecoregion represents a
suite of factors (climate, vegetation, landform) influencing, or
potentially influencing, the evolution of species within that
ecoregion. Since Missouri River Arctic grayling have existed for
thousands of years in an ecoregion quite different from the majority of
the taxon, they have likely developed adaptations during these
evolutionary timescales that distinguish them from the rest of the
taxon, even if we have yet to conduct the proper studies to measure
these adaptations. Second, the occurrence of Missouri River Arctic
grayling in a unique ecoregion helps reduce the risk of species-level
extinction, as the different regions may respond differently to
environmental change.
Arctic grayling in the upper Missouri River basin have existed for
at least 10,000 years in an ecological setting quite different from
that experienced by Arctic grayling elsewhere in the species' range.
The most salient aspects of this different setting relate to
temperature and climate, which can strongly and directly influence the
biology of ectothermic species (like Arctic grayling). Arctic grayling
in the upper Missouri River have experienced warmer temperatures than
most other populations. Physiological and life-history adaptation to
local temperature regimes are regularly documented in salmonid fishes
(Taylor 1991, pp. 191-193), but experimental evidence for adaptations
to temperature, such as unusually high temperature tolerance or lower
tolerance to colder temperatures, is lacking for Missouri River Arctic
grayling because the appropriate studies have not been conducted. Lohr
et al. (1996, p. 934) studied the upper thermal tolerances of Arctic
grayling from the Big Hole River, but their research design did not
include other populations from different thermal regimes, so it was not
possible to make between-population contrasts under a common set of
conditions. Arctic grayling from the upper Missouri River demonstrate
very high growth rates relative to other populations (Northcote 1995,
p. 157). Experimental evidence obtained by growing fish from
populations under similar conditions would be needed to measure the
relative influence of genetics (local adaptation) versus environment.
We conclude that the occurrence of Arctic grayling in the upper
Missouri River is biogeographically important to the species, that
grayling there have occupied a warmer and more temperate setting that
is distinctly different from the ecological settings relative to the
rest of the species (see Table 2, above), and that they have been on a
different evolutionary trajectory for at least 10,000 years. We
conclude that these differences are significant because they may
provide the species with additional evolutionary resiliency in the
future in light of the changing climate. Consequently, we believe that
Arctic grayling in the upper Missouri River occupy a unique ecological
setting for the species.
Gap in the Range
Arctic grayling in Montana (southern extent is approximately
44[deg]36'23'' N latitude) represent the southern-most extant
population of the species' distribution since the Pleistocene
glaciation. The next-closest native Arctic grayling population outside
the Missouri River basin is found in the Pembina River (approximately
52[deg]55'6.77'' N latitude) in central Alberta, Canada, west of
Edmonton (Blackburn and Johnson 2004, pp. ii, 17; ASRD 2005, p. 6). The
Pembina River drains into Hudson Bay and is thus disconnected from the
Missouri River basin. Loss of the native Arctic grayling of the upper
Missouri River would shift the southern distribution of Arctic grayling
by more than 8[deg] latitude (about 500 miles). Such a dramatic range
constriction would constitute a significant geographic gap in the
species' range and would eliminate a genetically distinct group of
Arctic grayling, which may limit the species' ability to cope with
future environmental change.
Marginal populations, defined as those on the periphery of the
species' range, are believed to have high conservation significance
(Mitikka et al. 2008; Gibson et al. 2009, entire; Haak et al. 2010,
entire; Osborne et al. 2012). Peripheral populations may occur in
suboptimal habitats and thus be subjected to very strong selective
pressures (Fraser 2000, p. 50). Consequently, individuals from these
populations may contain adaptations that may be important to the taxon
in the future. Lomolino and Channell (1998, p. 482) hypothesize that
because peripheral populations should be adapted to a greater variety
of environmental conditions, then they may be better suited to deal
with anthropogenic (human-caused) disturbances than populations in the
central part of a species' range. Arctic grayling in the upper Missouri
River have, for millennia, existed in a climate warmer than that
experienced by the rest of the taxon. If this selective pressure has
resulted in adaptations to cope with increased water temperatures, then
the population segment may contain genetic resources important to the
taxon. For example, if northern populations of Arctic grayling are less
suited to cope with increased water temperatures expected under climate
warming, then Missouri River Arctic grayling might represent an
important population for reintroduction in those northern regions. We
believe that Arctic grayling's occurrence at the southernmost extreme
of the range in the upper Missouri River contributes to the resilience
of the overall taxon because these peripheral populations may possess
increased adaptability relative to the rest of the taxon.
Only Surviving Natural Occurrence of the Taxon That May Be More
Abundant Elsewhere as an Introduced Population Outside of Its
Historical Range
This criterion does not directly apply to the Arctic grayling in
the upper Missouri River because it is not the only surviving natural
occurrence of the taxon; there are native Arctic grayling populations
in Canada, Alaska, and Asia.
Differs Markedly in Its Genetic Characteristics
Differences in genetic characteristics can be measured at the
molecular, genetic, or phenotypic level. Three different types of
molecular markers (allozymes, mtDNA, and microsatellites) demonstrate
that Arctic grayling from the upper Missouri River are genetically
different from those in Canada, Alaska, and Asia (Everett 1986, pp. 79-
80; Redenbach and Taylor 1999, p. 23; Stamford and Taylor 2004, p.
1538; Peterson and Ardren 2009, pp. 1764-1766; USFWS, unpublished
data). These
[[Page 49395]]
data confirm the reproductive isolation among populations that
establishes the discreteness of Missouri River Arctic grayling under
the DPS policy. Here, we speak to whether these data also establish
significance.
Allozymes
Using allozyme data, Everett (1986, entire) found marked genetic
differences among Arctic grayling collected from the Chena River in
Alaska; those descended from fish native to the Athabasca River
drainage in the Northwest Territories, Canada; and native upper
Missouri River drainage populations or populations descended from them
(see Leary 2005, pp. 1-2). The Canadian population had a high frequency
of two unique alleles (forms of a gene), which strongly differentiated
them from all the other samples (Everett 1986, p. 44). With the
exception of one introduced population in an irrigation canal
(Sunnyslope canal) in Montana that is believed to have experienced
extreme genetic bottlenecks, the Chena River (Alaskan) fish were highly
divergent from all the other samples as they possessed an unusually low
frequency of a specific allele (Everett 1986, p. 60; Leary 2005, p. 1),
and contained a unique variant of another allele (Leary 2005, p. 1).
Overall, each of the four native Missouri River populations examined
(Big Hole, Miner, Mussigbrod, and Centennial Valley) exhibited
statistically significant differences in allele frequencies relative to
both the Chena River (Alaska) and Athabasca River (Canada) populations
(Everett 1986, pp. 15, 67).
Combining the data of Everett (1986, entire), Hop and Gharrett
(1989, entire), and Leary (1990, entire) provides information from 21
allozyme loci (genes) from five native upper Missouri River drainage
populations, five native populations in the Yukon River drainage in
Alaska, and the one population descended from the Athabasca River
drainage in Canada (Leary 2005, pp. 1-2). Examination of the genetic
variation in these samples indicated that most of the genetic
divergence is due to differences among drainages (29 percent) and
comparatively little (5 percent) results from differences among
populations within a drainage (Leary 2005, p. 1).
Mitochondrial DNA
Analysis using mtDNA indicates that Arctic grayling in North
America represent at least three evolutionary lineages that are
associated with distinct glacial refugia (Redenbach and Taylor 1999,
entire; Stamford and Taylor 2004, entire). Arctic grayling in the upper
Missouri River basin belong to the so-called North Beringia lineage
(Redenbach and Taylor 1999, pp. 27-28; Samford and Taylor 2004, pp.
1538-1540) because they possess a form of mtDNA that was generally
absent from populations collected from other locations within the
species' range in North America (Redenbach and Taylor 1999, pp. 27-28;
Stamford and Taylor 2004, p. 1538). The notable exceptions were that
some fish from the lower Peace River drainage in British Columbia,
Canada, and all sampled individuals from the Saskatchewan River
drainage Saskatchewan, Canada, also possessed this form of mtDNA
(Stamford and Taylor 2004, p. 1538).
A form of mtDNA common in upper Missouri River Arctic grayling,
which occurs at lower frequencies in other populations, indicates that
Arctic grayling native to the upper Missouri River drainage probably
originated from a glacial refuge in the drainage and subsequently
migrated northwards when the Missouri River temporarily flowed into the
Saskatchewan River and was linked to an Arctic drainage (Cross et al.
1986, pp. 374-375; Pielou 1991, p. 195). When the Missouri River began
to flow southwards because of the advance of the Laurentide ice sheet
(Cross et al. 1986, p. 375; Pileou 1991, p. 10), the Arctic grayling in
the drainage became physically and reproductively isolated from the
rest of the species' range (Leary 2005, p. 2; Campton 2004, p. 6),
which would have included those populations in Saskatchewan.
Alternatively, the Missouri River Arctic grayling could have
potentially colonized Saskatchewan or the Lower Peace River (in British
Columbia) or both post-glacially (Stamford 2001, p. 49) via a gap in
the Cordilleran and Laurentide ice sheets (Pielou 1991, pp. 10-11),
which also might explain the low frequency 'Missouri River'' mtDNA in
Arctic grayling in the Lower Peace River and Upper Yukon River.
We do not interpret the observation that Arctic grayling in Montana
and Saskatchewan, and to lesser extent those from the Lower Peace and
Upper Yukon River systems, share a mtDNA haplotype to mean that these
groups of fish are genetically identical. Rather, we interpret it to
mean that these fish shared a common ancestor tens to hundreds of
thousands of years ago.
Microsatellite DNA
Recent analysis of microsatellite DNA (highly variable portions of
nuclear DNA) showed substantial divergence between Arctic grayling in
Missouri River and Saskatchewan populations (Peterson and Ardren 2009,
entire). This divergence between populations was measured in terms of
allele frequencies, using a metric called Fst (Allendorf and Luikart
2007, pp. 52-54, 198-199). An analogous metric, named Rst, also
measures genetic differentiation between populations based on
microsatellite DNA, but differs from Fst in that it also considers the
size differences between alleles (Hardy et al. 2003, p. 1468). An Fst
or Rst of 0 indicates that populations are the same genetically,
whereas a value of 1 indicates the populations share no genetic
material at the markers being surveyed. Fst values range from 0.13 to
0.31 (average 0.18) between Missouri River and Saskatchewan populations
(Peterson and Ardren 2009, pp. 1758, 1764-1765), whereas Rst values
range from 0.47 to 0.71 (average 0.54) for the same comparisons
(Peterson and Ardren 2009, pp. 1758, 1764-1765). These values indicate
that the two populations differ significantly in allele frequency and
also in the size of those alleles. This outcome indicates that the
observed genetic differences are due to mutational differences, which
suggests the groups may have been separated for millennia (Peterson and
Ardren 2009, pp. 1767-1768).
Analysis of Arctic grayling populations from Alaska, Canada, and
the Missouri River basin using nine of the same microsatellite loci as
Peterson and Ardren (2009, entire) further supports the distinction of
Missouri River Arctic grayling relative to populations elsewhere in
North America (USFWS, unpublished data). This analysis clearly
separated sample fish from 21 populations into two clusters: One
cluster representing populations from the upper Missouri River basin,
and another cluster representing populations from across Canada and
Alaska (USFWS, unpublished data). Divergence in size among these
alleles further supports the distinction between Missouri River Arctic
grayling and those in Canada and Alaska (USFWS, unpublished data). The
interpretation of these data is that the Missouri River populations and
the Canada/Alaska populations are highly genetically distinct at the
microsatellite loci considered.
Phenotypic Characteristics Influenced by Genetics--Meristics
Phenotypic variation can be evaluated by counts of body parts
(i.e., meristic counts of the number of gill rakers, fin rays, and
vertebrae characteristics of a population) that can vary within and
among species. These meristic traits are influenced by both genetics
and the environment (Allendorf and Luikart
[[Page 49396]]
2007, pp. 258-259). When the traits are controlled primarily by genetic
factors, then meristic characteristics can indicate significant genetic
differences among groups. Arctic grayling north of the Brooks Range in
Alaska and in northern Canada had lower lateral line scale counts than
those in southern Alaska and Canada (McCart and Pepper 1971, entire).
These two scale-size phenotypes are thought to correspond to fish from
the North and South Beringia glacial refuges, respectively (Stamford
and Taylor 2004, p. 1545). Arctic grayling from the Centennial Valley
had a phenotype intermediate to the large- and small-scale types
(McCart and Pepper 1971, pp. 749, 754). Arctic grayling populations
from the Missouri River (and one each from Canada and Alaska) could be
correctly assigned to their group 60 percent of the time using a suite
of seven meristic traits (Everett 1986, pp. 32-35). Those native
Missouri River populations that had high genetic similarity also tended
to have similar meristic characteristics (Everett 1986, pp. 80, 83).
Arctic grayling from the Big Hole River showed marked differences
in meristic characteristics relative to two populations from Siberia,
and were correctly assigned to their population of origin 100 percent
of the time (Weiss et al. 2006, pp. 512, 515-516, 518). The populations
that were significantly different in terms of their meristic
characteristics also exhibited differences in molecular genetic markers
(Weiss et al. 2006, p. 518).
Inference Concerning Genetic Differences in Arctic Grayling of the
Missouri River Relative to Other Examples of the Taxon
We believe the differences between Arctic grayling in the Missouri
River and sample populations from Alaska and Canada measured using
allozymes (Everett 1986, entire; Leary 2005, entire), mitochondrial DNA
(Redenbach and Taylor 1999, entire; Stamford and Taylor 2004, entire),
and microsatellite DNA markers (Peterson and Ardren 2009, pp. 1764-
1766; USFWS, unpublished data) represent ``marked genetic differences''
in terms of the extent of differentiation (e.g., Fst,
Rst) and the importance of that genetic legacy to the rest
of the taxon. The presence of morphological characteristics separating
Missouri River Arctic grayling from other populations also likely
indicates genetic differences, although this conclusion is based on a
limited number of populations (Everett 1986, pp. 32-35; Weiss et al.
2006, entire), and we cannot entirely rule out the influence of
environmental variation.
The intent of the DPS policy and the Act is to preserve important
elements of biological and genetic diversity, not necessarily to
preserve the occurrence of unique alleles in particular populations. In
Arctic grayling of the Missouri River, the microsatellite DNA data
indicate that the group is evolving independently from the rest of the
species. The extirpation of this group would mean the loss of the
genetic variation in one of the two most distinct groups identified in
the microsatellite DNA analysis, and the loss of the future
evolutionary potential that goes with it. Thus, the genetic data
support the conclusion that Arctic grayling of the upper Missouri River
represent a unique and irreplaceable biological resource of the type
the Act was intended to preserve. Thus, we conclude that Missouri River
Arctic grayling differ markedly in their genetic characteristics
relative to the rest of the taxon.
Upper Missouri River Arctic grayling satisfy the significance
criteria outlined in the Services' DPS policy because they occur in a
unique ecological setting, are separated from other Arctic grayling
populations by a large gap in their range, and differ markedly in their
genetic characteristics relative to other Arctic grayling populations.
Therefore, we consider the Arctic grayling in the upper Missouri River
basin significant to the taxon to which it belongs under the Service's
DPS policy.
Determination of Distinct Population Segment
We find that a population segment that includes all native ecotypes
of Arctic grayling in the upper Missouri River basin satisfies the
discreteness standard of the DPS policy. The segment is physically
isolated, and genetic data indicate that Arctic grayling in the
Missouri River basin have been separated from other populations for
thousands of years. The population segment occurs in an isolated
geographic area far south of all other Arctic grayling populations
worldwide, and we find that loss of this population segment would
create a significant gap in the species' range. Molecular genetic data
clearly differentiate Missouri River Arctic grayling from other Arctic
grayling populations, including those in Canada and Alaska.
Based on the best scientific and commercial information available,
as described above, we find that, under the Service's DPS policy, upper
Missouri River Arctic grayling are discrete and are significant to the
taxon to which they belong. Because the upper Missouri River population
of Arctic grayling is both discrete and significant, it qualifies as a
DPS under the Act.
As we described above, we are including introduced Arctic grayling
populations that occur in lakes in the upper Missouri River basin as
part of the DPS. The Service has interpreted the Act to provide a
statutory directive to conserve species in their native ecosystems (49
FR 33885; August 27, 1984) and to conserve genetic resources and
biodiversity over a representative portion of a taxon's historical
occurrence (61 FR 4722; February 7, 1996). The introduced Arctic
grayling populations occur within the boundaries of the upper Missouri
River basin and represent moderate to high levels of genetic diversity
from within the basin. The future adaptive capabilities represented by
this genetic diversity have conservation value, particularly given a
changing climate.
We define the historical range of this population segment to
include the major streams, lakes, and tributary streams of the upper
Missouri River (mainstem Missouri, Smith, Sun, Beaverhead, Jefferson,
Big Hole, and Madison Rivers, as well as their key tributaries, as well
as a few small lakes where Arctic grayling are or were believed to be
native (Elk Lake, Red Rock Lakes in the Centennial Valley, Miner Lake,
and Mussigbrod Lake, all in Beaverhead County, Montana)). We define the
current range of the DPS to consist of extant native populations in the
Big Hole River, Miner Lake, Mussigbrod Lake, Madison River-Ennis
Reservoir, and Centennial Valley, as well as all known introduced
populations within the upper Missouri River basin. We refer to this
entity as the Upper Missouri River DPS of Arctic grayling. The
remainder of this finding will thus focus on the population status of
and potential threats to this entity.
Population Status and Trends of Populations in the Upper Missouri River
DPS
The Upper Missouri River DPS of Arctic grayling is comprised of 20
populations, including 2 fluvial populations and 16 adfluvial
populations. Two other populations (Centennial Valley and Madison
River/Ennis Reservoir) appear to exhibit both fluvial and adfluvial
components (Table 3). Arctic grayling from the Centennial Valley (Long
Creek) and Ennis Reservoir/Madison River (mainstem Madison River) have
been documented well past the spawning period through autumn. These
occurrences are more prevalent in Long Creek in the Centennial Valley
than in the Madison
[[Page 49397]]
River population and do not appear to be linked to individual Arctic
grayling seeking thermal refugia during summer (Montana Arctic Grayling
Workgroup (AGW) 1995; p. 1; Cayer 2014a, pers. comm.; MFISH 2014b,
unpublished data). These occurrences include multiple age classes (Age-
1 to Age-3) of Arctic grayling in both Long Creek and the Madison River
and are located in stream reaches that are considerable distances (up
to 15 miles in the Madison River) from adfluvial habitats (Cayer 2014a,
pers. comm.; MFISH 2014b, unpublished data). Eighteen of the 20
populations occur solely on Federal or majority Federal land; the
remaining two (Big Hole River and Ennis Reservoir/Madison River) occur
on primarily private land.
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[GRAPHIC] [TIFF OMITTED] TP20AU14.001
BILLING CODE 4310-55-C
Estimated abundance of reproductively mature individuals in the two
fluvial populations varies from about one hundred to several thousand
[[Page 49399]]
Arctic grayling (Table 3). Where quantitative data are available,
estimated abundance of mature individuals in adfluvial populations
(including the two populations exhibiting both life histories) varies
from a few hundred to around 25,000 Arctic grayling. Most populations
are currently stable or increasing in abundance, with the exception of
the Ennis Reservoir/Madison River population (Table 3).
Distinct Population Segment Five-Factor Analysis
Since the Arctic grayling in the upper Missouri River basin
qualifies as a DPS, we will now evaluate its status with regard to its
potential for listing as endangered or threatened based on the five
factors enumerated in section 4(a) of the Act. Our evaluation of the
Upper Missouri River DPS of Arctic grayling follows.
Summary of Information Pertaining to the Five Factors
Section 4 of the Act (16 U.S.C. 1533) and implementing regulations
(50 CFR 424) set forth procedures for adding species to, removing
species from, or reclassifying species on the Federal Lists of
Endangered and Threatened Wildlife and Plants. Under section 4(a)(1) of
the Act, a species may be determined to be endangered or threatened
based on any of the following 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.
In making this finding, information pertaining to the Upper
Missouri River DPS of Arctic grayling in relation to the five factors
provided in section 4(a)(1) of the Act is discussed below. In
considering what factors might constitute threats, we must look beyond
the mere exposure of the species to the factor to determine whether the
species responds to the factor in a way that causes actual impacts to
the species. If there is exposure to a factor, but no response, or only
a positive response, that factor is not a threat. If there is exposure
and the species responds negatively, the factor may be a threat and we
then attempt to determine how significant a threat it is. If the threat
is significant, it may drive or contribute to the risk of extinction of
the species such that the species warrants listing as endangered or
threatened as those terms are defined by the Act. This does not
necessarily require empirical proof of a threat. The combination of
exposure and some corroborating evidence of how the species is likely
impacted could suffice. The mere identification of factors that could
impact a species negatively is not sufficient to compel a finding that
listing is appropriate; we require evidence that these factors are
operative threats that act on the species to the point that the species
meets the definition of an endangered or threatened species under the
Act.
In making our revised 12-month finding on the petition, we consider
and evaluate the best available scientific and commercial information.
This evaluation includes all factors we previously considered in the
2010 finding and, at the end of this analysis, explains how the
Services' conclusions differ now.
Factor A. The Present or Threatened Destruction, Modification, or
Curtailment of Its Habitat or Range
Curtailment of Range and Distribution
The range and distribution of fluvial Arctic grayling in the upper
Missouri River basin was reduced over the past 100 years (Kaya 1992, p.
51), primarily due to historical habitat fragmentation by dams and
irrigation diversions and by habitat degradation or modification from
unregulated land use (Vincent 1962, pp. 97-121). Fluvial Arctic
grayling typically need large expanses of connected habitat to fulfill
their life-history stages (Armstrong 1986, p. 8). For example, fluvial
Arctic grayling in the Big Hole River have been documented migrating
over 60 miles (97 km) between overwintering, spawning, and foraging
habitats (Shepard and Oswald 1989, pp. 18-21, 27). These past
reductions in range and distribution reproductively isolated fluvial
Arctic grayling populations within the basin (Peterson and Ardren 2009,
p. 1770).
Although the range and distribution of fluvial Arctic grayling has
contracted from historical levels, expression of the fluvial life
history is represented, at least in part, in four Arctic grayling
populations within the Upper Missouri River DPS. Whether strictly
fluvial (e.g., Big Hole and Ruby River) or partially fluvial (e.g.,
Centennial Valley (Long Creek) and Ennis Reservoir/Madison River
(mainstem Madison River)), these populations occur in four watersheds
where large reaches of connected habitat remain and still permit the
expression of the fluvial life history, despite the presence of
mainstem dams in three of four watersheds (Kaya 1992, entire; see
Figure 1). Thus, despite historical curtailment of range, the amount of
connected habitat in some systems is adequate to permit the expression
of the fluvial life history.
Of the four Arctic grayling populations still expressing a fluvial
life history, three of four populations (Big Hole River, Centennial
Valley, and Ruby River) are currently increasing in abundance (see
Table 3). In each of these populations, as abundance increases, there
is a corresponding increase in distribution. Natural reproduction is
occurring in all three of these populations. In the Big Hole River and
the Centennial Valley, remote site incubators (RSIs) have been used as
a conservation tool to help facilitate increased abundance and
distribution of Arctic grayling. Thus, observed increases in abundance
and distribution may be partially attributable to the use of RSIs (for
more in-depth discussion on RSI use, see ``Native Arctic Grayling
Genetic Reserves and Translocation,'' below). Given the above
information, it appears that three of four fluvial, or partly fluvial,
populations are viable and have the necessary configuration and amount
of habitat to fulfill their life-history needs. Thus, effects of past
range curtailment on the fluvial component of Arctic grayling in the
upper Missouri River basin are present, but there appears to be
sufficient adequate habitat remaining to support expression of the
fluvial life history.
Adfluvial Arctic grayling populations in the upper Missouri River
basin are present in all lakes originally thought to have had native
populations historically (Miner, Mussigbrod, Upper Red Rock, and Elk
Lakes (present but not included in Table 3, above, because of uncertain
viability)). Thus, there has been no contraction of the range of
adfluvial populations. Given the above information, curtailment of
range and distribution is not precluding the expression of either
fluvial or adfluvial life history. Although curtailment of range and
distribution occurred historically, Arctic grayling populations are
still present in 7 of 10 historically occupied watersheds in the upper
Missouri River basin (see ``Drainage'' column in Table 3). Accordingly,
we have no evidence that curtailment of range and distribution is a
current threat to the DPS. In addition, we have no information
suggesting curtailment of range and distribution will be a threat in
the future.
Dams on Mainstem Rivers
Much of the historical range of the Upper Missouri River DPS of
Arctic
[[Page 49400]]
grayling has been altered by the construction of dams and reservoirs
(Kaya 1990, pp. 51-52; Kaya 1992, p. 57). The construction of large
dams on mainstem river habitats throughout the upper Missouri River
system fragmented river corridors necessary for the expression of
Arctic grayling migratory life histories in some systems. Construction
of dams that obstructed fish passage on the mainstem Missouri River
(Hauser, Holter, Canyon Ferry, and Toston dams), Madison River
(Madison-Ennis, Hebgen dams), Beaverhead River and its tributary Red
Rock River (Clark Canyon, Lima dams), Ruby River (Ruby dam), and Sun
River (Gibson dam) all likely contributed to the historical decline of
fluvial Arctic grayling in the DPS (Vincent 1962, pp. 127-128; Kaya
1992, p. 57). Lack of fish passage at these dams contributed to the
extirpation of fluvial Arctic grayling from some waters by blocking
migratory corridors (Vincent 1962, p. 128), curtailing access to
important spawning and rearing habitats, and impounding water over
former spawning locations (Vincent 1962, p. 128). Most dams within the
upper Missouri River basin were constructed between 1905 and 1960 (Kaya
1990, entire).
Despite the construction of multiple dams throughout the historical
range of Arctic grayling, multiple populations, or portions of
populations, of the fluvial ecotype are still represented in the DPS.
These populations reside in areas where sufficient quantity and quality
of habitat exist and permit the expression of this life history. In
some cases, dams may be providing a benefit, because currently many of
the dams that historically affected fluvial Arctic grayling populations
are now precluding invasion by nonnative fish from downstream sources.
For example, Lima Dam in the Centennial Valley is currently precluding
brown trout invasion from downstream sources (Mogen 2014, pers. comm).
Currently, there are five Arctic grayling populations within the DPS
that occur above mainstem dams (Centennial Valley, Ruby River, Hyalite
Lake, Diversion Lake, and Gibson Reservoir) with at least one nonnative
fish species occurring downstream of these dams (MFISH 2014d,
unpublished data).
Some reservoirs created by dams are currently being used by Arctic
grayling as overwintering, rearing and foraging areas. Both adult and
juvenile Arctic grayling use Ennis Reservoir for overwintering,
rearing, and foraging (Byorth and Shepard 1990, entire). In the
Centennial Valley, Arctic grayling have recently been detected in Lima
Reservoir (MFISH 2014e, unpublished data). The movements of Arctic
grayling within and out of Lima Reservoir are unknown; however, Lima
Reservoir is a large reservoir and, as such, is likely used for
overwintering purposes.
Arctic grayling have been documented in stream and river reaches
below some dams, most likely indicating downstream passage of fish over
or through dams. These fish are essentially ``lost'' to the population
residing above the dam, because none of the mainstem river dams in the
upper Missouri River basin provides upstream fish passage. Substantial
losses from a population resulting from downstream entrainment of fish
through dams could cause declines in reproductive potential and
abundance in the reservoir population above the dam (Kimmerer 2008,
entire). However, it is unknown what entrainment rates currently are in
populations residing near dams. Rate of entrainment is likely dependent
on a number of factors, including dam operations, season, water
conditions in the reservoir, initial population size above the dam,
etc. Recent monitoring data and angler reports of Arctic grayling
observed downstream of reservoirs supporting Arctic grayling
populations are sporadic (Horton 2014c, pers. comm.; SSA 2014); thus it
appears the threat of mainstem dams is likely affecting some
individuals, but not affecting populations or the DPS as a whole.
Historically, operational practices at Madison Dam have likely
affected the Arctic grayling population in Ennis Reservoir/Madison
River. A population decline in Arctic grayling appeared to coincide
with a reservoir drawdown in the winter of 1982-1983 (Byorth and
Shepard 1990, pp. 52-53). This drawdown likely affected the forage
base, rearing habitat, and spawning cycle of Arctic grayling in the
reservoir. However, under a new licensing agreement dated September 27,
2000, between the Federal Energy Regulatory Commission and Ennis Dam
operators, such substantial drawdowns in elevation of Ennis Reservoir
are no longer permitted (Clancey 2014, pers. comm.).
Given the above information, mainstem dams were a historical threat
to Arctic grayling populations in the upper Missouri River basin. Dams
still impact individuals, because some Arctic grayling are currently
being entrained and lost from their source population. In Ennis
Reservoir, the new licensing agreement is expected to reduce the
effects of dam operations on the Arctic grayling population. Most
Arctic grayling populations residing above dams are stable or
increasing; thus, it does not appear this impact is acting at the
population or DPS level. We have no information to conclude that
mainstem dams will be a threat in the future at the population or DPS
level.
Water Management in the Upper Missouri River Basin
The predominant use of private lands in the upper Missouri River
basin is irrigated agriculture and ranching. These activities have
historically had significant effects on aquatic habitats, primarily
changes in water availability and alteration of the structure and
function of aquatic habitats. Changes in water availability can affect
Arctic grayling reproduction, survival, and movements among habitat
types (Kaya 1990, entire).
In contrast to most of the Arctic grayling populations in the Upper
Missouri River DPS that occur on Federal land, the fluvial population
of Arctic grayling in the Big Hole River occurs on primarily (~90
percent) private land. Thus, any conservation efforts conducted in the
Big Hole River Valley need support from involved agencies and private
landowners. In 2006, a candidate conservation agreement with assurances
(CCAA; Montana Fish, Wildlife, and Parks et al. 2006, entire) was
developed for Arctic grayling in the Big Hole River. The conservation
goal of this CCAA is to secure and enhance the fluvial population of
Arctic grayling in the upper Big Hole River drainage. Conservation
projects conducted under the CCAA are prioritized and guided by the Big
Hole Arctic Grayling Strategic Habitat Conservation Plan (SHCP) (for
more specific information, see ``Conservation Efforts to Reduce Habitat
Destruction, Modification, or Curtailment of Its Range,'' below).
Since 2006, many conservation and restoration projects have been
completed in the upper Big Hole River under the direction of the CCAA
and SHCP (Table 4). Below, we describe and evaluate the implementation
and effectiveness of these projects relative to the potential stressors
analyzed under Factor A for the Big Hole River population. We also
analyze the effects of potential stressors under Factor A for the other
Arctic grayling populations in the DPS.
[[Page 49401]]
Table 4--Conservation Projects and Results, and Arctic Grayling Response in the Big Hole River Since
Implementation of the Big Hole CCAA in 2006
[All information on conservation projects and conservation results cited from the Big Hole Arctic Grayling
Strategic Habitat Conservation Plan]
----------------------------------------------------------------------------------------------------------------
Conservation projects Arctic grayling
Threat factor Stressor \a\ Conservation result response
----------------------------------------------------------------------------------------------------------------
A..................... Dams/habitat Fish ladders: 41..... Stream miles (%) Number of
fragmentation. Bridges: 7........... accessible to breeding adults has
Grade control grayling \b\: increased from ~100
structures: 2. Tier I- (2007-2011) to 500-
82(98%; pre- 900 \c\ (2013)
CCAA=87%).. (Leary 2014,
Tier II- unpublished data).
61(67%; pre-
CCAA=27%)..
Tier III-
32(20%: pre-
CCAA=6%)..
Dewatering/Thermal PODs: 343 of 504 with Arctic
stress. signed SSPs. Achievement of grayling abundance
Irrigation instream flow \d\ (catch per unit
improvements: 88. goals increased effort) increased
Water measuring from 50% (pre- from 0.2 fish/mile
devices: 67. CCAA) to 78% (2008) to 1.4 fish/
Stock water systems: (post-CCAA). mile (2012) in the
63. Landowner CCAA monitoring
Stream restoration: contributions to reaches of the
26 miles. streamflow mainstem Big Hole
Rock Creek increasing as River (MFWP 2013a,
restoration. of PODs unpublished data).
with signed SSPs
increase [landowner
contribution to
instream flows in
Big Hole River (pre-
2006 = 0 cfs; 2013
= 250 cfs)].
Temperature
reductions in
tributaries (see
Rock Creek example
below).
Pre-restoration Arctic
(2007):. grayling abundance
36 days \d\ (catch per unit
max. temp >70 effort) increased
[deg]F. from 2.9 fish/mile
16 days (2008) to 7.4 fish/
max. temp >77 mile (2012) in the
[deg]F. CCAA monitoring
Post-restoration tributaries (MFWP
(2013):. 2013a, unpublished
0 days max data).
temp. >70 [deg]F. Arctic
grayling
distribution has
increased 4 miles
in Rock Creek
(young-of-year and
Age 1+) and 2 miles
in Big Lake Creek
(Age 1+) since 2006
(SHCP 2013, p. 12).
Entrainment........... Fish screens: 2...... No
Prioritized entrainment
monitoring protocol. documented since
2010.
Observed low
entrainment rates in
unscreened ditches
(73 Arctic grayling/
138 ditch miles).
Riparian habitat loss. Stream restoration: 110 miles
26 miles. (65%) of riparian
Riparian fencing: 108 habitat on enrolled
miles. lands improving.
Stock water systems: 15% increase
63. in sustainable
Grazing mgmt. plans: riparian areas from
21 landowners 32% (2006) to 47%
(85,000 ac.). (2013).
Noxious weed Adaptive
management. management in place
Willow planting to address non-
(72,200 planted). improving areas.
----------------------------------------------------------------------------------------------------------------
\a\ PODs = Points of Diversion, SSPs = Site-specific plans; \b\ Tier I is core spawning, rearing and adult
habitat that is currently occupied by Arctic grayling, Tier II is periphery habitat intermittently used by
Arctic grayling, Tier III is suitable, but currently unoccupied historical habitat; \c\ The estimate of number
of breeding adults in the Big Hole River in 2013 is reported as a range because of uncertainty in the
frequency rate of rare alleles in the analysis; \d\ Abundance estimates from 2013 were lower than those
reported for 2012 likely due to unusually high flows (3X normal) concurrent with fall sampling that likely
decreased capture efficiency, resulting in lower abundance estimates in 2013.
[[Page 49402]]
Habitat Fragmentation/Smaller Seasonal Barriers
Big Hole River: Smaller dams or diversions associated with
irrigation structures historically posed a threat to Arctic grayling
migratory behavior, especially in the Big Hole River drainage. In the
Big Hole River, numerous diversion structures have been identified as
putative fish migration barriers (Petersen and Lamothe 2006, pp. 8, 12-
13, 29) that may limit the ability of Arctic grayling to migrate to
spawning, rearing, or sheltering habitats under certain conditions. As
with the larger dams, these smaller fish passage barriers can reduce
reproduction (access to spawning habitat is blocked), reduce growth
(access to feeding habitat is blocked), and increase mortality (access
to refuge habitat is blocked). Historically, these types of barriers
were numerous and widespread across the Big Hole River drainage.
Currently, habitat fragmentation due to irrigation diversion
structures in the Big Hole is being systematically reduced under the
CCAA for Fluvial Arctic Grayling in the upper Big Hole River
(hereafter, Big Hole CCAA or CCAA; for more specific information, see
``Conservation Efforts to Reduce Habitat Destruction, Modification, or
Curtailment of Its Range'') and Big Hole Arctic Grayling SHCP. Since
2006, 41 fish ladders have been installed in the mainstem Big Hole
River and tributaries (Table 4). Multiple culverts have been replaced
with bridges and several grade control structures have been installed
(Table 4). As a result, no fish barriers now exist in the mainstem
upper Big Hole River. Almost all (98 percent) of tier I habitat and the
majority (68 percent) of tier II habitat is connected and accessible to
Arctic grayling (Table 4): 67 miles of stream have been reconnected in
the Big Hole River system since 2006 (MFWP 2014a, unpublished data).
Other populations: Smaller fish passage barriers also have been
noted to affect Arctic grayling in the Centennial Valley (Unthank 1989,
p. 9). Historically, spawning Arctic grayling migrated from the
Jefferson River system, through the Beaverhead River and Red Rock River
through the Red Rock Lakes and into the upper drainage, and then
returned downstream after spawning (Henshall 1907, p. 5). The
construction of a water control structure (sill) at the outlet of Lower
Red Rock Lake in 1930 (and reconstruction in 1957 (USFWS 2009, p. 74))
created an upstream migration barrier that blocked these migrations
(Unthank 1989, p. 10; Gillin 2001, p. 4-4). However, recent changes in
water management at the Red Rock Lakes National Wildlife Refuge (NWR)
have resulted in year-round fish passage through the control structure
at the outlet of Lower Red Rock Lake (West 2013, pers. comm.).
In Mussigbrod Lake, Arctic grayling occasionally pass downstream
over a diversion structure at the lake outlet, and become trapped in an
isolated pool (Olsen 2014, pers. comm.). During high-snowpack years,
Arctic grayling likely can swim back up to the lake from the pool, but
in low snowpack years, some Arctic grayling perish when the isolated
pool dries up (Olsen 2014, pers. comm.). However, this phenomenon has
occurred periodically in recent history and has had no discernible
impacts on Arctic grayling abundance in Mussigbrod Lake (Olsen 2014,
pers. comm.).
All 16 adfluvial Arctic grayling populations in the upper Missouri
River basin occur on Federal land (U.S. Forest Service) and are not
influenced by irrigation structures because none are present. The
effect of a barrier at the outlet of Mussigbrod Lake is likely
impacting individuals, but not the population because of the robust
population size in Mussigbrod Lake and historical stability of that
population since the outlet structure was created. Based on this
information, we conclude that the threats from habitat fragmentation
have been sufficiently mitigated or minimized and are no longer are
acting as a stressor at the population or DPS level.
Degradation of Riparian Habitat
Riparian corridors are important for maintaining habitat for Arctic
grayling in the upper Missouri River basin, and in general are critical
for the ecological function of aquatic systems (Gregory et al. 1991,
entire). Riparian zones are important for Arctic grayling because of
their effect on water quality and water temperature, and their role in
maintaining natural ecological process responsible for creating and
maintaining necessary physical habitat features (i.e., pools, riffles,
and scour areas) used by the species to meet its life-history
requirements.
Big Hole: Arctic grayling abundance in the upper Big Hole River is
positively related to the presence of overhanging vegetation, primarily
willows (Salix spp.), that is associated with pool habitat (Lamothe and
Magee 2004, pp. 21-22). Removal of willows and riparian clearing
concurrent with livestock and water management along the upper Big Hole
River has led to a shift in channel form (i.e., braided channels
becoming a single wide channel), increased erosion rates, reduced
cover, increased water temperatures, and reduced recruitment of large
wood debris into the active stream channel (Confluence Consulting et
al. 2003, pp. 24-26). These factors combine to reduce the suitability
of the habitat for species like Arctic grayling (Hubert 1985, entire).
Currently, restoration of riparian areas in the upper Big Hole
River system is a priority under the CCAA (for more specific
information, see ``Conservation Efforts to Reduce Habitat Destruction,
Modification, or Curtailment of Its Range,'' below). Since 2006,
efforts to restore and conserve riparian habitats have been numerous
and multi-faceted (see Table 4). About 170 miles (274 km) of riparian
habitat are currently enrolled in the Big Hole CCAA, out of a total of
about 340 miles (547 km) of total riparian habitat in the CCAA
Management Area. Of the enrolled riparian habitat, 65 percent (110
miles (177 km)) is improving in condition, as rated by a standardized
riparian protocol (NRCS 2004, entire). Further, 47 percent of enrolled
riparian habitat (80 miles (129 km)) is functioning at a sustainable
level, which is a 15 percent increase in 5 years (MTFWP et al. 2006, p.
92; see Table 4). A sustainable rating indicates that the stream can
access its flood plain, transport its sediment load, build banks, store
water, and dissipate flood energy in conjunction with a healthy
riparian zone (NRCS 2004, p. 7). Riparian habitats are reassessed every
5 years and are scored on 10 stability and sustainability metrics (for
example, stream incisement), with any reach scoring at 80 percent or
above rated as sustainable (NRCS 2004, entire). In addition, adaptive
management within the CCAA framework will allow for reevaluation of
conservation measures being implemented in non-improving habitat.
Other populations: In the Centennial Valley, historical livestock
grazing both within the Red Rock Lakes NWR and on adjacent private
lands negatively affected the condition of riparian habitats on
tributaries to the Red Rock Lakes (Mogen 1996, pp. 75-77; Gillin 2001,
pp. 3-12, 3-14). In general, degraded riparian habitat limits the
creation and maintenance of aquatic habitats, especially pools, which
are preferred habitats for adult Arctic grayling (Lamothe and Magee
2004, pp. 21-22; Hughes 1992, entire), although many spawning adult
Arctic grayling in Red Rock Creek outmigrate soon after spawning and
likely do not use available pool habitat (Jordan 2014, pers. comm.).
Loss of riparian vegetation
[[Page 49403]]
increases bank erosion, which can lead to siltation of spawning
gravels, which may in turn harm Arctic grayling by reducing the extent
of suitable spawning habitat and reducing survival of Arctic grayling
embryos already present in the stream gravels.
Recently, the Red Rock Lakes NWR acquired land on Red Rock Creek,
upstream of the refuge boundary (West 2014a, pers. comm.). Much of this
parcel was riparian habitat that was historically heavily grazed; thus,
the refuge implemented a rest-rotation grazing system where more
durable lands are grazed while more sensitive lands (e.g., riparian
areas) are rested for up to 4 years. On average, grazing intensities on
the refuge have decreased from 20,000 Animal Unit Months (AUMs, number
of cow/calf pairs multiplied by the number of months grazed) to about
5,000 AUMs. As a result of these changes, riparian habitat within the
refuge has dramatically improved (West 2014b, pers. comm.) and is
expected to continue improving under the new grazing regime. Concurrent
with riparian improvement within Red Rock Lakes NWR, the number of
adult Arctic grayling migrating up Red Rock Creek to spawn has
increased from fewer than 500 to more than 2,000 (Patterson 2014,
unpublished data). Given the riparian improvements within Red Rock
Lakes NWR, and that the refuge represents the vast majority of current
Arctic grayling habitat in the Centennial Valley, the effects of
degraded riparian habitat do not appear to be acting on the core of the
Centennial Valley population at the individual or population level.
Most of the riparian habitat surrounding high-elevation lakes on
Federal land where the remaining populations are found is intact and of
high quality (MFISH 2014a, unpublished data; MFWP 2014e, unpublished
data; USFS 2014, p. 2), because these habitats are in remote locations
or wilderness areas with little anthropogenic disturbance. Given that
riparian degradation is being systematically addressed in the Big Hole
River and Centennial Valley on the National Refuge land where the
majority of Arctic grayling reside, we conclude that riparian
degradation is not a current threat to the DPS. Riparian habitat is
expected to remain intact on Federal land because of existing
regulatory mechanisms (see in Factor D discussion, below). Riparian
habitat in the Big Hole River is expected to continue improving because
of the proven track record of conservation evidenced by the current
upward trend in riparian habitat quality. As more site-specific plans
are signed under the Big Hole CCAA, more riparian improvement is
expected because conservation measures will be similar between
currently implemented and future site-specific plans. Given that
riparian habitat is intact or improving for populations of Arctic
grayling occurring on Federal land and the Big Hole population, and
these populations account for 19 of 20 populations in the DPS, we
conclude riparian habitat degradation is not a current rangewide threat
and is not expected to become a threat in the future.
Dewatering From Irrigation and Consequent Increased Water Temperatures
Demand for irrigation water in the semi-arid upper Missouri River
basin historically dewatered many rivers formerly or currently occupied
by Arctic grayling. The primary effects of this dewatering were: (1)
Increased water temperatures, and (2) reduced habitat capacity. In
ectothermic species like salmonid fishes, water temperature sets basic
constraints on species' distribution and physiological performance,
such as activity and growth (Coutant 1999, pp. 32-52). Increased water
temperatures can reduce the growth and survival of Arctic grayling
(physiological stressor). Reduced habitat capacity can concentrate
fishes and thereby increase competition and predation (ecological
stressor). Below we discuss the potential effects of increased water
temperature on the Upper Missouri River DPS of Arctic grayling. For
discussion of the potential effects of reduced habitat capacity, see
Cumulative Effects from Factors A through E, below.
Big Hole: In the Big Hole River system, surface-water (flood)
irrigation has altered the natural hydrologic function of the river
(Shepard and Oswald 1989, p. 29; Byorth 1993, p. 14; 1995, pp. 8-10;
Magee et al. 2005, pp. 13-15). An inverse relationship between flow
volume and water temperature (i.e., lower flows can lead to higher
water temperatures) is apparent in the Big Hole River (Flynn et al.
2008, pp. 44, 46, but see Sladek 2013, p. 31). Summer water
temperatures exceeding 21 [deg]C (70 [deg]F) are considered to be
physiologically stressful for cold-water fish species, such as Arctic
grayling (Hubert et al. 1985, pp. 7, 9). Summer water temperatures
consistently exceed 21 [deg]C (70 [deg]F) in the mainstem of Big Hole
River (Cayer and McCullough 2012, p. 7; (Cayer and McCullough 2013, p.
6) and have exceeded the upper incipient lethal temperature (UILT; the
temperature that is survivable for periods longer than 1 week by 50
percent of a ``test population'' in an experimental setting) for Arctic
grayling (e.g., 25 [deg]C or 77 [deg]F) (Lohr et al. 1996). As a
result, thermal fish kills have been documented in the Big Hole River
(Lohr et al. 1996, p. 934) in the past. The most recent fish kill in
the Big Hole River that we are aware of occurred in 1994, and included
eight fish species, including Arctic grayling (Lohr et al. 1996, p.
934).
Arctic grayling in the Big Hole River use tributaries as a thermal
refuge when summer water temperatures in the mainstem become stressful
(Vatland et al. 2009, p. 11). Summer water temperatures within most
tributaries are cooler than those observed in some reaches of the
mainstem Big Hole River (Vatland et al. 2009, entire; MFWP 2014b,
unpublished data).
Since 2006, water conservation and restoration projects associated
with the Big Hole Arctic grayling CCAA (for more specific information,
see ``Conservation Efforts to Reduce Habitat Destruction, Modification,
or Curtailment of Its Range,'' below) have been implemented to increase
instream flows and reduce water temperatures in the Big Hole River and
tributaries. Varying flow targets for different management segments of
the Big Hole River were outlined in the CCAA, based on the wetted
perimeter method, a biologically based method for determining instream
flow requirements to provide necessary resources for all life stages of
Arctic grayling. Over 300 irrigation diversions are operated under flow
agreements within finalized site-specific plans (Table 4). The 10
remaining site-specific plans representing the remainder of points of
diversion are expected to be signed in August 2014. Although we are
aware of the future potential of more points of diversion being managed
under signed site plans to contribute to Arctic grayling conservation,
we do not consider these anticipated future efforts to contribute to
Arctic grayling conservation currently, and have not considered them as
part of this status review or our listing determination for this DPS.
Multiple other projects designed to decrease dewatering and thermal
stress have been implemented since 2006 (Table 4). The collective
result of these efforts are increasing streamflows, increased access to
cold-water refugia via fish ladders, and marked temperature reductions,
particularly in some tributaries (Table 4).
Specific flow targets were developed for the different Management
Segments in the CCAA Management Area (see MFWP et al. 2006, pp. 7, 9,
13, for more information on CCAA Management
[[Page 49404]]
Segments). The goal for increasing instream flow was to achieve flow
targets 75 percent of days in each Management Segment during years of
average or greater snowpack. This goal was based on a comparison
between minimum flow targets and historical streamflows recorded in
Management Segments C and D. Achieving flow targets 75 percent of days
in each Management Segment was intended to be a general goal because
many other factors influence instream flows in the Big Hole River that
are outside the control of landowners (e.g., snowpack, precipitation).
Before implementation of the CCAA (2000-2005), average flow targets
were met among all Management Segments 50 percent of the time, and
since implementation of the CCAA (2006-2012), they have been met 78
percent of the time (SHCP 2013, p. 12). Thus, the targets are being
met.
Consistently since 2006, one management area, known as Management
Segment C, has exhibited the lowest instream flows among all Management
Segments. In part, instream flows in Management Segment C are
influenced by several large diversions immediately upstream of the flow
measuring device at the downstream boundary of Management Segment C
(Robert 2014, pers. comm.). Some of this diverted water is returned to
the Big Hole River downstream of the flow measuring device (Robert
2014, pers. comm.). As such, instream flows in Management Segment C
represent the ``worst case'' scenario among all Management Segments.
The Montana Department of Natural Resources and Conservation conducted
an analysis of this ``worst case'' scenario, to explore how instream
flows in Management Segment C have changed since the inception of the
Big Hole CCAA. Given that natural factors such as summer precipitation
and annual snowpack influence instream flows in the Big Hole River, the
analysis of instream flows in Management Segment C included comparisons
among several years of similar (but below average) snow pack and
similar summer precipitation, both before and after CCAA implementation
(Table 5).
Table 5--Comparison of Number of Days Varying Flow Targets Were Achieved
Among Similar Years of Below Average Snowpack in the Big Hole River CCAA
Management Segment C, Pre- and Post CCAA. All Information in This Table
Cited From Roberts 2014, Unpublished Data
------------------------------------------------------------------------
Pre-CCAA Post-CCAA
-----------------------------------
1988 2003 2012 2013
------------------------------------------------------------------------
Peak snowpack (percent of average).. \a\73 108 81 \a\75
May-Aug. precipitation (in.)........ 4.14 3.85 4.74 5.14
July-Aug. temps (degrees F; -1.3 8.0 1.4 1.9
departure from normal).............
Signed SSPs......................... 0 0 12 15
Landowner contributions (cfs)....... 0 0 252 260
Days <160 \b\ cfs................... 50 8 11 40
Days <60 \b\ cfs.................... 123 123 87 69
Days <20 cfs........................ 79 68 0 28
Days <10 cfs........................ 65 7 0 1
Mean discharge (cfs; July-Sept.).... 8.4 19.7 45 39
Mean discharge (cfs; Aug.).......... 1.1 14.2 33.7 21
-----------------------------------
Total Days 60 years) with no observed declines in abundance.
Predation by Birds and Mammals
In general, the incidence and effect of predation by birds and
mammals on Arctic grayling is not well understood because few detailed
studies have been completed (Northcote 1995, p. 163). Black bear (Ursus
americanus), mink (Neovison vison), and river otter (Lontra canadensis)
are present in southwestern Montana, but direct evidence of predatory
activity by these species is often lacking (Kruse 1959, p. 348). Osprey
(Pandion haliaetus) can capture Arctic grayling during the summer
(Kruse 1959, p. 348). In the Big Hole River, Byorth and Magee (1998, p.
926) attributed the loss of Arctic grayling from artificial enclosures
used in a competition experiment to predation by minks, belted
kingfisher (Ceryle alcyon), osprey, and great blue heron (Ardea
herodia). In addition, American white pelican (Pelecanus
erythrorhynchos) are seasonally present in the Big Hole River, and they
also may feed on Arctic grayling. The aforementioned mammals and birds
can be effective fish predators; however, Arctic grayling evolved with
these native predator species and have developed life-history and
reproductive strategies to mitigate for predation losses. We have no
data demonstrating any of these species historically or currently
consume Arctic grayling at levels sufficient to exert a measureable,
population-level impact on native Arctic grayling in the upper Missouri
River system. We expect the current situation to continue, so we
conclude that predation by birds and mammals does not constitute a
threat to Missouri River Arctic grayling now or in the future.
Summary of Factor C
Based on the information available at this time, we conclude
disease does not represent a past or current threat to the Upper
Missouri River DPS of Arctic grayling. We have no basis for concluding
that disease may become a future threat.
Predation and competition can influence the distribution,
abundance, and diversity of species in ecological communities.
Predation by and competition with nonnative species can negatively
affect native species, particularly those that are stressed or
occurring at low densities due to unfavorable environmental conditions.
Historically, the impact of predation and competition from nonnatives
was likely greater because many of the habitats used by Arctic grayling
were degraded. Thus, predation and competition likely played a role
historically in decreasing the abundance and distribution of Arctic
grayling. Currently, habitat conditions have improved markedly for
those Arctic grayling populations on Federal land (18 of 20
populations) and for the Big Hole River population on primarily private
land. Predation and competition with nonnative species are still
occurring in these systems, although the extent and magnitude of these
effects appears to be mediated by habitat quality. Abundance of Arctic
grayling and nonnative brown trout are increasing in the Big Hole
River. Before suppression efforts began, Yellowstone cutthroat hybrids
and Arctic grayling spawners were both at 40 year highs in Red Rock
Creek in the Centennial Valley. We acknowledge nonnative trout
densities are high in the Madison River and may be contributing to the
decline of that Arctic grayling population; however, most other
adfluvial populations appear to have stable abundance of Arctic
grayling and nonnatives. Thus, based on our review we have no
information that predation or competition represents a threat at the
DPS level on the Upper Missouri River DPS of Arctic grayling. Further,
Arctic grayling experts project only a small effect of predicted
nonnative trout densities on Arctic grayling recruitment in the future.
Thus, we have no information that predation or competition from
nonnative trout represents a future threat at the population or species
level.
Little is known about the effect of predation on Arctic grayling by
birds and mammals. Such predation likely does occur, but we are not
aware of any situation where an increase in fish-eating birds or
mammals has coincided with the decline of Arctic grayling.
Consequently, the available information does not support a conclusion
that predation by birds or mammals represents a substantial past,
present, or future threat to native Arctic grayling in the upper
Missouri River.
Factor D. The Inadequacy of Existing Regulatory Mechanisms
Section 4(b)(1)(A) of the Act requires the Service to take into
account ``those efforts, if any, being made by any State or foreign
nation, or any political subdivision of a State or foreign nation, to
protect such species . . .'' We consider relevant Federal, State, and
Tribal laws, and regulations when evaluating the status of the species.
Regulatory mechanisms, if they exist, may preclude the need for listing
if we determine that such mechanisms adequately address the threats to
the species such that listing is not warranted. Only existing
ordinances, regulations, and laws, that have a direct connection to a
law, are enforceable and permitted are discussed in this section. All
other measures are discussed under the specific relevant factor.
U.S. Federal Laws and Regulations
No Federal laws in the United States specifically address the
Arctic grayling, but several, in their implementation, may affect the
species' habitat.
National Environmental Policy Act
All Federal agencies are required to adhere to the National
Environmental Policy Act (NEPA) of 1970 (42 U.S.C. 4321 et seq.) for
projects they fund, authorize, or carry out. The Council on
Environmental Quality's regulations for implementing NEPA (40 CFR parts
1500-1518) state that, when preparing environmental impact statements,
agencies shall include a discussion on the environmental impacts of the
various project alternatives, any adverse environmental effects which
cannot be avoided, and any irreversible or irretrievable commitments of
resources involved (40 CFR part 1502). The NEPA itself is a disclosure
law, and does not require subsequent minimization or mitigation
measures by the Federal agency involved. Although Federal agencies may
include conservation measures for Arctic grayling as a result of the
NEPA process, any such measures are typically voluntary in nature and
are not required by NEPA.
[[Page 49415]]
Federal Land Policy and Management Act
The Federal Land Policy and Management Act (FLPMA) of 1976 (43
U.S.C. 1701 et seq.), as amended, states that the public lands shall be
managed in a manner that will protect the quality of scientific,
scenic, historical, ecological, environmental, air and atmospheric,
water resource, and archeological values. This statute protects lands
within the range of the Arctic grayling managed by the Bureau of Land
Management (BLM).
The BLM considers the fluvial Arctic grayling a sensitive species
requiring special management consideration for planning and
environmental analysis (BLM 2009a, entire, BLM 2009b, entire). The BLM
has recently developed a resource management plan (RMP) for the Dillon
Field Office Area that provides guidance for the management of over
900,000 acres of public land administered by BLM in southwest Montana
(BLM 2006a, p. 2). The Dillon RMP area thus includes the geographic
area that contains the Big Hole, Miner, Mussigbrod, Madison River, and
Centennial Valley populations of Arctic grayling. A RMP planning area
encompasses all private, State, and Federal lands within a designated
geographic area (BLM 2006a, p. 2), but the actual implementation of the
RMP focuses on lands administered by the BLM that typically represent
only a fraction of the total land area within that planning area (BLM
2006b, entire). Restoring Arctic grayling habitat and ensuring the
long-term persistence of both fluvial and adfluvial ecotypes are among
the RMP's goals (BLM 2006a, pp. 30-31). However, there is little actual
overlap between the specific parcels of BLM land managed by the Dillon
RMP and the current distribution of Arctic grayling (BLM 2006b,
entire).
The BLM also has a RMP for the Butte Field Office Area, which
includes more than 300,000 acres in south-central Montana (BLM 2008,
entire), including portions of the Big Hole River in Deerlodge and
Silver Bow counties (BLM 2008, p. 8; 2009c, entire). The Butte RMP
considers conservation and management strategies and agreements for
Arctic grayling in its planning process and includes a goal to
opportunistically enhance or restore habitat for Arctic grayling (BLM
2008, pp. 10, 30, 36). However, the Butte RMP does not mandate specific
actions to improve habitat for Arctic grayling in the Big Hole River
and little overlap exists between BLM-managed lands and Arctic grayling
occupancy in this planning area.
National Forest Management Act
Under the U.S. Forest Service (USFS) National Forest Management Act
(NFMA) of 1976, as amended (16 U.S.C. 1600 et seq.), the USFS strives
to provide for a diversity of plant and animal communities when
managing national forest lands. Individual national forests may
identify species of concern that are significant to each forest's
biodiversity. The USFS considers fluvial Arctic grayling a sensitive
species (USFS 2004, entire) for which population viability is a
concern. However, this designation provides no special regulatory
protections.
Most of the upper Missouri River grayling populations occur on
National Forest land; all 16 adfluvial populations and the fluvial Ruby
River population (majority on National Forest) occur on USFS-managed
lands. These populations occur across four different National Forests;
consequently the riparian habitats surrounding the lakes and
tributaries are managed according to the standards and guidelines
outlined in each National Forest Plan. All Forest Plans do not contain
the same standards and guidelines; however, each Plan has standards and
guidelines for protecting riparian areas around perennial water
sources. In the Beaverhead-Deerlodge and Helena National Forest Plans,
the Inland Native Fish Strategy (INFS) standards and guidelines have
been incorporated. The INFS, in part, defines widths of riparian buffer
zones adequate to protect streams and lakes from non-channelized
sediment inputs and contribute to other riparian functions, such as
stream shading and bank stability. These protections have been
incorporated into the Beaverhead-Deerlodge and Helena National Forest
Plans through amendments and are currently preserving intact riparian
areas around most, if not all, adfluvial Arctic grayling habitats.
Exceptions to the riparian protections outlined in INFS are
occasionally granted; however, these exceptions require an analysis of
potential effects and review by a USFS fish biologist.
On the Gallatin National Forest, standards and guidelines in the
Forest Plan include using ``best management practices (BMPs)'' to
protect water sources and riparian areas. Similar to INFS, BMPs outline
buffer strips along watercourses where disturbance and activity is
minimized to protect riparian areas and water quality. On the Lewis and
Clark National Forest, standards and guidelines are in place to leave
timbered buffer strips adjacent to waterbodies to protect riparian
areas. Grayling habitat on the Gallatin and Lewis and Clark National
Forests consists of seven high-elevation mountain lakes.
The NFMA and INFS are adequately protecting riparian habitat on
National Forest land, given the intact nature of most riparian areas
surrounding the high-elevation lake populations and the Ruby River.
National Park Service (NPS) Organic Act
The NPS Organic Act of 1916 (16 U.S.C. 1 et seq.), as amended,
states that the NPS ``shall promote and regulate the use of the Federal
areas known as national parks, monuments, and reservations . . . to
conserve the scenery and the national and historic objects and the wild
life therein and to provide for the enjoyment of the same in such
manner and by such means as will leave them unimpaired for the
enjoyment of future generations.'' Arctic grayling are native to the
western part of Yellowstone National Park and habitats are managed
accordingly for the species under the Native Species Management Plan
(NPS 2010, entire). One adfluvial Arctic grayling population, Grebe
Lake, currently occurs in Yellowstone National Park. The Grebe Lake
population is one of the larger adfluvial populations (see Table 3,
above) in the DPS. The habitat in Grebe Lake and the tributaries is
managed for conservation (NPS 2010, p. 44). Further, it is expected
that these habitats will be managed for conservation in the future,
based on provisions in the Organic Act and guidance outlined in the
Native Species Management Plan.
National Wildlife Refuge System Improvement Act of 1997
The National Wildlife Refuge Systems Improvement Act (NWRSIA) of
1997 (Pub. L. 105-57) amends the National Wildlife Refuge System
Administration Act of 1966 (16 U.S.C. 668dd et seq.). The NWRSIA
directs the Service to manage the Refuge System's lands and waters for
conservation. The NWRSIA also requires monitoring of the status and
trends of refuge fish, wildlife, and plants. The NWRSIA requires
development of a comprehensive conservation plan (CCP) for each refuge
and management of each refuge consistent with its plan.
The Service has developed a final CCP to provide a foundation for
the management and use of Red Rock Lakes NWR (USFWS 2009, entire) in
the Centennial Valley. Since the development of the CCP, Refuge staff
have conducted numerous habitat conservation/restoration projects to
benefit Arctic grayling, including:
[[Page 49416]]
Removal of an earthen dam whose reservoir inundated several hundred
meters of historical Arctic grayling spawning habitat in Elk Springs
Creek, and subsequent reintroductions and tracking of young-of-year
Arctic grayling in Elk Springs Creek (West 2014a, pers. comm.). However
to date, the reintroductions in Elk Springs Creek have not established
a spawning run. Other conservation projects conducted on the Refuge
include the acquisition of new land and decreases in grazing
intensities from 20,000 AUMs to about 5,000 AUMs. The Refuge has
implemented a rest-rotation grazing system where more durable lands are
grazed while more sensitive lands (e.g., riparian areas) are rested for
up to 4 years (West 2014a, pers. comm.). Some active riparian
restoration has also occurred, including a project to reconnect Red
Rock Creek to a historical channel and replacement of four culverts to
allow for natural tributary migration across alluvial fans (West 2014a,
pers. comm.). The Refuge is also actively engaged in supporting ongoing
graduate research efforts to explore potential limiting factors for
Arctic grayling in the Centennial Valley.
Other conservation projects under the CCP have been focused on
potential nonnative species effects on Arctic grayling, namely a 5-year
project removing hybrid cutthroat trout captured during their upstream
spawning run and a study of dietary overlap between Arctic grayling and
Yellowstone cutthroat trout (West 2014a, pers. comm.). The Refuge also
operates a sill dam (previous upstream fish barrier) to provide
upstream fish passage and operates one irrigation ditch only when
snowpack is average or above and timing is such that young Arctic
grayling are not present near the diversion (West 2014a, pers. comm.).
The NWRSIA is adequately protecting habitat for Arctic grayling on
the Refuge because riparian habitats are improving and the Centennial
Valley population is increasing in both abundance and distribution. The
proven track record of completed conservation projects on the refuge
and currently expanding Arctic grayling population indicate that the
continued implementation of the CCP during the next 15 years (which is
the life of the CCP) will continue to improve habitat conditions on the
refuge.
Federal Power Act (FPA)
The Federal Power Act of 1920 (16 U.S.C. 791 et seq., as amended)
provides the legal authority for the Federal Energy Regulatory
Commission (FERC), as an independent agency, to regulate hydropower
projects. In deciding whether to issue a license, FERC is required to
give equal consideration to mitigation of damage to, and enhancement
of, fish and wildlife (16 U.S.C. 797(e)). A number of FERC-licensed
dams exist in the Missouri River basin in current (i.e., Ennis Dam on
the Madison River) and historical Arctic grayling habitat (e.g., Hebgen
Dam on the Madison River; Hauser, Holter, and Toston dams on the
mainstem Missouri River; and Clark Canyon Dam on the Beaverhead River).
The FERC license expiration dates for these dams range from 2024
(Toston) to 2059 (Clark Canyon) (FERC 2010, entire). None of these
structures provides upstream passage of fish, and such dams are
believed to be one of the primary factors that led to the historical
decline of Arctic grayling in the Missouri River basin (see discussion
under Factor A, above). However, recent monitoring data indicate
multiple stable Arctic grayling populations occurring above mainstem
dams, with the exception of the Ennis Reservoir/Madison River
population. The drawdowns in reservoir water level believed to have
historically affected the Ennis Reservoir/Madison River Arctic grayling
population are not permitted under a new licensing agreement between
the Federal Energy Regulatory Commission and Madison Dam operators, as
we described previously in this finding (Clancey 2014, pers. comm.).
This change in water management in Ennis Reservoir will ensure adequate
rearing and foraging habitat for this population. The fluvial ecotype
is still represented in the DPS and both strictly fluvial Arctic
grayling populations appear to be stable or increasing. Thus, we
conclude the Federal Power Act is currently adequate to protect the
Upper Missouri River DPS of Arctic grayling at the population and DPS
level.
Clean Water Act
The Clean Water Act (CWA) of 1972 (33 U.S.C. 1251 et seq.)
establishes the basic structure for regulating discharges of pollutants
into the waters of the United States and regulating quality standards
for surface waters. The CWA's general goal is to ``restore and maintain
the chemical, physical, and biological integrity of the Nation's
waters'' (33 U.S.C. 1251(a)). The CWA requires States to adopt
standards for the protection of surface water quality and establishment
of total maximum daily load (TMDL) guidelines for rivers. The Big Hole
River has approved TMDL plans for its various reaches (MDEQ 2009a,
entire; 2009b, entire); thus, complete implementation of this plan
should improve water quality (by reducing water temperatures, and
reducing sediment and nutrient inputs) in the Big Hole River in the
future. As of September 2013, there was no significant TMDL plan
development activity in the Madison River or Red Rock watershed in the
Centennial Valley (see MDEQ 2014). Currently, TMDL documents have been
approved for the Ruby River. All planning areas containing other
adfluvial Arctic grayling populations in the upper Missouri River basin
have approved TMDLs, including the Gallatin, Lake Helena, and Sun
watersheds (see MDEQ 2014).
Currently, water temperatures in the Big Hole River exceed levels
outlined in the TMDL. However, reductions in water temperature within
tributaries have been demonstrated (see discussion under Factor A and
Table 4). Given that most Arctic grayling populations within the upper
Missouri River basin are stable or increasing and habitats are largely
being managed in a manner that benefits the species, we have no
evidence that the CWA is inadequately protecting Arctic grayling at the
population or DPS level.
State Laws
Montana Environmental Policy Act
The legislature of Montana enacted the Montana Environmental Policy
Act (MEPA) as a policy statement to encourage productive and enjoyable
harmony between humans and their environment, to protect the right to
use and enjoy private property free of undue government regulation, to
promote efforts that will prevent or eliminate damage to the
environment and biosphere and stimulate the health and welfare of
humans, to enrich the understanding of the ecological systems and
natural resources important to the State, and to establish an
environmental quality council (MCA 75-1-102). Part 1 of the MEPA
establishes and declares Montana's environmental policy. Part 1 has no
legal requirements, but the policy and purpose provide guidance in
interpreting and applying statutes. Part 2 requires State agencies to
carry out the policies in Part 1 through the use of systematic,
interdisciplinary analysis of State actions that have an impact on the
human environment. This is accomplished through the use of a
deliberative, written environmental review. In practice, MEPA provides
a basis for the adequate review of State actions in order to ensure
that environmental concerns are fully considered (MCA 75-1-102).
Similar to NEPA, the MEPA is largely a disclosure
[[Page 49417]]
law and a decision-making tool that does not specifically require
subsequent minimization or mitigation measures.
Laws Affecting Physical Aquatic Habitats
A number of Montana State laws have a permitting process applicable
to projects that may affect stream beds, river banks, or floodplains.
These include the Montana Stream Protection Act (SPA), the Streamside
Management Zone Law (SMZL), and the Montana Natural Streambed and Land
Preservation Act (Montana Department of Natural Resources (MDNRC) 2001,
pp. 7.1-7.2). The SPA requires that a permit be obtained for any
project that may affect the natural and existing shape and form of any
stream or its banks or tributaries (MDNRC 2001, p. 7.1). The Montana
Natural Streambed and Land Preservation Act (i.e., MNSLPA or 310
permit) requires private, nongovernmental entities to obtain a permit
for any activity that physically alters or modifies the bed or banks of
a perennially flowing stream (MDNRC 2001, p. 7.1). The SPA and MNSLPA
laws do not mandate any special recognition for species of concern, but
in practice, biologists that review projects permitted under these laws
usually stipulate restrictions to avoid harming such species (Horton
2010, pers. comm.). The SMZL regulates forest practices near streams
(MDNRC 2001, p. 7.2). The Montana Pollutant Discharge Elimination
System (MPDES) Stormwater Permit applies to all discharges to surface
water or groundwater, including those related to construction,
dewatering, suction dredges, and placer mining, as well as to
construction that will disturb more than 1 acre within 100 ft (30.5 m)
of streams, rivers, or lakes (MDNRC 2001, p. 7.2).
Review of applications by MFWP, MTDEQ, or MDNRC is required prior
to issuance of permits under the above regulatory mechanisms (MDNRC
2001, pp. 7.1-7.2). These regulatory mechanisms are expected to limit
impacts to aquatic habitats in general. Given that most Arctic grayling
populations are stable or increasing in abundance in the presence of
these regulatory mechanisms, we have no basis for concluding that these
regulatory mechanisms are inadequate to protect the Arctic grayling and
their habitat now or in the future.
Montana Water Use Act
The purpose of the Montana Water Use Act (Title 85: Chapter 2,
Montana Codes Annotated) is to provide water for existing and future
beneficial use and to maintain minimum flows and water quality in
Montana's streams. The Missouri River system is generally believed to
be overappropriated, and water for additional consumptive uses is only
available for a few months during very wet years (MDNRC 1997, p. 12).
However, the upper Missouri River basin and Madison River basin have
been closed to new water appropriations because of water availability
problems, overappropriation, and a concern for protecting existing
water rights (MDNRC 2009, p. 45). In addition, recent compacts (a legal
agreement between Montana, a Federal agency, or an Indian tribe
determining the quantification of federally or tribally claimed water
rights) have been signed that close appropriations in specific waters
in or adjacent to Arctic grayling habitats. For example, the USFWS-Red
Rock Lakes-Montana Compact includes a closure of appropriations for
consumptive use in the drainage basins upstream of the most downstream
point on the Red Rock Lakes NWR and the Red Rock Lakes Wilderness Area
(MDNRC 2009, pp. 18, 47). The NPS-Montana Compact specifies that
certain waters will be closed to new appropriations when the total
appropriations reach a specified level, and it applies to Big Hole
National Battlefield and adjacent waters (North Fork of the Big Hole
River and its tributaries including Ruby and Trail Creeks), and the
portion of Yellowstone National Park that is in Montana (MDNRC 2009, p.
48).
The State of Montana is currently engaged in a Statewide effort to
adjudicate (finalize) water rights claimed before July 1, 1973. The
final product of adjudication in a river basin is a final decree. To
reach completion, a decree progresses through several stages: (1)
Examination, (2) temporary preliminary decree, (3) preliminary decree,
(4) public notice, (5) hearings, and (6) final decree (MDNRC 2009, pp.
9-14). As of February 2014, the Centennial Valley has a preliminary
decree, and the Big Hole and Madison Rivers have preliminary temporary
decrees (MDNRC 2014, entire). We anticipate the final adjudication of
all the river basins in Montana that currently contain native Arctic
grayling will be completed in the next 5 years, but we do not know if
this process will eliminate the overallocation of water rights. We note
that the overallocation of water in some systems within the upper
Missouri river basin is of general concern to Arctic grayling because
of the species' need for adequate quantity and quality of water for all
life stages. However, we have no information indicating that
overallocation of water in the upper Missouri River basin is a current
threat at the individual or DPS level because most populations are
stable or increasing at this time. Therefore, we conclude that the
Montana Water Use Act is adequate to protect the Arctic grayling and
its habitat.
Angling Regulations
Arctic grayling is considered a game fish (MFWP 2010, p. 16), but
is subject to special catch-and-release regulations in streams and
rivers within its native range, as was described under Factor B, above
(MFWP 2014d, p. 51). Catch-and-release regulations also are in effect
for Ennis Reservoir on the Madison River and Red Rock Creek in the
Centennial Valley (MFWP 2014d, p. 63). Arctic grayling in other
adfluvial populations are subject to more liberal regulations; anglers
can keep up to 5 per day and have up to 10 in possession in accordance
with standard daily and possession limits for that angling management
district (MFWP 2014d, p. 51). We have no evidence to indicate that
current fishing regulations are inadequate to protect native Arctic
grayling in the Missouri River basin (see discussion under Factor B,
above).
Summary of Factor D
Current Federal and State regulatory mechanisms are adequate to
protect Arctic grayling of the upper Missouri River. We conclude this
because the majority of populations are on Federal land where
regulatory mechanisms are in place to preserve intact habitats and are
expected to remain in place. In the Big Hole River, fluvial Arctic
grayling generally occupy waters adjacent to private lands (MFWP et al.
2006, p. 13; Lamothe et al. 2007, p. 4), so Federal regulations may
have limited ability to protect that population. However, some Federal
regulations (e.g., CWA, FPA, NMFA, NWRSIA, NPS Organic Act) in concert
with other existing conservation efforts (e.g., Big Hole CCAA) are
adequate to sustain and improve habitat conditions for Arctic grayling.
Arctic grayling in the Big Hole River appear to be responding
positively to these improvements. In addition, we did not identify
other threats to the DPS that would require regulatory protections.
For the reasons described above, we conclude that existing
regulatory mechanisms are adequate to protect the Upper Missouri River
DPS of Arctic grayling. We do not anticipate any changes to the
existing regulatory mechanisms; thus we conclude that existing
regulatory mechanisms will remain adequate in the future.
[[Page 49418]]
Factor E. Other Natural or Manmade Factors Affecting Its Continued
Existence
Drought
Drought is a natural occurrence in the interior western United
States (see National Drought Mitigation Center 2010). The duration and
severity of drought in Montana appears to have increased during the
last 50 years, and precipitation has tended to be lower than average in
the last 20 years (National Climatic Data Center 2010). Drought can
affect fish populations by reducing stream flow volumes. This leads to
dewatering and high temperatures that can limit connectivity among
spawning, rearing, and sheltering habitats. Drought can also reduce the
volume of thermally suitable habitat and increase the frequency of
water temperatures above the physiological limits for optimum growth
and survival in Arctic grayling. In addition, drought can interact with
human-caused stressors (e.g., irrigation withdrawals, riparian habitat
degradation) to further reduce stream flows and increase water
temperatures.
Reduced stream flows and elevated water temperatures during drought
have been most apparent in the Big Hole River system (Magee and Lamothe
2003, pp. 10-14; Magee et al. 2005, pp. 23-25; Rens and Magee 2007, pp.
11-12, 14). In the Big Hole River, evidence for the detrimental effects
of drought on Arctic grayling populations is primarily inferential;
observed declines in fluvial Arctic grayling and nonnative trout
abundances in the Big Hole River coincide with periods of drought
(Magee and Lamothe 2003, pp. 22-23, 28) and fish kills (Byorth 1995,
pp. 10-11, 31).
Although the response of stream and river habitats to drought is
expected to be most pronounced because of the strong seasonality of
flows in those habitats, effects in lake environments can occur. For
example, both the Upper and Lower Red Rock Lakes are very shallow
(Mogen 1996, p. 7). Increased frequency or duration of drought could
lead to increased warming in shallower lakes, such as Upper Red Rock
Lake. However, the Centennial Valley has many springs sources that
could, at least in part, mitigate for increases in water temperature
due to increased drought frequency and magnitude. Other potential
effects from drought could include a reduction in overall lake depth,
which could in turn affect summer or overwintering habitat. Adfluvial
populations in high mountain lakes would likely not be affected
significantly by drought because air (and thus water) temperatures in
these habitats are relatively cool due to the greater distance from sea
level at high elevations (~ a 3.6 [deg]F (6.5 [deg]C) decrease in air
temperature for every 3,200 ft. (1 kilometer) above sea level; Physics
2014). In addition, most of these habitats are relatively large bodies
of water volumetrically, thus are resistant to warming, given the high
specific heat of water (USGS 2014). Further, intact riparian areas in
these habitats buffer against water temperature increases in
tributaries by blocking incoming solar radiation (Sridhar et al. 2004,
entire; Cassie 2006, p. 1393).
Given the climate of the intermountain West, we conclude that
drought has been and will continue to be a natural occurrence. We
assume that negative effects of drought on Arctic grayling populations,
such as reduced connectivity among habitats or increased water
temperatures at or above physiological thresholds for growth and
survival, are more frequent in stream and river environments and in
very shallow lakes relative to larger, deeper lakes. As discussed under
Factor A, the implementation of the Big Hole Arctic grayling CCAA is
likely to minimize some of the effects of drought in the Big Hole
River, by reducing the likelihood that human-influenced actions or
outcomes (irrigation withdrawals, destruction of riparian habitats, and
fish passage barriers) will interact with the natural effects of
drought (reduced stream flows and increased water temperatures). We
expect the impact of drought may act at the individual level, but not
at the population or DPS level because most grayling populations reside
in drought-resistant habitats in high mountain lakes. Some populations
will likely be affected by drought, but implemented conservation
measures (Big Hole River population) and natural spring sources
(Centennial Valley) are expected to minimize the impact. Overall, we
conclude that drought has been a past threat when many historical
habitats were degraded, but is not a current threat because of the
intact nature of most habitats occupied by Arctic grayling in the upper
Missouri River basin. Drought is expected to increase in both duration
and severity in the future; however, resiliency currently being
incorporated into riparian and aquatic habitats through conservation
projects will likely buffer the effects of drought. Thus, drought is
not expected to pose a threat to the DPS in the future.
Stochastic (Random) Threats, Genetic Diversity and Small Population
Size
A principle of conservation biology is that the presence of larger
and more productive (resilient) populations can reduce overall
extinction risk. To minimize extinction risk due to stochastic (random)
threats, life-history diversity should be maintained, populations
should not all share common catastrophic risks, and both widespread and
spatially close populations are needed (Fausch et al. 2006, p. 23;
Allendorf et al. 1997, entire).
The Upper Missouri River DPS of Arctic grayling exists largely as a
collection of isolated populations (Peterson and Ardren 2009, entire),
with little to no gene flow among populations. While the inability of
fish to move between populations limits genetic exchange and
demographic support (Hilderbrand 2003, p. 257), large population sizes
coupled with adequate number of breeding individuals minimize the
effects of isolation. For example, Grebe Lake, a large population,
receives no genetic infusion from any other population in the upper
Missouri River basin, yet has a very large estimated effective
population size (see Table 3, above). Loss of genetic diversity from
genetic drift is not a concern for this population, despite it being
reproductively isolated.
Abundance among the 20 Arctic grayling populations varies widely
(see Table 3, above). Individually, small populations like Ruby River
need to maintain enough adults to minimize loss of variability through
genetic drift and inbreeding (Rieman and McIntyre 1993, pp. 10-11). The
point estimates of the effective number of breeders observed in all
populations (where data are available) are above the level at which
inbreeding is an immediate concern (Leary 2014, pers. comm.). The Ruby
River population exhibits a low effective number of breeders, but
contains the second highest genetic diversity among all populations
(Leary 2014, unpublished data). Thus, inbreeding depression is probably
not a concern for this population in the near future (Leary 2014, pers.
comm.).
Effective population size estimates for other Arctic grayling
populations vary from 162 to 1,497 (see Table 3, above). There has been
considerable debate about what effective population size is adequate to
conserve genetic diversity and long-term adaptive potential (see
Jamieson and Allendorf 2012 for review, p. 579). However, loss of
genetic diversity is typically not an immediate threat even in isolated
populations with an Ne >100 (Palstra and Ruzzante 2008, p.
3441), but rather is a symptom of deterministic processes acting on the
population (Jamieson and Allendorf
[[Page 49419]]
2012, p. 580). In other words, loss of genetic diversity due to small
effective population size typically does not drive species to
extinction (Jamieson and Allendorf 2012, entire); other processes, such
as habitat degradation, have a more immediate and greater impact on
species persistence (Jamieson and Allendorf 2012). We acknowledge that
loss of genetic diversity can occur in small populations; however, in
this case, it appears that there are adequate numbers of breeding
adults to minimize loss of genetic diversity. Thus, we conclude that
loss of genetic diversity is not a threat at the DPS level.
Conservation of life-history diversity is important to the
persistence of species confronted by habitat change and environmental
perturbations (Beechie et al. 2006, entire). Therefore, the
reintroductions of fluvial Arctic grayling into the upper Ruby River
that have occurred provide redundancy of the fluvial ecotype. The
number of breeding individuals in the Ruby River population has
increased over the last 3 years (Leary 2014, unpublished data). Thus,
there is now a viable replicate of the fluvial ecotype.
Populations of Arctic grayling in the Upper Missouri River DPS are
for the most part widely separated from one another, occupying 7 of 10
historically occupied watersheds (see Table 3, above). Thus, risk of
extirpation by a rare, high-magnitude environmental disturbance (i.e.,
catastrophe) is relatively low. In addition, multiple spawning
locations exist for 11 of the 20 populations in the Upper Missouri
River DPS. The 11 populations with access to multiple spawning
tributaries include all the largest populations in terms of abundance,
except Mussigbrod Lake (see Table 3). Abundance and number of breeding
individuals is adequate in most populations to sustain moderate to high
levels of genetic diversity currently observed. Based on this
information, we conclude that stochastic processes are not a threat to
the Upper Missouri River DPS of Arctic grayling and are not expected to
be in the future.
Summary of Factor E
Overall, we conclude that the Upper Missouri River DPS of Arctic
grayling has faced historical threats from drought, loss of genetic
diversity, and small population size. However, the DPS currently exists
as multiple, isolated populations across a representative portion of
its historical range. While reproductive isolation can lead to
detrimental genetic effects, the current size of most Arctic grayling
populations, trends in effective population size, and number of
breeders suggest these effects will be minimal. Redundancies within and
among populations are present: Multiple spawning tributaries,
geographic separation, life-history replication. Given this
information, we conclude the redundant nature of multiple resilient
populations across a representative portion of the species' historical
range minimizes the impacts of drought, low abundance, reduced genetic
diversity, and lack of a fluvial ecotype replicate. Thus, these are not
current threats, and are not expected to be threats in the future.
Cumulative Effects From Factors A Through E
We limit our discussion of cumulative effects from Factors A
through E to interactions involving climate change. Our rationale for
this is that climate change has the highest level of uncertainty among
other factors considered, and likely has the most potential to affect
Arctic grayling populations when interacting with other factors.
Climate Change and Nonnative Species Interactions
Changes in water temperature due to climate change may influence
the distribution of nonnative trout species (Rahel and Olden 2008, p.
524) and the outcome of competitive interactions between those species
and Arctic grayling. Brown trout are generally considered to be more
tolerant of warm water than many salmonid species common in western
North America (Coutant 1999, pp. 52-53; Selong et al. 2001, p. 1032),
and higher water temperatures may favor brown trout where they compete
against salmonids with lower thermal tolerances (Rahel and Olden 2008,
p. 524). Recently, observed increases in the abundance and distribution
of brown trout in the upper reaches of the Big Hole River (MFWP 2013,
unpublished data) may be consistent with the hypothesis that stream
warming is facilitating encroachment. However, the effect of increased
abundance and distribution of brown trout on Arctic grayling in the Big
Hole River is unknown.
Currently, brown trout are at relatively low densities (<20 fish/
mile) in the upper Big Hole River, where Arctic grayling densities are
highest (MFWP 2013e, unpublished data). At densities of 100 brown trout
per mile (a plausible future scenario), Arctic grayling experts
predicted a 5 percent reduction in Arctic grayling recruitment in the
Big Hole River, due to competition and predation (SSA 2014, p. 2).
Given that natural mortality of salmonid fry is typically high (>90
percent) (Kruse 1959, pp. 329, 333; Bradford 1995, p. 1330), the
predicted reductions in Arctic grayling recruitment by current and
future densities of brown trout in the Big Hole River will likely not
impact Arctic grayling at the population level. Thus, the potential
cumulative effect of climate change and nonnative species interactions
is not a current or future threat for the Upper Missouri River DPS of
Arctic grayling.
Climate Change and Dewatering
Synergistic interactions are possible between effects of climate
change and effects of other potential stressors such as dewatering.
Increases in temperature and changes in precipitation are likely to
affect the availability of water in the West. However, it is difficult
to project how climate change will affect water availability because
increased air and water temperatures may be accompanied and tempered by
more frequent precipitation events. Uncertainty about how different
temperature and precipitation scenarios could affect water availability
make projecting possible synergistic effects of climate change on the
Arctic grayling too speculative at this time.
Summary
Recent genetic analyses have concluded that many of the introduced
populations of Arctic grayling in the upper Missouri River basin
contain moderate to high levels of genetic diversity and that these
populations were created from local sources within the basin. These
introduced populations currently occur within the confines of the upper
Missouri River basin and occupy high quality habitats on Federal land,
the same places the Service would look to for long-term conservation of
the species, if needed. As such, these populations and their future
adaptive potential have conservation value and are included in the
Upper Missouri River DPS of Arctic grayling.
Currently, we recognize 20 populations of Arctic grayling in the
Upper Missouri River DPS, 18 of which occur on Federal land. Adequate
regulatory mechanisms exist to ensure the conservation of habitat on
Federal land for these populations. Historical habitat degradation on
private land has affected the Big Hole River population; however,
habitat conditions have been improving since the implementation of the
Big Hole CCAA in 2006. Conservation actions associated with the Big
Hole CCAA and SHCP have reduced water temperatures in tributaries,
increased instream flows in
[[Page 49420]]
tributaries and the mainstem Big Hole River, connected almost all core
habitat for Arctic grayling, and improved riparian health. Arctic
grayling have responded favorably to these improvements because
abundance and distribution have increased throughout the upper Big Hole
River, and number of breeding adults has increased by a factor of at
least 5 since 2006. The Service is encouraged by the successful track
record of conservation actions implemented under the Big Hole CCAA and
SHCP over the past 7 years.
Riparian restoration efforts in the Big Hole River and Centennial
Valley are ongoing and will continue to be key in mitigating the
anticipated effects of drought and climate change. Increased shading of
tributaries and decreased width-to-depth ratios in stream channels can
effectively minimize effects from increasing air temperatures and
drought. In addition, these changes to habitat can alter predation and
competition potential where both nonnative species and Arctic grayling
coexist, as they have for over 100 years in some populations.
We acknowledge the uncertainty regarding the current status of the
Ennis Reservoir/Madison River population and probable declining trend
in abundance. The factors influencing the current demographics of this
population are unclear. However, we are encouraged by the recent FERC
relicensing agreement precluding reservoir drawdowns that likely
affected this population and its habitat in the past.
In conclusion, we find viable populations of both ecotypes present
in the DPS, the majority of which occur on Federal land and are
protected by Federal land management measures. Numbers of breeding
adults are currently increasing in both strictly fluvial populations
and in the Centennial Valley. High-quality habitat is present for most
populations or is improving where it is not optimal (e.g., Big Hole
River). Health of riparian areas is trending upward and will be key to
minimizing effects of climate change and drought. All Arctic grayling
populations are genetically diverse, are of Montana-origin, and occur
in 7 of 10 historically occupied watersheds.
In 2010, we identified multiple threats as acting on the Upper
Missouri River DPS of Arctic grayling. At that time, we determined that
habitat-related threats included habitat fragmentation, dewatering,
thermal stress, entrainment, riparian habitat loss, and effects from
climate change. Since 2010, we have 4 additional years of monitoring
data and have gained new insight. It is now apparent that these threats
are being effectively mitigated on private land (Big Hole River) by
conservation actions under the Big Hole CCAA and do not appear to be
present or acting at a level to warrant concern on most of the
adfluvial populations. Almost all (98 percent) of Arctic grayling core
habitat in the Big Hole River is now connected. Recent riparian
restoration activities have appreciably reduced water temperatures and
improved riparian habitat in tributaries to the Big Hole River and are
expected to buffer the effects of climate change. Entrainment of Arctic
grayling into irrigation canals in the Big Hole system is low, with no
documented entrainment occurring since 2010. Habitats on Federal land
are largely intact and these populations are not subject to many of the
stressors historically identified for other populations because no
irrigation diversions are present, habitats are primarily high-
elevation lakes that have cool water temperatures, and riparian areas
are largely intact.
In 2010, another threat identified as acting on the Upper Missouri
River DPS of Arctic grayling was the presence of nonnative trout. We
considered nonnative trout a threat at that time because we were aware
of several instances where Arctic grayling declines had occurred
following nonnative trout introductions. Currently, we have a better
understanding of the interactions between nonnative trout and Arctic
grayling. Our review of these interactions and case histories suggests
that habitat degradation, concurrent with nonnative trout
introductions, likely contributed to historical declines in Arctic
grayling in those instances. Further, it appears the effect of
nonnative trout on Arctic grayling are likely habitat-mediated;
nonnative trout affect Arctic grayling disproportionately when habitat
conditions are degraded, but both Arctic grayling and nonnatives can
coexist at viable levels when habitat conditions are improved. The
primary evidence supporting this assertion is the increasing abundance
and distribution of both Arctic grayling and nonnatives in the Big Hole
River (brown trout) and Centennial Valley (Yellowstone cutthroat trout
before suppression began). Another line of evidence to support this
assertion is observed spatial segregation between nonnatives and Arctic
grayling in the core Arctic grayling areas in the Big Hole River,
especially spawning and rearing areas (SSA 2014). In addition, Arctic
grayling in adfluvial habitats have maintained stable or increasing
population levels in the presence of brook, rainbow, and Yellowstone
cutthroat trout for over 100 years in many instances in the upper
Missouri River basin, where habitat degradation has not occurred or
been extensive.
In 2010, we stated that existing regulatory mechanisms were
inadequate to protect the Upper Missouri River DPS of Arctic grayling.
The primary reason for this assertion was that Arctic grayling
populations were reported as declining; thus existing regulatory
mechanisms were believed to be inadequate because they had failed to
halt or reverse this decline. Currently, we have updated information
indicating that 19 of 20 populations of Arctic grayling are either
stable or increasing. Existing regulatory mechanisms have precluded
riparian habitat destruction on Federal lands or mandated restoration
of impaired areas and are expected to provide similar protections in
the future. Given the updated information, we now believe these
regulatory mechanisms are adequate.
In 2010, we identified reduced genetic diversity, low abundance,
random events, drought, and lack of a fluvial replicate as threats to
the Upper Missouri River DPS of Arctic grayling. Updated genetic
information that was not available in 2010 indicates moderate to high
levels of genetic diversity within most Arctic grayling populations in
the DPS. Further, abundance estimates derived from this updated genetic
information indicate higher Arctic grayling abundances than previously
thought. Adequate redundancy exists within the DPS to minimize the
effects of random events and drought; lake habitats occupied by most
Arctic grayling populations are drought-resistant. Lastly, a viable
fluvial replicate now exists (Ruby River), with 5 years of natural
reproduction documented and an increasing number of breeding adults.
Finding
As required by the Act, we considered the five factors in assessing
whether the Upper Missouri River DPS of Arctic grayling is endangered
or threatened throughout all of its range. We examined the best
scientific and commercial information available regarding the present
and future threats faced by the Upper Missouri River DPS of Arctic
grayling. We reviewed the petition, information available in our files
and other available published and unpublished information, including
information submitted by the public, and we consulted with recognized
Arctic grayling experts and other Federal and State agencies. Habitat-
related threats previously identified,
[[Page 49421]]
including habitat fragmentation, dewatering, thermal stress,
entrainment, riparian habitat loss, and effects from climate change,
have been sufficiently ameliorated and the information indicates that
19 of 20 populations of Arctic grayling are either stable or
increasing. On the basis of the best scientific and commercial
information available and the analysis provided above, we find that the
magnitude and imminence of threats do not indicate that the Upper
Missouri River DPS of Arctic grayling is in danger of extinction
(endangered), or likely to become endangered within the foreseeable
future (threatened), throughout its range. Therefore, we find that
listing the Upper Missouri River DPS throughout its range as a
threatened or an endangered species is not warranted at this time.
Significant Portion of the Range
Under the Act and our implementing regulations, a species may
warrant listing if it is an endangered or a threatened species
throughout all or a significant portion of its range. The Act defines
``endangered species'' as any species which is ``in danger of
extinction throughout all or a significant portion of its range,'' and
``threatened species'' as any species which is ``likely to become an
endangered species within the foreseeable future throughout all or a
significant portion of its range.'' The term ``species'' includes ``any
subspecies of fish or wildlife or plants, and any distinct population
segment [DPS] of any species of vertebrate fish or wildlife which
interbreeds when mature.'' On July 1, 2014, we published a final policy
interpreting the phrase ``Significant Portion of its Range'' (SPR) (79
FR 37578). The final policy states that (1) if a species is found to be
an endangered or a threatened species throughout a significant portion
of its range, the entire species is listed as an endangered or a
threatened species, respectively, and the Act's protections apply to
all individuals of the species wherever found; (2) a portion of the
range of a species is ``significant'' if the species is not currently
an endangered or a threatened species throughout all of its range, but
the portion's contribution to the viability of the species is so
important that, without the members in that portion, the species would
be in danger of extinction, or likely to become so in the foreseeable
future, throughout all of its range; (3) the range of a species is
considered to be the general geographical area within which that
species can be found at the time FWS or NMFS makes any particular
status determination; and (4) if a vertebrate species is an endangered
or a threatened species throughout an SPR, and the population in that
significant portion is a valid DPS, we will list the DPS rather than
the entire taxonomic species or subspecies.
The SPR policy is applied to all status determinations, including
analyses for the purposes of making listing, delisting, and
reclassification determinations. The procedure for analyzing whether
any portion is an SPR is similar, regardless of the type of status
determination we are making. The first step in our analysis of the
status of a species is to determine its status throughout all of its
range. If we determine that the species is in danger of extinction, or
likely to become so in the foreseeable future, throughout all of its
range, we list the species as an endangered (or threatened) species and
no SPR analysis will be required. If the species is neither an
endangered nor a threatened species throughout all of its range, we
determine whether the species is an endangered or a threatened species
throughout a significant portion of its range. If it is, we list the
species as an endangered or a threatened species, respectively; if it
is not, we conclude that listing the species is not warranted.
When we conduct an SPR analysis, we first identify any portions of
the species' range that warrant further consideration. The range of a
species can theoretically be divided into portions in an infinite
number of ways. However, there is no purpose to analyzing portions of
the range that are not reasonably likely to be significant and either
an endangered or a threatened species. To identify only those portions
that warrant further consideration, we determine whether there is
substantial information indicating that (1) the portions may be
significant and (2) the species may be in danger of extinction in those
portions or likely to become so within the foreseeable future. We
emphasize that answering these questions in the affirmative is not a
determination that the species is an endangered or a threatened species
throughout a significant portion of its range--rather, it is a step in
determining whether a more detailed analysis of the issue is required.
In practice, a key part of this analysis is whether the threats are
geographically concentrated in some way. If the threats to the species
are affecting it uniformly throughout its range, no portion is likely
to warrant further consideration. Moreover, if any concentration of
threats apply only to portions of the range that clearly do not meet
the biologically based definition of ``significant'' (i.e., the loss of
that portion clearly would not be expected to increase the
vulnerability to extinction of the entire species), those portions will
not warrant further consideration.
If we identify any portions that may be both (1) significant and
(2) endangered or threatened, we engage in a more detailed analysis to
determine whether these standards are indeed met. The identification of
an SPR does not create a presumption, prejudgment, or other
determination as to whether the species in that identified SPR is an
endangered or a threatened species. We must go through a separate
analysis to determine whether the species is an endangered or a
threatened species in the SPR. To determine whether a species is an
endangered or a threatened species throughout an SPR, we will use the
same standards and methodology that we use to determine if a species is
an endangered or a threatened species throughout its range.
Depending on the biology of the species, its range, and the threats
it faces, it may be more efficient to address the ``significant''
question first, or the status question first. Thus, if we determine
that a portion of the range is not ``significant,'' we do not need to
determine whether the species is an endangered or a threatened species
there; if we determine that the species is not an endangered or a
threatened species in a portion of its range, we do not need to
determine if that portion is ``significant.''
We evaluated the current range of the Upper Missouri River DPS of
Arctic grayling to determine if there is any apparent geographic
concentration of potential threats. We examined potential threats from
curtailment of range, dams, habitat fragmentation, dewatering and
thermal stress, entrainment, riparian habitat loss, sediment,
exploitation, disease and competition/predation, drought, climate
change, stochastic events, reduced genetic diversity, low abundance,
and lack of a fluvial ecotype replicate. The type and magnitude of
stressors acting on the Arctic grayling populations in the DPS are
varied.
Currently, nineteen of the twenty Arctic grayling populations in
the DPS are stable or increasing in abundance. Given this trend, we
conclude that there is no concentration of threats acting on these
nineteen populations because these populations are able to maintain
viability, despite some stressors acting at the individual level on
some of these populations. However, we acknowledge the probable
declining population trend in the Ennis Reservoir/Madison River
population. It is unclear what factor or
[[Page 49422]]
combination of factors is contributing to this decline. Nonnative trout
abundance is highest in the Madison River, relative to all other
systems occupied by nonnative trout and Arctic grayling, and this
factor may be contributing to the decline of Arctic grayling in Ennis
Reservoir/Madison River.
Given the probable decline of Arctic grayling in Ennis Reservoir/
Madison River, we analyzed the potential significance of this
population to the overall Upper Missouri River DPS of Arctic grayling.
To do this analysis, we evaluated whether the Ennis Reservoir/Madison
River population's contribution to the viability of the DPS is so
important that, without the members in this portion, the DPS would be
in danger of extinction, or likely to become so in the foreseeable
future, throughout all of its range. The Ennis Reservoir/Madison River
population occupies a small portion of the range within the DPS and
represents only 1 of 20 populations in the overall DPS. We conclude
that the DPS would still be viable if the Ennis Reservoir/Madison River
population were extirpated because adequate redundancy (3 other fluvial
or partially fluvial and 16 other adfluvial populations) of Arctic
grayling populations would still exist. In addition, representation of
resilient populations would remain in the Madison drainage (Grebe Lake
population) and rangewide in 7 of 10 historically occupied watersheds
in the Upper Missouri River basin. Further, resiliency of the DPS would
not be compromised by the loss of the Ennis Reservoir/Madison River
population because all remaining Arctic grayling populations are
widespread and viable. Therefore, in the hypothetical absence of the
Ennis Reservoir/Madison River population, the remainder of the Upper
Missouri River DPS of Arctic grayling would not meet the definition of
threatened or endangered under the Act. For the reasons stated above,
the Ennis Reservoir/Madison River population does not meet the
definition of ``significant'' for the purposes of this SPR analysis.
In conclusion, we find no concentration of stressors acting on
nineteen of twenty Arctic grayling populations in the DPS. The Ennis
Reservoir/Madison River population does appear to have a stressor or
combination of stressors acting at the population level. However,
further analysis indicates that the Ennis Reservoir/Madison River does
not meet the definition of ``significant'' in our SPR policy because
adequate redundancy, representation, and resiliency would still exist
within the DPS if the Ennis Reservoir/Madison River population were
extirpated. Thus, the remainder of the Upper Missouri River DPS of
Arctic grayling would not meet the definition of threatened or
endangered. Therefore, we find that there is not a significant portion
of the range of the Upper Missouri River DPS of Arctic grayling that
warrants listing.
Our review of the best available scientific and commercial
information indicates that the Upper Missouri River DPS of Arctic
grayling is not in danger of extinction (endangered), nor likely to
become endangered within the foreseeable future (threatened),
throughout all or a significant portion of its range. Therefore, we
find that listing the Upper Missouri River DPS of Arctic grayling as an
endangered or threatened species under the Act is not warranted at this
time.
We request that you submit any new information concerning the
status of, or threats to, the Upper Missouri River DPS of Arctic
grayling to our Montana Ecological Services Office (see ADDRESSES)
whenever it becomes available. New information will help us monitor the
Upper Missouri River DPS of Arctic grayling and encourage its
conservation. If an emergency situation develops for the Upper Missouri
River DPS of Arctic grayling, we will act to provide immediate
protection.
References Cited
A complete list of references cited is available on the Internet at
https://www.regulations.gov and upon request from the Montana Ecological
Services Office (see ADDRESSES).
Authors
The primary authors of this document are the staff members of the
Montana Ecological Services Office.
Authority
The authority for this section is section 4 of the Endangered
Species Act of 1973, as amended (16 U.S.C. 1531 et seq.).
Dated: August 6, 2014.
David Cottingham,
Acting Director, Fish and Wildlife Service.
[FR Doc. 2014-19353 Filed 8-19-14; 4:15 pm]
BILLING CODE 4310-55-P