Endangered and Threatened Wildlife and Plants; Revised 12-Month Finding for Upper Missouri River Distinct Population Segment of Fluvial Arctic Grayling, 20305-20314 [E7-7484]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife
and Plants; Revised 12-Month Finding
for Upper Missouri River Distinct
Population Segment of Fluvial Arctic
Grayling
Fish and Wildlife Service,
Interior.
ACTION: Notice of revised 12-month
finding.
AGENCY:
SUMMARY: We, the Fish and Wildlife
Service (Service), announce our revised
12-month finding on a petition to list
the upper Missouri River Distinct
Population Segment (DPS) of fluvial
Arctic grayling (Thymallus arcticus) as
threatened or endangered under the
Endangered Species Act of 1973, as
amended (Act). After a review of the
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best available scientific and commercial
information, we find that fluvial Arctic
grayling of the upper Missouri River
does not constitute a species,
subspecies, or distinct population
segment under the Act. Therefore, we
find that the petition to list the upper
Missouri River DPS of fluvial Arctic
grayling is not warranted, and we
withdraw the fluvial Arctic grayling
from the candidate list. The Service
continues to seek new information on
the taxonomy, biology, ecology, and
status of fluvial Arctic grayling and to
support cooperative conservation of
fluvial Arctic grayling in the upper
Missouri River system.
This finding was made on April
24, 2007.
DATES:
The complete file for this
finding is available for inspection, by
appointment, during normal business
hours, at the U.S. Fish and Wildlife
Service, Montana Field Office, 585
Shepard Way, Helena, MT 59601;
telephone (406) 449–5225. Submit new
information, materials, comments, or
questions concerning this species to us
at this address (Attention: Arctic
grayling).
ADDRESSES:
FOR FURTHER INFORMATION CONTACT:
Mark Wilson, Field Supervisor,
Montana Field Office, at the address and
telephone listed above.
SUPPLEMENTARY INFORMATION:
Species Information
Description
The Arctic grayling (Thymallus
arcticus) belongs to the family
Salmonidae (salmon, trout, charr,
whitefishes), subfamily Thymallinae
(graylings), and is represented by a
single genus, Thymallus, which
contains three other recognized species
in addition to T. arcticus (Scott and
Crossman 1973, pp. 301–302; Behnke
2002, pp. 327–331). Arctic grayling have
elongate, laterally compressed bodies
with deeply forked tails, and adults
typically average 254 to 330 millimeters
(10 to 13 inches) in length. Coloration
varies from silvery or iridescent blue
and lavender, to dark blue (Behnke
2002, pp. 327–328). During the
spawning period, the colors darken and
the males become more brilliantly
colored than the females. 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. Dark spots are often
evident on the body towards the head
(Behnke 2002, pp. 327–328).
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Distribution
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Arctic grayling have a primarily
holarctic distribution and are native to
Arctic Ocean drainages of northwestern
Canada and Alaska, from the Peace,
Saskatchewan, and Athabasca River
drainages in Alberta eastward to
Hudson Bay and westward to the Bering
Straits and eastern Siberia and northern
Eurasia (Scott and Crossman 1973, pp.
301–302). Arctic grayling also are native
to Pacific coast drainages of Alaska and
Canada as far south as the Stikine River
in British Columbia (Scott and
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Crossman 1973, pp. 301–302; Nelson
and Paetz 1991, pp. 253–256; Behnke
2002, pp. 327–331). Arctic grayling
generally occur throughout their native
range though the species is extirpated in
some locations (Michigan) and has
experienced local range contraction in
others (e.g., Peace-Willison watershed in
British Columbia (Blackman et al. (1990,
pp. 15, 17, 34), portions of Alberta
(Alberta Sustainable Resource
Development (2005; pp. iv, 5–18), and
Montana).
In North America, two populations of
Arctic grayling, believed to have been
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isolated by Pleistocene glaciations, have
been recorded outside of Canada and
Alaska (Vincent 1962, pp. 23–31). One
population was found in streams and
rivers of the Great Lakes region of
northern Michigan, but those grayling
were extirpated in the 1930s (Hubbs and
Lagler 1949, p. 44; Scott and Crossman
1973, p. 301). The second population
historically inhabited watersheds in the
upper Missouri River basin upstream of
Great Falls, Montana (Figure 1).
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Genetic data indicate Arctic grayling
native to the Missouri River system
were most likely isolated geographically
from Hudson Bay and Arctic Ocean
drainages by the onset of Wisconsin
glaciation approximately 70,000 years
ago (Redenbach and Taylor 1999, p. 32).
Arctic grayling native to the upper
Missouri River system are genetically
diverged from Arctic grayling in the
northern part of the species’ range
(Lynch and Vyse 1979, pp. 268–270,
275; Everett 1986, pp. 15–16, 79–80;
Redenbach and Taylor 1999, pp. 23, 28–
29, 32–33; reviewed by Leary 2005, pp.
1–3; reviewed by Campton 2006, pp. 5–
6), and appear to be most closely related
evolutionarily to populations in the
Fond du Lac area of northeastern
Saskatchewan, Canada (Stamford and
Taylor 2004, p. 1538). Genetic
divergence happens when two or more
genetic characteristics that have
occurred naturally over time are passed
from one generation to subsequent
generations. Arctic grayling in the upper
Missouri River basin are commonly
referred to as ‘‘Montana grayling’’ and
have been variously categorized as a
separate species (Thymallus montanas;
Scott and Crossman 1973, p. 301) or
subspecies (T. arcticus montanus;
Williams et al. 1989, p. 4), but these
designations are of uncertain validity
(Scott and Crossman 1973, p. 301) and
not widely accepted (Kaya 1990, pp. 3–
4; Integrated Taxonomic Information
System 2006). The lack of accepted
subspecific designations is based on
morphological similarity among
disjunct populations (Kaya 1990, p. 4).
Arctic grayling in the upper Missouri
River basin currently represent the
southern extent of the species’ range
(Scott and Crossman 1973, pp. 301–
302), and both migratory, river-dwelling
(fluvial) and lake-dwelling (adfluvial
and lacustrine) populations are native to
the upper Missouri River. For
simplicity, the term ‘‘adfluvial’’ will be
used to refer to all Arctic grayling
populations associated with lakes or
reservoirs. The migratory, stream- and
river-dwelling form of Arctic grayling
native to the upper Missouri River is
hereafter referred to as ‘‘fluvial’’ Arctic
grayling of the upper Missouri River.
Arctic Grayling Distribution in the
Upper Missouri River Basin
Fluvial Arctic grayling reside in the
Big Hole River and the lower reaches of
connected tributaries (see Figure 1
above). Adfluvial Arctic grayling native
to the upper Missouri River system are
known to reside in the Red Rock Lakes
system, in the upper reaches of the
Beaverhead River within the Centennial
Valley, Montana (Vincent 1962, p. 120;
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see Figure 1 above). An indigenous
Arctic grayling population exhibiting
adfluvial characteristics also is present
in the Madison River upstream from
Ennis Reservoir (see Figure 1 above).
The adfluvial characteristics expressed
by the Madison River-Ennis Reservoir
population may reflect recent
divergence away from the presumed
ancestral fluvial form resulting from the
construction of Ennis Dam (Kaya 1990,
p. 33; Kaya 1992a, p. 53). A few
adfluvial populations found in small
lakes within the Big Hole River system
(in particular Miner and Mussigbrod
Lakes; see Figure 1 above) may be
remnant native populations derived
from fluvial Arctic grayling from the Big
Hole River and isolated by recent
habitat fragmentation, but widespread
stocking of these and other locations
with hatchery-reared Arctic grayling
during the 1930s–1950s (e.g., Everett
1986, p. 4, 16; Kaya 1990, pp. 31, 75–
80) also makes it possible that these fish
are introduced populations or that the
existing populations are a mixture of
native and introduced Arctic grayling.
Ecology
Northcote (1995) and Kaya (1990)
reviewed the ecology of Arctic grayling
and fluvial Arctic grayling of the upper
Missouri River, respectively. Much of
the information on fluvial Arctic
grayling in the upper Missouri River
system comes from the Big Hole River,
Montana (see Figure 1 above), which
contains a fluvial population. Arctic
grayling exhibit life history and
migratory forms present in other species
of inland trout and charr, including
fluvial and adfluvial. Fluvial
populations are characterized by a cycle
of migratory behavior over their lifespan
between spawning, feeding, and
overwintering habitats within rivers or
streams (Northcote 1995, pp. 156–160).
Fluvial Arctic grayling typically migrate
upstream to spawn in tributary or
mainstem river locations and
downstream to overwintering habitats.
Such movement patterns have been
observed in fluvial Arctic grayling in
Big Hole River, Montana (Shepard and
Oswald 1989, pp. 18, 27–28). Migrations
to feeding habitats may occur if these
locations differ from spawning or
overwintering habitats (Kaya 1990, pp.
9–11). Overall, movements by fluvial
populations within and among
tributaries and mainstem rivers may
cover hundreds of kilometers
(Armstrong 1986, p. 7). Fluvial Arctic
grayling in the Big Hole River system
have been shown to migrate in excess of
80 km (50 mi) between spawning,
feeding and wintering areas (Shepard
and Oswald 1989, pp. 18, 21; Lamothe
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and Magee 2003, pp. 7, 11, 17).
Adfluvial Arctic grayling feed and
overwinter in lakes, but migrate to inlet
or outlet streams to spawn (Northcote
1995, p. 148–149; Northcote 1997, pp.
1030–1034).
Age at maturity and longevity in
Arctic grayling varies among systems
and is probably related to growth rate,
with populations in colder, less
productive habitats maturing at later
ages and having a greater lifespan
(Northcote 1995, pp. 155–157). Fluvial
Arctic grayling in the Big Hole River
system typically mature at 2 years of age
(males) or 3 years of age (females), and
individuals older than 6 years of age are
rare (Liknes 1981, pp. 16–18; Kaya 1990,
pp. 18–20; Magee and Lamothe 2003, p.
22). Arctic grayling are spring spawners.
In Montana, Arctic grayling typically
spawn from late April to mid-May by
depositing adhesive eggs over gravel
substrate without excavating a nest or
redd (Shepard and Oswald 1989, pp.
24–25, 29; Kaya 1990, pp. 15–16). In
general, the reproductive ecology of
Arctic grayling is somewhat different
from other salmonid species (trout and
salmon) in that Arctic grayling eggs tend
to be comparatively small (Behnke 2002,
p. 328), and males establish and defend
spawning territories rather than
defending access to females (Northcote
1995, p. 150). The time required for
development of eggs from embryo until
they emerge from stream gravel and
become swim-up fry varies with water
temperature, but averages about 3 weeks
for Arctic grayling in the upper Missouri
River basin (Kaya 1990, pp. 16–17).
Small, weakly swimming fry of fluvial
Arctic grayling prefer low velocity
stream habitats (Kaya 1990, pp. 23–24;
Northcote 1995, pp. 152–153).
Arctic grayling of all ages feed
primarily on aquatic and terrestrial
invertebrates captured on or near the
water surface (Northcote 1995, pp. 153–
154; Behnke 2002, p. 328). They also
will feed opportunistically on fish and
fish eggs (Northcote 1995, p. 154;
Behnke 2002, p. 328). Feeding locations
for individual fish are typically
established and maintained through
size-mediated dominance hierarchies
(e.g., Hughes 1992, pp. 1994–1995).
Although fluvial Arctic grayling may
have specific habitat requirements
depending on their life stage (e.g., fry)
and ecological activity (e.g., spawning),
individuals inhabiting streams and
rivers often exhibit a preference for pool
habitats (Liknes 1981, pp. 22, 28; Kaya
1990, pp. 20–21; Lamothe and Magee
2003, pp. 13–14, 17; Lamothe and
Magee 2004, p. 24). Vincent (1962, pp.
39, 42) concluded that fluvial Arctic
grayling in Montana typically reside in
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streams with low-to-moderate gradient
(<4 percent) and prefer low-to-moderate
water velocities (<60 centimeters/sec).
Observations of fluvial Arctic grayling
habitat use in the Big Hole River by
Liknes (1981, p. 28) and Liknes and
Gould (1987, p. 128) are consistent with
these generalizations.
Arctic grayling generally prefer cool
or coldwater habitats (Hubert et al.
1985, pp. 9, 14, 25, 27). Selong et al.
(2001, p. 1032) placed Arctic grayling in
a ‘‘coldwater’’ group of salmonids, along
with Arctic charr and bull trout, based
on critical thermal maximum values.
Genetic Relationships Among Arctic
Grayling Populations in the Upper
Missouri River Basin
Discussion of genetic divergence
among Arctic grayling populations is
complicated by the extensive hatchery
propagation and transplantation of
stocks from location to location (Everett
1986, p. 40). Over 10 million grayling of
unknown origin were stocked in the Big
Hole River over a 30-year period from
the 1930s to the 1950s (Kaya 1990, pp.
31, 75–80). Everett (1986 pp. 42, 43, 47)
concluded that the effect of grayling
introductions on local genetics appears
stronger in lake populations than in the
Big Hole River. Nonetheless, the limited
available genetic data suggest the
presence of two or more groups—
clusters or sets of populations that are
genetically more closely related to each
other than they are to other populations
of the same species—of Arctic grayling
within the upper Missouri River that
may not be strictly delineated by
geography and life history (Leary 2005,
p. 3; Campton 2006, pp. 6–9, 12).
Inferences about genetic differences
among Arctic grayling populations
within the upper Missouri River basin
are primarily based on data collected by
Everett (1986) and Leary (1990). These
two studies examined how a particular
form (allele) of a protein molecule
(allozyme) varied in frequency across
Arctic grayling populations in Montana.
Allozymes are gene products coded by
DNA, so allozyme variation can be used
to infer genetic relationships among
populations, subspecies or species.
Campton (2006, pp. 6, 12), in his review
of those data, suggested the existence of
two possible genetic groups: (a) A Big
Hole-Madison River group that includes
the fluvial population in the Big Hole
River, certain populations in adjoining
waters of the Big Hole River system
(e.g., Bobcat, Miner, and Mussigbrod
Lakes, and Steel Creek; see Figure 1
above; see Everett 1986, p. 7; Leary
1990, pp. 6–8), and fish from the
Madison River-Ennis Reservoir; and (b)
a Red Rock Lakes group that includes
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native adfluvial populations from the
Red Rock and Elk Lakes system in the
upper Beaverhead River system, and a
number of introduced adfluvial
populations (Agnes, Grebe, Rogers,
Odell, and Elizabeth Lakes; see Leary
1990, pp. 7–8) believed to be derived
from human introductions of Red Rock
Lakes grayling and/or associated
hatchery stocks. The two groups (Big
Hole-Madison and Red Rock Lakes) are
differentiated by divergent allele
frequencies for two allozymes (Campton
2006, p. 6). The relative genetic
difference between these two groups
within the upper Missouri River basin is
less than the difference between upper
Missouri River Arctic grayling and
sample populations from Alaska and
Canada (Everett 1986, p. 80; Leary 1990,
pp. 1, 7–8). The level of genetic
divergence observed among populations
within the upper Missouri River is
consistent with what would be expected
for populations within a geographic area
that share a recent ancestry but have
since diverged, as compared with the
greater divergence observed among
populations from different geographic
areas or river systems that have been
separated from each other for a much
longer period of time (i.e., upper
Missouri River versus Alaskan and
Canadian populations).
Campton (2006, p. 12) also noted that
a few adfluvial populations of Arctic
grayling in the Big Hole River drainage,
including Miner Lake (see Figure 1
above), appear to share recent ancestry
with the mainstem Big Hole River
fluvial population.
Like Campton, Leary also concluded
that Big Hole River and Madison River
grayling samples appear to be quite
similar (Leary 2005, p. 3). Leary’s
interpretation of the genetic
relationships among Miner Lake, Red
Rock Lakes, and Elk Lake populations
was different from Campton’s. Leary
found Miner Lake to be very divergent
from all the others, but also concluded
that there was significant divergence
between the Red Rock Lakes and Elk
Lake samples (Leary 2005, p. 3). He
interpreted the allozyme data to mean
that the adfluvial samples do not appear
to form a genetically distinct group and
consequently concluded that the data do
not support the premise that the fluvial
and adfluvial life histories fall into two
distinct genetic lineages (Leary 2005, p.
3). Rather, he contended the data
represent divergence among populations
regardless of life history (Leary 2005, p.
3). In his review, Campton (2006)
concurred that the apparent genetic
divergence between the two groups (Big
Hole-Madison River and Red Rock
Lakes) was not completely consistent
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with life histories because several
adfluvial populations belonged to the
Big Hole River-Madison River genetic
group.
An Arctic grayling population
residing in the Sunnyslope irrigation
canal in Teton County, Montana, is
thought to be derived from an
introduction into Pishkin Reservoir
(Kaya 1990, p. 41; see Figure 1 above)
and is not easily assigned to either of
the two genetic groups suggested by
Campton. These fish appear to be
genetic outliers relative to the two other
native genetic groups of Arctic grayling
(Leary 1990, p. 8; Campton 2006, p. 7).
Overall, both Campton and Leary
observe that: (a) Fluvial Arctic grayling
from the Big Hole River are genetically
different from native adfluvial Arctic
grayling in Red Rock Lakes based on
observed differences in allozyme allele
frequencies even if the genetic
divergence between these populations
appears to be low (average Nei’s genetic
distance of the cluster containing these
populations equals 0.0132 (Leary 1990,
pp. 1,8)); (b) the existing genetic data do
not strongly support the hypothesis that
the fluvial form of Arctic grayling in the
upper Missouri River represents a
unique genetic lineage, because it is
genetically similar to adfluvial
populations in Miner Lake and in the
Madison River (Leary 2005, pp. 3–4;
Campton 2006, p. 12); and (c) the low
allozyme variability in upper Missouri
River Arctic grayling samples results in
a weak dataset for resolving ancestries
among recently diverged populations
(Leary 2005, pp. 3–4; Campton 2006, p.
10). The Service views Campton’s and
Leary’s conclusions about the ancestral
relationships among Arctic grayling
populations in the upper Missouri River
as tentative, given the inherent
limitations of the existing genetic data.
However, it is the best available
scientific information at this time.
Further investigations with more
variable genetic markers, such as
microsatellite DNA, may clarify genetic
relationships (Campton 2006, pp. 10,
14).
Heritable, Behavioral Differences
Between Fluvial and Adfluvial Arctic
Grayling in the Upper Missouri River
Basin
Arctic grayling exhibit at least two life
histories in the upper Missouri River
system—a river-dwelling fluvial form
and a lake-dwelling adfluvial form. Life
history variation in salmonid fishes
(trout and salmon) may or may not be
related to genetic differentiation (e.g.,
Fausch and Young 1995, p. 365).
However, experiments designed to
determine whether behavioral
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differences were due to genetic or
environmental influences found that the
behavioral differences between fluvial
and adfluvial Arctic grayling in
Montana were heritable. In tests of
swimming behavior of young-of-year
Arctic grayling raised in common
conditions in captivity, progeny of
fluvial Big Hole River fish behaved
significantly differently, on average,
than adfluvial progeny from Red Rock
Lakes and Madison River-Ennis
Reservoir populations (Kaya 1989, 1991;
Kaya and Jeanes 1995). The Big Hole
River progeny exhibited a greater
tendency to hold position in flowing
water (Kaya and Jeanes 1995, pp. 453–
456). Because the test fish from the Big
Hole River population were progeny of
parents reared in a non-fluvial
environment, retention of this rheotactic
behavior (behavior in response to
flowing water) was taken as evidence
that such behavior has a genetic
(heritable) basis (Kaya and Jeanes 1995,
p. 456), consistent with conclusions of
previous investigations (Kaya 1989, pp.
474, 478–479; Kaya 1991, pp. 53, 55–
58).
Expression of rheotactic
characteristics in Arctic grayling also
can be influenced by ontogeny, or the
developmental history of an individual
(in this case, time from emergence from
gravel as fry until maturity; Kaya 1991,
pp. 53, 55–57), and environmental
conditions, such as time of day (Kaya
1989, p. 56), light intensity (Kaya 1989,
p. 478; Kaya 1991, p. 56), or water
temperature (Kaya 1989, p. 478).
However, the collective results are
nonetheless consistent with the
hypothesis that heritable, behavioral
differences in the test populations exist
between the fluvial and adfluvial
populations and those associated with
lakes or reservoirs.
Adfluvial Arctic grayling repeatedly
introduced into rivers have failed to
establish viable populations (Kaya
1992b, pp. 12–14). Adaptive divergence
and lack of ecological exchangeability
between life history types are among the
factors that may have contributed to
these failures (Campton 2006, p. 13).
However, introductions of fluvial
grayling into other rivers within the
native range have not been successful
either, so success may be due to other
factors (e.g., habitat degradation or
competition with nonnative fish (Kaya
1992b, pp. 10–12, 60)). In general, life
history expression in salmonid species
can be flexible, and Arctic grayling
exhibit variation in migratory behavior
across the range of the species
(Northcote 1997, p. 1030). Geography
may be a stronger determinant of
ancestral relationships than life history
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for Arctic grayling. Native Arctic
grayling populations within the upper
Missouri River basin may be similar
based on genetics, because they reside
in the same river basin and presumably
share a recent evolutionary ancestry
(Campton 2006, p. 12), while at the
same time expressing different life
histories in response to local habitat
conditions.
Previous Federal Action
The Service initiated a status review
for the Montana Arctic grayling
(Thymallus arcticus montanus) through
a notice of review published on
December 30, 1982 (47 FR 58454). In
that notice, Montana Arctic grayling was
designated a Category 2 species, which
included taxa for which information in
possession of the Service at that time
indicated that proposing to list the
species as Endangered or Threatened
was possibly appropriate, but for which
substantial data were not currently
available to biologically support a
proposed rule (47 FR 58454). We
received a petition, dated October 2,
1991, from the Biodiversity Legal
Foundation and George Wuerthner on
October 9, 1991. The petition requested
that the ‘‘fluvial Arctic grayling’’ be
listed as an endangered species
throughout its historic range ‘‘in the
conterminous United States.’’
We published a notice of a 90-day
finding in the January 19, 1993, Federal
Register (58 FR 4975). In that 90-day
finding we found that 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. We also found that
because the Michigan population of
Arctic grayling is extinct and, therefore,
by definition cannot be listed, the
finding would address only the fluvial
population of the Arctic grayling in the
upper Missouri River drainage.
On July 25, 1994, we published a
notice of a 12-month petition finding in
the Federal Register concluding that
listing the fluvial Arctic grayling
indigenous to the upper Missouri River
was warranted but precluded by other
higher priority listing actions (59 FR
37738). This finding stated that the
Service viewed adfluvial Arctic grayling
as not under consideration in the
Service’s finding as it was believed to be
a distinct population from the fluvial
Arctic grayling. This 1994 status review
identified the fluvial form of Arctic
grayling in the upper Missouri River
drainage as a DPS based on its
geographic isolation and behavioral
distinctiveness (59 FR 37738–37741,
July 25, 1994). This status review
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occurred prior to the finalization of the
Service and the National Marine
Fisheries Service’s joint DPS policy in
1996 (61 FR 4722, February 7, 1996).
Since 1994, and based on the best
available information and the
assessment that we conduct during our
candidate review process, we have
continued to preliminarily recognize the
fluvial Arctic grayling of the upper
Missouri River as a DPS, and has
maintained it as a candidate species
through the annual Candidate Notice of
Review. In 2004, the Service elevated
the listing priority number of the fluvial
Arctic grayling to 3 (69 FR 24881, May
4, 2004) because the abundance of the
remnant population in the Big Hole
River declined substantially and
reestablishment efforts had not yet
produced self-sustaining populations
elsewhere in the upper Missouri River.
On May 31, 2003, the Center for
Biological Diversity and Western
Watersheds Project (collectively
plaintiffs) filed a complaint in United
States District Court in Washington, DC
(1:03-cv-01110), challenging the
Service’s continuing ‘‘warranted but
precluded’’ determination for fluvial
Arctic grayling contained in the 2002
Candidate Notice of Review (67 FR
40657, June 13, 2002). Plaintiffs filed an
amended complaint on July 22, 2004,
challenging the Service’s failure to use
its emergency listing authority to protect
the fluvial Arctic grayling under the Act
(16 U.S.C. 1531 et seq.). The litigation
with plaintiffs was settled in August
2005. In this settlement agreement, the
Service agreed that on or before April
16, 2007, it shall submit for publication
in the Federal Register a final
determination made pursuant to the Act
as to whether or not the ‘‘Montana
fluvial Arctic grayling’’ is an
endangered or threatened species.
During the evaluation of the petition,
the Service considered the term
‘‘Montana fluvial Arctic grayling’’ as
synonymous with ‘‘fluvial Arctic
grayling of the upper Missouri River.’’
In this finding, as in the past, the fluvial
form of the indigenous Arctic grayling
from the upper Missouri River drainage
in Montana and Wyoming is referred to
as the fluvial Arctic grayling. This
revised 12-month finding is being
published as a final listing
determination in accordance with the
settlement agreement.
Distinct Vertebrate Population Segment
Pursuant to the Act, we must consider
for listing any species, subspecies, or,
for vertebrates, any DPS of these taxa if
there is sufficient information to
indicate that such action may be
warranted. The petition we received
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concerns a potential DPS of fluvial
Arctic grayling. Under our Policy
Regarding the Recognition of Distinct
Vertebrate Population Segments (61 FR
4722, February 7, 1996) (known as the
DPS Policy), three elements are
considered in a decision regarding the
status of a possible DPS as endangered
or threatened under the Act. These
factors are applied similarly for
additions to the Lists of Endangered and
Threatened Wildlife and Plants (Lists),
reclassification, and removal from the
Lists. They are: (1) Discreteness of the
population segment 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 (i.e., is
the population segment, when treated as
if it were a species, endangered or
threatened?). Discreteness refers to the
isolation of a population from other
members of the species, and we evaluate
this based on specific criteria that are
also contained in the DPS Policy and are
listed below. If the population segment
is determined to be discrete, then we
evaluate significance by using the
available scientific information to
determine the population segment’s
importance to the taxon to which it
belongs. If we determine that a
population segment is discrete and
significant, we subsequently evaluate it
for endangered or threatened status
based on the Act’s standards.
Discreteness
Under our DPS Policy, a population
segment of a vertebrate species may be
considered discrete if it satisfies either
one of the following conditions: (1) It is
markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors.
Quantitative measures of genetic or
morphological discontinuity may
provide evidence of this separation; or
(2) It is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the Act.
The subject of this DPS evaluation is
the fluvial Arctic grayling of the upper
Missouri River. In response to a
petition, the fluvial Arctic grayling was
the subject of a status review by the
Service in 1994, which identified Arctic
grayling indigenous to the Big Hole and
Madison Rivers as elements of a fluvial
DPS in the upper Missouri River (59 FR
37738–37741, July 25, 1994). However,
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this status review occurred prior to the
finalization of the Service and the
National Marine Fisheries Service’s
joint DPS policy in 1996 (61 FR 4722,
February 7, 1996). Since 1994, and most
recently in 2004 and 2005, the Service
reviewed the available information
concerning the taxonomic status of the
species in relation to the DPS policy and
again preliminarily determined that the
fluvial Arctic grayling of the upper
Missouri River was a valid DPS (Service
2004, 2005). This DPS evaluation
considers the information used in the
previous assessments as well as a
solicited review (Campton 2006) and
unsolicited review (Leary 2005) of the
available genetic data for Arctic grayling
in Montana.
(1) Fluvial Arctic Grayling Are Discrete
as a Consequence of Physical Features
Fluvial arctic grayling native to the
upper Missouri River are ‘‘markedly
separated’’ from other grayling, both
those in Canada and Alaska, and from
the adfluvial form in the Missouri River
drainage because of physical and
reproductive isolation. Fluvial actic
grayling are geographically disjunct and
reproductively isolated from
populations inhabiting Arctic Ocean
and Hudson Bay drainages in Canada
and Alaska (Scott and Crossman 1973,
p. 301). Arctic grayling in the upper
Missouri River are reproductively
isolated from their nearest conspecifics
by at least 800 kilometers (km) (500
miles (mi)) (Nelson and Paetz 1991, p.
255) and have been separated from
Arctic Ocean populations for perhaps
70,000 years as a result of glacial
activity (Lynch and Vyse 1979, p. 263;
Redenbach and Taylor 1999, p. 32). This
long period of reproductive isolation
coupled with genetic drift and
environmental selection pressures has
resulted in genetic differences between
Arctic grayling from the Missouri River
and elsewhere based on analyses of
allozymes and mitochondrial DNA
(Lynch and Vyse 1979, pp. 263, 268,
275; Everett and Allendorf 1985, pp. 22–
23, 26; Everett 1986, pp. 79–80;
Redenbach and Taylor 1999, p. 23;
reviewed by Campton 2006, pp. 5–6;
reviewed by Leary 2005, pp. 1–3).
Fluvial and adfluvial Arctic grayling
within the upper Missouri River basin
are ‘‘markedly separated’’ from each
other as a result of physical features.
The fluvial form was once widespread
in the upper Missouri River basin, but
the adfluvial form was native only to the
Red Rocks Lakes and possible Elk Lake
in the headwaters of the Beaverhead
River (Kaya 1990). Extant populations of
native fluvial and adfluvial Arctic
grayling within the upper Missouri
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River are reproductively isolated, and
the available genetic data are consistent
with the hypothesis of two genetic
groups of Arctic grayling (the Big Hole—
Madison River and Red Rock Lakes
genetic groups) within the upper
Missouri River (Leary 2005, p. 3;
Campton 2006, pp. 6–9, 12)
(2) Fluvial Arctic Grayling Are Not
Discrete as a Consequence of
Physiological Features
We do not believe that fluvial Arctic
grayling are discrete because of unique
or different physiological
characteristics. Lohr et al. (1996)
examined the thermal tolerance of
juvenile fluvial Arctic grayling from the
Big Hole River to elevated temperatures
in laboratory tests. However, grayling
from the Big Hole River did not appear
to be more tolerant of warm stream
temperatures than grayling from Alaska
(Lohr et al. 1996, p. 937).
Arctic grayling from the upper
Missouri River tend to grow more
quickly than individuals from northern
populations (Northcote 1995, pp. 156–
157). However, experimental data are
lacking that permit these differences to
be attributed to environmental versus
genetic influences.
(3) Fluvial Arctic Grayling Are Not
Discrete as a Consequence of Ecological
Features
The Arctic grayling of the upper
Missouri River represent the only
natural example of the taxon inhabiting
an Atlantic Ocean drainage (via the
Missouri and Mississippi Rivers and
Gulf of Mexico). All other wild
populations of Arctic grayling inhabit
drainages of the Arctic Ocean, Hudson
Bay, or north Pacific Ocean (USFWS
2005, p. 10). However, fluvial Arctic
grayling of the upper Missouri River
basin are not discrete from adfluvial
Arctic grayling of the upper Missouri
River basin as a consequence of
ecological features as they exist within
a common drainage.
(4) Fluvial Arctic Grayling Are Discrete
as a Consequence of Behavioral
Features
Under historical conditions within
the upper Missouri River basin, native
fluvial and adfluvial populations of
Arctic grayling spawned in different
locations (Vincent 1962, pp. 98–121;
Kaya 1990, pp. 24–30; Kaya 1992a, pp.
47–53). Homing behavior to natal (birth)
habitats that is typically expressed by
Arctic grayling (e.g., Carl et al. 1992, p.
245) would presumably result in the
reproductive isolation of historical
fluvial and adfluvial populations even if
occasional exchange was possible. In
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addition, genetic differences between
the extant fluvial population in the Big
Hole River and the native adfluvial
population in Red Rock Lakes (e.g.,
Everett 1986, pp. 79–30; Leary 1990, pp.
7–8) are consistent with reproductive
isolation between those populations
based on observed differences in
allozyme allele frequencies.
Fluvial and adfluvial Arctic grayling
do not appear to represent distinct
lineages based strictly on life histories
within the upper Missouri River system
(e.g., Leary 2005, p. 3; Campton 2006, p.
12); there are clearly some heritable
differences in juvenile swimming
behavior among fluvial Arctic grayling
and the native adfluvial populations in
terms of rheotactic response to flowing
water (Kaya 1989, pp. 474, 478–479;
Kaya 1991, pp. 53, 55–58; Kaya and
Jeanes 1995, pp. 453–456). These
differences in behavior are sufficient to
satisfy the discreteness criterion of the
DPS policy.
On the basis of the available
information, we conclude that the
fluvial Arctic grayling of the upper
Missouri River drainage is discrete from
other populations of the same taxon as
a consequence of physical and
behavioral factors. Since a population
segment of a vertebrate species may be
considered discrete if the first factor is
met (marked separateness), we need not
address the second factor (delimitation
by an international boundary).
Therefore, we considered the potential
significance of this discrete population
to the remainder of the taxon.
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Significance
If a population segment is determined
to be discrete, the Service considers the
available scientific evidence of its
significance to the taxon to which it
belongs. Our policy states that this
consideration may include, but is not
limited to, the following:
(1) Persistence of the discrete
population segment in an ecological
setting unusual or unique for the taxon;
(2) Evidence that loss of the discrete
population segment would result in a
significant gap in the range of the 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
historic 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 criteria to be
considered significant. Furthermore, the
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list of criteria is not exhaustive; other
criteria may be used, as appropriate.
(1) Fluvial Arctic Grayling Do Not
Persist in an Ecological Setting Unusual
or Unique for the Taxon
As discussed above, Arctic grayling
generally occur throughout their native
range in the holarctic region of Canada
and Alaska to eastern Siberia and
northern Eurasia (Scott and Crossman
1973, pp. 301–302). In our 2005
candidate assessment, we asserted that
the fluvial Arctic grayling of the upper
Missouri River persist in an ecological
setting unusual or unique for the taxon
as they represent the only natural
example of the taxon inhabiting an
Atlantic Ocean drainage via the
Missouri and Mississippi Rivers and
Gulf of Mexico. We noted that all other
wild populations of Arctic grayling
inhabit drainages of the Arctic Ocean,
Hudson Bay, or north Pacific Ocean
(USFWS 2005, p. 10). However, as
established above, we now note that
adfluvial Arctic grayling also persist in
the upper Missouri River drainage. Our
prior finding did not take these fish into
account in its discussion of ecological
setting. Because both the fluvial and
adfluvial forms are found in the upper
Missouri drainage, we cannot find that
the population persists in an ecological
setting unique or unusual to the taxon
as a whole.
Further, existence of the species in a
different drainage, or different rivers
and lakes, from those grayling found in
Canada and Alaska is not necessarily
evidence of a unique ecological setting.
Arctic grayling in the neararctic region
are found in the same habitat type as
those in Montana. Grayling inhabit clear
water streams, rivers, and lakes.
Riverine populations depend on large
streams, deep pools of small streams, or
spring-fed reaches that are not
completely frozen in winter for
overwinter survival. Populations not
associated with lakes are found in both
Alaska and Montana (Hubert 1985, p. 1).
For this reason also, we find that fluvial
Arctic grayling do not persist in an
ecological setting unique or unusual for
the taxon.
(2) The Loss of the Fluvial Arctic
Grayling Would Not Result in a
Significant Gap in the Range of the
Taxon
Loss of the fluvial Arctic grayling in
the upper Missouri River, when
considered in relation to grayling
throughout the remainder of the nearctic
region, would mean the loss of a small
percentage of the entire range of the
taxon. Due to the broad geographic
range of Arctic grayling, the gap in the
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range of Arctic grayling resulting from
the loss of fluvial Arctic grayling in the
upper Missouri River basin would not
result in a significant gap in the range
of the taxon as a whole.
In our 2005 candidate assessment, we
asserted that the loss of the fluvial
Arctic grayling of the upper Missouri
River would result in a significant gap
in the range of the taxon as these fish
are the only extant fluvial grayling
population in the contiguous United
States and represent the southernmost
extent of the species (USFWS 2005, p.
10). However, the Ninth Circuit Court
has rejected this argument as a
misconstruction of this criterion in the
case of National Association of Home
Builders v. Norton, 340 F. 3d 835, 852
(9th Cir. 2003) concerning the cactus
ferruginous pygmy-owl (Glaucidium
brasilianum cactorum) (70 FR 44551,
August 3, 2005). The Court found that
in designating a DPS under the DPS
policy, we must find that a discrete
population is significant to the taxon as
a whole, not to the United States.
Therefore, we have determined, based
on the information available to the
Service, the loss of the fluvial Arctic
grayling in the upper Missouri River
would not result in a significant gap in
the range of the species on the basis of
the significance of the Montana
population to the species as a whole.
(3) Fluvial Arctic Grayling Do Not
Represent the Only Surviving Natural
Occurrence of the Taxon
This criterion from the DPS policy
does not apply to the fluvial Arctic
grayling in the upper Missouri River
because it is clearly not a population
segment representing the only surviving
natural occurrence of a taxon that may
be more abundant elsewhere as an
introduced population outside its
historic range. Consequently, this
population of grayling is not significant
according to this standard.
(4) Fluvial Arctic Grayling in the
Missouri River Drainage Do Not Differ
Markedly in Genetic Characteristics
From Adfluvial Populations in the
Missouri River Drainage
As noted above, analyses of allozymes
and mitochondrial DNA show genetic
divergence between Arctic grayling in
the upper Missouri River and Arctic
grayling in Canada and Alaska (Lynch
and Vyse 1979, pp. 263, 268, 275;
Everett and Allendorf 1985, pp. 22–23,
26; Everett 1986, pp. 79–80; Redenbach
and Taylor 1999, p. 23; reviewed by
Campton 2006, pp. 5–6; reviewed by
Leary 2005, pp. 1–3) and appear to be
most closely related evolutionarily to
populations in northeastern
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Saskatchewan, Canada (Stamford and
Taylor 2004, p. 1538).
In addition, fluvial Arctic grayling
from the Big Hole River are genetically
different from native adfluvial Arctic
grayling in Red Rock Lakes based on
observed differences in allozyme allele
frequencies (Campton 2006, p. 6).
However, the relative genetic difference
between these two groups within the
upper Missouri River basin is less than
that between upper Missouri River
Arctic grayling and sample populations
from Alaska and Canada (Leary 1990,
pp. 1, 7–8).
Resolving ancestries among recently
diverged upper Missouri River Arctic
grayling populations is difficult due to
the low allozyme variability among
samples (Leary 2005, pp. 3–4; Campton
2006, p. 10). In this case, although
allozyme data from 39 loci are available
from these populations, only 2 of the
loci analyzed were generally variable
among them (Everett 1986; Leary 1990;
Leary 2005, p. 3). Information from only
two loci may cause chance similarities
or differences and require cautious
interpretation (Leary 2005, p. 3).
Likewise, the paucity of genetic
variation detected by Redenbach and
Taylor (1999, p. 27) in their restriction
enzyme analysis of mtDNA of upper
Missouri River basin Arctic grayling
precludes making any inferences about
genetic similarities or differences among
the upper Missouri River populations
sampled except that they all appear to
share a common maternal lineage (Leary
2005, p. 4). The level of genetic
divergence observed among populations
within the upper Missouri River is
consistent with what would be expected
for populations within a geographic area
that share a recent ancestry (Campton
2006, p. 12).
Discerning genetic divergence among
Arctic grayling populations is further
complicated by the extensive hatchery
propagation and transplantation of
stocks, as discussed above (Everett 1986,
p. 40). The Service does not regard the
introduced, lake-dwelling grayling to be
part of the indigenous upper Missouri
River fluvial Arctic grayling population
(59 FR 37739, July 25, 1994). However,
widespread stocking of hatchery-reared
Arctic grayling in the Big Hole River
system and other locations (e.g., Everett
1986, pp. 4, 16; Kaya 1990, pp. 31, 75–
80) makes it possible that some fish are
introduced populations or that the
existing populations are a mixture of
native and introduced Arctic grayling.
We find that, based on the genetic
information currently available, the
fluvial Arctic grayling of the upper
Missouri River drainage do not differ
markedly from adfluvial populations of
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the species in their genetic
characteristics such that they should be
considered biologically or ecologically
significant based simply on genetic
characteristics. Biological and ecological
significance under the DPS policy is
always considered in light of
Congressional guidance (see Senate
Report 151, 96th Congress, 1st Session)
that the authority to list DPSs be used
‘‘sparingly’’ while encouraging the
conservation of genetic diversity.
Conclusion on DPS
Under section 3 of the Act and our
implementing regulations at 50 CFR
424.02, a ‘‘species’’ is defined to include
any species or subspecies of fish,
wildlife, or plant, and any distinct
population segment of any vertebrate
species which interbreeds when mature.
Our implementing regulations provide
further guidance on determining
whether a particular taxon or
population is a species or subspecies for
the purposes of the Act: ‘‘The Secretary
shall rely on standard taxonomic
distinctions and the biological expertise
of the Department and the scientific
community concerning the relevant
taxonomic group’’ (50 CFR 424.11). As
noted above, Arctic grayling in the
upper Missouri River basin have been
classified into separate species and
subspecies, but these designations are
not widely accepted. Therefore, we do
not consider the subject of this petition
to constitute a distinct species or
subspecies.
The 1994 status review identified the
fluvial form of Arctic grayling in the
upper Missouri River drainage as a DPS
based on its geographic isolation and
behavioral distinctiveness (59 FR 37738,
July 25, 1994). On the basis of the best
available information, we continue to
conclude that the fluvial Arctic grayling
of the upper Missouri River drainage is
‘‘markedly separated’’ from all other
populations of the same taxon as a
consequence of physical and behavioral
factors. Consequently, the Service
concludes that the petitioned entity is
discrete according to the 1996 DPS
policy. However, on the basis of the four
significance criteria in the 1996 DPS
Policy, the Service is unable to conclude
at this time that the petitioned entity is
significant. Therefore, we find that the
fluvial Arctic grayling of the upper
Missouri River does not qualify as a
distinct population segment under the
Act.
Significant Portion of the Range
Pursuant to the Act and our
implementing regulations, a species
may warrant listing if it is threatened or
endangered in a significant portion of its
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20313
range. However, the petition did not
request that we determine whether the
grayling was threatened or endangered
in a significant portion of its range.
Rather, it asked that we list the fluvial
Arctic grayling in the U.S. as an
endangered species. Consistent with the
petition, our previous petition findings
have uniformly addressed possible
listing in the context of whether the
fluvial Arctic grayling in Montana
constitutes a DPS, and therefore a
‘‘species’’ under the Act. As discussed
above, we have now determined that the
fluvial Arctic grayling is not a DPS.
Thus, we have disposed of the question
raised by the petition: we have no
obligation under the Act to address the
separate question of whether the fluvial
Arctic grayling in Montana constitutes a
significant portion of the range of some
of the entire grayling species, or some
valid but currently undefined DPS. If
the Service determines in the future that
the grayling is threatened or endangered
in a significant portion of its range, we
will add the species to the candidate list
and propose its listing. However, that
would be a future action. Because the
petition and our prior finding were with
respect to a DPS, and we have found
that there is not a valid DPS, we do not
need to address significant portion of
the range at this time.
Finding
On the basis of the discussion
presented in this document, we find
that the fluvial Arctic grayling of the
upper Missouri River does not qualify as
a distinct population segment. As a
result, we find that the petition to list
the fluvial Arctic grayling of the upper
Missouri River is not warranted. Based
on this determination, we withdraw the
fluvial Arctic grayling of the upper
Missouri River from the candidate list.
Although no further action will result
from this finding, we request that you
submit new information concerning the
taxonomy, biology, ecology, and status
of the Arctic grayling of the upper
Missouri River system to the Montana
Field Office (see ADDRESSES below)
whenever it becomes available. We will
accept additional information and
comments from all concerned
governmental agencies, the scientific
community, industry, or any other
interested party concerning this finding;
and will reconsider this determination
in the event of new information as
appropriate. The Service continues to
strongly encourage cooperative
conservation and restoration of fluvial
Arctic grayling in the upper Missouri
River.
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References
A complete list of all references cited
herein is available upon request from
the Montana Field Office, U.S. Fish and
Wildlife Service (see ADDRESSES).
Author
The authors of this finding are
biologists in Region 6 of the U.S. Fish
and Wildlife Service.
Authority
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Dated: April 13, 2007.
H. Dale Hall,
Director, Fish and Wildlife Service.
[FR Doc. E7–7484 Filed 4–23–07; 8:45 am]
BILLING CODE 4310–55–P
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
Administration
50 CFR Part 648
[Docket No. 070409081–7081–01; I.D.
032907A]
RIN 0648–AS22
Magnuson-Stevens Fishery
Conservation and Management Act
Provisions; Fisheries of the
Northeastern United States; Summer
Flounder, Scup, and Black Sea Bass
Fishery Management Plan;
Amendment 14
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce.
ACTION: Proposed rule; request for
comments.
jlentini on PROD1PC65 with PROPOSAL
AGENCY:
SUMMARY: NMFS proposes regulations to
implement Amendment 14 to the
Summer Flounder, Scup, and Black Sea
Bass Fishery Management Plan (FMP)
developed by the Mid-Atlantic Fishery
Management Council (Council). The
proposed measures include a plan to
rebuild the scup stock from an
overfished condition to the level
associated with maximum sustainable
yield, as required by the MagnusonStevens Fishery Conservation and
Management Act (Magnuson-Stevens
Act). This action also proposes to allow
the regulations concerning the Gear
Restricted Areas (GRAs) to be modified
through framework adjustments to the
FMP. The intended effect of this change
would improve the timing of developing
and implementing modifications to the
GRAs.
VerDate Aug<31>2005
18:31 Apr 23, 2007
Jkt 211001
Comments must be received by
5 p.m. local time, on May 24, 2007.
ADDRESSES: You may submit comments
by any of the following methods:
• E-mail: FSBAmendment14Proposed
Rule@noaa.gov. Include in the subject
line the following identifier:
‘‘Comments on Amendment 14
Proposed Rule (Scup Rebuilding Plan).’’
• Federal e-rulemaking portal: http:/
www.regulations.gov
• Mail: Patricia A. Kurkul, Regional
Administrator, NMFS, Northeast
Regional Office, One Blackburn Drive,
Gloucester, MA 01930. Mark the outside
of the envelope: ‘‘Comments on
Amendment 14 Proposed Rule (Scup
Rebuilding Plan).’’
• Fax: (978) 281–9135
Copies of Amendment 14 and of the
draft Environmental Assessment,
preliminary Regulatory Impact Review,
and Initial Regulatory Flexibility
Analysis (EA/RIR/IRFA) are available
from Daniel T. Furlong, Executive
Director, Mid-Atlantic Fishery
Management Council, Room 2115,
Federal Building, 300 South New Street,
Dover, DE 19901–6790. The EA/RIR/
IRFA is also accessible via the Internet
at https://www.nero.noaa.gov.
FOR FURTHER INFORMATION CONTACT:
Michael P. Ruccio, Fishery Policy
Analyst, (978) 281–9104.
SUPPLEMENTARY INFORMATION: On August
18, 2005, NMFS notified the Council
that the scup (Stenotomus chrysops)
stock had been designated as overfished
and that, within 1 year of that notice, an
amendment or proposed regulations for
the scup fishery to end overfishing and
to rebuild the stock must be prepared in
accordance with the Magnuson-Stevens
Act. In response, the Council has
developed, and submitted for Secretarial
review, Amendment 14 to propose two
actions: (1) A 7–year plan to rebuild the
scup stock from an overfished condition
to a level associated with maximum
sustained yield (Bmsy), as required by the
Magnuson-Stevens Act; and (2) an
administrative change to the regulations
on framework adjustments.
DATES:
Background
The scup stock was determined to be
overfished in 1998 when the
Sustainable Fisheries Act (SFA)
amendments to the Magnuson-Stevens
Act were implemented. The Council
developed and proposed Amendment
12 (64 FR 16891, April 7, 1999) to
rebuild the scup stock in accordance
with the provisions outlined in the SFA.
The Council proposed in Amendment
12 that the management measures in
place to rebuild the scup fishery,
established by Amendment 8, were
PO 00000
Frm 00026
Fmt 4702
Sfmt 4702
adequate under SFA guidelines. NMFS
disagreed, and the rebuilding plan
proposed in Amendment 12 was
disapproved on April 28, 1999.
Following the disapproval, the
management measures previously
implemented by Amendment 8
remained in place for the scup fishery.
In years subsequent to the disapproval
of Amendment 12, the scup stock
exhibited signs of recovery. The
Northeast Fisheries Science Center
(NEFSC) spring survey index 3–year
average value for 2001–2003 indicated
that scup spawning stock biomass (SSB)
had increased to 3.31 kg/tow, above the
minimum biomass threshold (1/2 Bmsy)
of 2.77 kg/tow. The scup stock was no
longer considered overfished, although
the 35th Stock Assessment Review
Committee (SARC 35) indicated that the
status of the stock with respect to
overfishing could not be evaluated.
Although the condition of the scup
stock was improving, the stock had not
yet been rebuilt, as required by the
Magnuson-Stevens Act, to the Bmsy
proxy rebuilding target of 5.54 kg/tow.
In 2005, the NEFSC 3–year SSB index
value decreased to 0.69 kg/tow,
indicating that the stock was again
below the minimum biomass threshold
(1/2 Bmsy) and considered overfished.
NMFS formally notified the Council of
the overfished status of the scup stock,
thus initiating the Magnuson-Stevens
Act requirement that the Council
develop regulations or an amendment to
the FMP to rebuild the scup stock to the
Bmsy proxy level. The rebuilding plan
implemented by such regulations or
amendment must achieve the rebuilding
target within 10 years to comply with
the Magnuson-Stevens Act. In response,
the Council has developed, and
submitted for Secretarial review,
Amendment 14.
Proposed Scup Rebuilding Plan
Under Amendment 14, a constant
fishing mortality rate (F) of 0.10 would
be applied each year during a 7–year
rebuilding time period. Under this
approach, the NEFSC 3–year SSB index
value for the rebuilding period ending
December 31, 2014, is projected to be
5.96 kg/tow, approximately 8 percent
above the Bmsy proxy rebuilding target
(5.54 kg/tow).
Applying a constant F=0.10 for 7
years is projected to achieve the
required stock rebuilding to comply
with the Magunuson-Stevens Act;
however, because scup is a relatively
data poor stock and uncertainty exists
around estimates of fishing mortality,
stock size, and discards, Amendment 14
contains additional criteria to be
E:\FR\FM\24APP1.SGM
24APP1
Agencies
[Federal Register Volume 72, Number 78 (Tuesday, April 24, 2007)]
[Proposed Rules]
[Pages 20305-20314]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: E7-7484]
=======================================================================
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Revised 12-Month
Finding for Upper Missouri River Distinct Population Segment of Fluvial
Arctic Grayling
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of revised 12-month finding.
-----------------------------------------------------------------------
SUMMARY: We, the Fish and Wildlife Service (Service), announce our
revised 12-month finding on a petition to list the upper Missouri River
Distinct Population Segment (DPS) of fluvial Arctic grayling (Thymallus
arcticus) as threatened or endangered under the Endangered Species Act
of 1973, as amended (Act). After a review of the best available
scientific and commercial information, we find that fluvial Arctic
grayling of the upper Missouri River does not constitute a species,
subspecies, or distinct population segment under the Act. Therefore, we
find that the petition to list the upper Missouri River DPS of fluvial
Arctic grayling is not warranted, and we withdraw the fluvial Arctic
grayling from the candidate list. The Service continues to seek new
information on the taxonomy, biology, ecology, and status of fluvial
Arctic grayling and to support cooperative conservation of fluvial
Arctic grayling in the upper Missouri River system.
DATES: This finding was made on April 24, 2007.
ADDRESSES: The complete file for this finding is available for
inspection, by appointment, during normal business hours, at the U.S.
Fish and Wildlife Service, Montana Field Office, 585 Shepard Way,
Helena, MT 59601; telephone (406) 449-5225. Submit new information,
materials, comments, or questions concerning this species to us at this
address (Attention: Arctic grayling).
FOR FURTHER INFORMATION CONTACT: Mark Wilson, Field Supervisor, Montana
Field Office, at the address and telephone listed above.
SUPPLEMENTARY INFORMATION:
Species Information
Description
The Arctic grayling (Thymallus arcticus) belongs to the family
Salmonidae (salmon, trout, charr, whitefishes), subfamily Thymallinae
(graylings), and is represented by a single genus, Thymallus, which
contains three other recognized species in addition to T. arcticus
(Scott and Crossman 1973, pp. 301-302; Behnke 2002, pp. 327-331).
Arctic grayling have elongate, laterally compressed bodies with deeply
forked tails, and adults typically average 254 to 330 millimeters (10
to 13 inches) in length. Coloration varies from silvery or iridescent
blue and lavender, to dark blue (Behnke 2002, pp. 327-328). During the
spawning period, the colors darken and the males become more
brilliantly colored than the females. 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. Dark spots are often evident on the body towards
the head (Behnke 2002, pp. 327-328).
[[Page 20306]]
Distribution
Arctic grayling have a primarily holarctic distribution and are
native to Arctic Ocean drainages of northwestern Canada and Alaska,
from the Peace, Saskatchewan, and Athabasca River drainages in Alberta
eastward to Hudson Bay and westward to the Bering Straits and eastern
Siberia and northern Eurasia (Scott and Crossman 1973, pp. 301-302).
Arctic grayling also are native to Pacific coast drainages of Alaska
and Canada as far south as the Stikine River in British Columbia (Scott
and Crossman 1973, pp. 301-302; Nelson and Paetz 1991, pp. 253-256;
Behnke 2002, pp. 327-331). Arctic grayling generally occur throughout
their native range though the species is extirpated in some locations
(Michigan) and has experienced local range contraction in others (e.g.,
Peace-Willison watershed in British Columbia (Blackman et al. (1990,
pp. 15, 17, 34), portions of Alberta (Alberta Sustainable Resource
Development (2005; pp. iv, 5-18), and Montana).
In North America, two populations of Arctic grayling, believed to
have been isolated by Pleistocene glaciations, have been recorded
outside of Canada and Alaska (Vincent 1962, pp. 23-31). One population
was found in streams and rivers of the Great Lakes region of northern
Michigan, but those grayling were extirpated in the 1930s (Hubbs and
Lagler 1949, p. 44; Scott and Crossman 1973, p. 301). The second
population historically inhabited watersheds in the upper Missouri
River basin upstream of Great Falls, Montana (Figure 1).
BILLING CODE 4310-55-P
[[Page 20307]]
[GRAPHIC] [TIFF OMITTED] TP24AP07.000
[[Page 20308]]
Genetic data indicate Arctic grayling native to the Missouri River
system were most likely isolated geographically from Hudson Bay and
Arctic Ocean drainages by the onset of Wisconsin glaciation
approximately 70,000 years ago (Redenbach and Taylor 1999, p. 32).
Arctic grayling native to the upper Missouri River system are
genetically diverged from Arctic grayling in the northern part of the
species' range (Lynch and Vyse 1979, pp. 268-270, 275; Everett 1986,
pp. 15-16, 79-80; Redenbach and Taylor 1999, pp. 23, 28-29, 32-33;
reviewed by Leary 2005, pp. 1-3; reviewed by Campton 2006, pp. 5-6),
and appear to be most closely related evolutionarily to populations in
the Fond du Lac area of northeastern Saskatchewan, Canada (Stamford and
Taylor 2004, p. 1538). Genetic divergence happens when two or more
genetic characteristics that have occurred naturally over time are
passed from one generation to subsequent generations. Arctic grayling
in the upper Missouri River basin are commonly referred to as ``Montana
grayling'' and have been variously categorized as a separate species
(Thymallus montanas; Scott and Crossman 1973, p. 301) or subspecies (T.
arcticus montanus; Williams et al. 1989, p. 4), but these designations
are of uncertain validity (Scott and Crossman 1973, p. 301) and not
widely accepted (Kaya 1990, pp. 3-4; Integrated Taxonomic Information
System 2006). The lack of accepted subspecific designations is based on
morphological similarity among disjunct populations (Kaya 1990, p. 4).
Arctic grayling in the upper Missouri River basin currently
represent the southern extent of the species' range (Scott and Crossman
1973, pp. 301-302), and both migratory, river-dwelling (fluvial) and
lake-dwelling (adfluvial and lacustrine) populations are native to the
upper Missouri River. For simplicity, the term ``adfluvial'' will be
used to refer to all Arctic grayling populations associated with lakes
or reservoirs. The migratory, stream- and river-dwelling form of Arctic
grayling native to the upper Missouri River is hereafter referred to as
``fluvial'' Arctic grayling of the upper Missouri River.
Arctic Grayling Distribution in the Upper Missouri River Basin
Fluvial Arctic grayling reside in the Big Hole River and the lower
reaches of connected tributaries (see Figure 1 above). Adfluvial Arctic
grayling native to the upper Missouri River system are known to reside
in the Red Rock Lakes system, in the upper reaches of the Beaverhead
River within the Centennial Valley, Montana (Vincent 1962, p. 120; see
Figure 1 above). An indigenous Arctic grayling population exhibiting
adfluvial characteristics also is present in the Madison River upstream
from Ennis Reservoir (see Figure 1 above). The adfluvial
characteristics expressed by the Madison River-Ennis Reservoir
population may reflect recent divergence away from the presumed
ancestral fluvial form resulting from the construction of Ennis Dam
(Kaya 1990, p. 33; Kaya 1992a, p. 53). A few adfluvial populations
found in small lakes within the Big Hole River system (in particular
Miner and Mussigbrod Lakes; see Figure 1 above) may be remnant native
populations derived from fluvial Arctic grayling from the Big Hole
River and isolated by recent habitat fragmentation, but widespread
stocking of these and other locations with hatchery-reared Arctic
grayling during the 1930s-1950s (e.g., Everett 1986, p. 4, 16; Kaya
1990, pp. 31, 75-80) also makes it possible that these fish are
introduced populations or that the existing populations are a mixture
of native and introduced Arctic grayling.
Ecology
Northcote (1995) and Kaya (1990) reviewed the ecology of Arctic
grayling and fluvial Arctic grayling of the upper Missouri River,
respectively. Much of the information on fluvial Arctic grayling in the
upper Missouri River system comes from the Big Hole River, Montana (see
Figure 1 above), which contains a fluvial population. Arctic grayling
exhibit life history and migratory forms present in other species of
inland trout and charr, including fluvial and adfluvial. Fluvial
populations are characterized by a cycle of migratory behavior over
their lifespan between spawning, feeding, and overwintering habitats
within rivers or streams (Northcote 1995, pp. 156-160). Fluvial Arctic
grayling typically migrate upstream to spawn in tributary or mainstem
river locations and downstream to overwintering habitats. Such movement
patterns have been observed in fluvial Arctic grayling in Big Hole
River, Montana (Shepard and Oswald 1989, pp. 18, 27-28). Migrations to
feeding habitats may occur if these locations differ from spawning or
overwintering habitats (Kaya 1990, pp. 9-11). Overall, movements by
fluvial populations within and among tributaries and mainstem rivers
may cover hundreds of kilometers (Armstrong 1986, p. 7). Fluvial Arctic
grayling in the Big Hole River system have been shown to migrate in
excess of 80 km (50 mi) between spawning, feeding and wintering areas
(Shepard and Oswald 1989, pp. 18, 21; Lamothe and Magee 2003, pp. 7,
11, 17). Adfluvial Arctic grayling feed and overwinter in lakes, but
migrate to inlet or outlet streams to spawn (Northcote 1995, p. 148-
149; Northcote 1997, pp. 1030-1034).
Age at maturity and longevity in Arctic grayling varies among
systems and is probably related to growth rate, with populations in
colder, less productive habitats maturing at later ages and having a
greater lifespan (Northcote 1995, pp. 155-157). Fluvial Arctic grayling
in the Big Hole River system typically mature at 2 years of age (males)
or 3 years of age (females), and individuals older than 6 years of age
are rare (Liknes 1981, pp. 16-18; Kaya 1990, pp. 18-20; Magee and
Lamothe 2003, p. 22). Arctic grayling are spring spawners. In Montana,
Arctic grayling typically spawn from late April to mid-May by
depositing adhesive eggs over gravel substrate without excavating a
nest or redd (Shepard and Oswald 1989, pp. 24-25, 29; Kaya 1990, pp.
15-16). In general, the reproductive ecology of Arctic grayling is
somewhat different from other salmonid species (trout and salmon) in
that Arctic grayling eggs tend to be comparatively small (Behnke 2002,
p. 328), and males establish and defend spawning territories rather
than defending access to females (Northcote 1995, p. 150). The time
required for development of eggs from embryo until they emerge from
stream gravel and become swim-up fry varies with water temperature, but
averages about 3 weeks for Arctic grayling in the upper Missouri River
basin (Kaya 1990, pp. 16-17). Small, weakly swimming fry of fluvial
Arctic grayling prefer low velocity stream habitats (Kaya 1990, pp. 23-
24; Northcote 1995, pp. 152-153).
Arctic grayling of all ages feed primarily on aquatic and
terrestrial invertebrates captured on or near the water surface
(Northcote 1995, pp. 153-154; Behnke 2002, p. 328). They also will feed
opportunistically on fish and fish eggs (Northcote 1995, p. 154; Behnke
2002, p. 328). Feeding locations for individual fish are typically
established and maintained through size-mediated dominance hierarchies
(e.g., Hughes 1992, pp. 1994-1995).
Although fluvial Arctic grayling may have specific habitat
requirements depending on their life stage (e.g., fry) and ecological
activity (e.g., spawning), individuals inhabiting streams and rivers
often exhibit a preference for pool habitats (Liknes 1981, pp. 22, 28;
Kaya 1990, pp. 20-21; Lamothe and Magee 2003, pp. 13-14, 17; Lamothe
and Magee 2004, p. 24). Vincent (1962, pp. 39, 42) concluded that
fluvial Arctic grayling in Montana typically reside in
[[Page 20309]]
streams with low-to-moderate gradient (<4 percent) and prefer low-to-
moderate water velocities (<60 centimeters/sec). Observations of
fluvial Arctic grayling habitat use in the Big Hole River by Liknes
(1981, p. 28) and Liknes and Gould (1987, p. 128) are consistent with
these generalizations.
Arctic grayling generally prefer cool or coldwater habitats (Hubert
et al. 1985, pp. 9, 14, 25, 27). Selong et al. (2001, p. 1032) placed
Arctic grayling in a ``coldwater'' group of salmonids, along with
Arctic charr and bull trout, based on critical thermal maximum values.
Genetic Relationships Among Arctic Grayling Populations in the Upper
Missouri River Basin
Discussion of genetic divergence among Arctic grayling populations
is complicated by the extensive hatchery propagation and
transplantation of stocks from location to location (Everett 1986, p.
40). Over 10 million grayling of unknown origin were stocked in the Big
Hole River over a 30-year period from the 1930s to the 1950s (Kaya
1990, pp. 31, 75-80). Everett (1986 pp. 42, 43, 47) concluded that the
effect of grayling introductions on local genetics appears stronger in
lake populations than in the Big Hole River. Nonetheless, the limited
available genetic data suggest the presence of two or more groups--
clusters or sets of populations that are genetically more closely
related to each other than they are to other populations of the same
species--of Arctic grayling within the upper Missouri River that may
not be strictly delineated by geography and life history (Leary 2005,
p. 3; Campton 2006, pp. 6-9, 12).
Inferences about genetic differences among Arctic grayling
populations within the upper Missouri River basin are primarily based
on data collected by Everett (1986) and Leary (1990). These two studies
examined how a particular form (allele) of a protein molecule
(allozyme) varied in frequency across Arctic grayling populations in
Montana. Allozymes are gene products coded by DNA, so allozyme
variation can be used to infer genetic relationships among populations,
subspecies or species. Campton (2006, pp. 6, 12), in his review of
those data, suggested the existence of two possible genetic groups: (a)
A Big Hole-Madison River group that includes the fluvial population in
the Big Hole River, certain populations in adjoining waters of the Big
Hole River system (e.g., Bobcat, Miner, and Mussigbrod Lakes, and Steel
Creek; see Figure 1 above; see Everett 1986, p. 7; Leary 1990, pp. 6-
8), and fish from the Madison River-Ennis Reservoir; and (b) a Red Rock
Lakes group that includes native adfluvial populations from the Red
Rock and Elk Lakes system in the upper Beaverhead River system, and a
number of introduced adfluvial populations (Agnes, Grebe, Rogers,
Odell, and Elizabeth Lakes; see Leary 1990, pp. 7-8) believed to be
derived from human introductions of Red Rock Lakes grayling and/or
associated hatchery stocks. The two groups (Big Hole-Madison and Red
Rock Lakes) are differentiated by divergent allele frequencies for two
allozymes (Campton 2006, p. 6). The relative genetic difference between
these two groups within the upper Missouri River basin is less than the
difference between upper Missouri River Arctic grayling and sample
populations from Alaska and Canada (Everett 1986, p. 80; Leary 1990,
pp. 1, 7-8). The level of genetic divergence observed among populations
within the upper Missouri River is consistent with what would be
expected for populations within a geographic area that share a recent
ancestry but have since diverged, as compared with the greater
divergence observed among populations from different geographic areas
or river systems that have been separated from each other for a much
longer period of time (i.e., upper Missouri River versus Alaskan and
Canadian populations).
Campton (2006, p. 12) also noted that a few adfluvial populations
of Arctic grayling in the Big Hole River drainage, including Miner Lake
(see Figure 1 above), appear to share recent ancestry with the mainstem
Big Hole River fluvial population.
Like Campton, Leary also concluded that Big Hole River and Madison
River grayling samples appear to be quite similar (Leary 2005, p. 3).
Leary's interpretation of the genetic relationships among Miner Lake,
Red Rock Lakes, and Elk Lake populations was different from Campton's.
Leary found Miner Lake to be very divergent from all the others, but
also concluded that there was significant divergence between the Red
Rock Lakes and Elk Lake samples (Leary 2005, p. 3). He interpreted the
allozyme data to mean that the adfluvial samples do not appear to form
a genetically distinct group and consequently concluded that the data
do not support the premise that the fluvial and adfluvial life
histories fall into two distinct genetic lineages (Leary 2005, p. 3).
Rather, he contended the data represent divergence among populations
regardless of life history (Leary 2005, p. 3). In his review, Campton
(2006) concurred that the apparent genetic divergence between the two
groups (Big Hole-Madison River and Red Rock Lakes) was not completely
consistent with life histories because several adfluvial populations
belonged to the Big Hole River-Madison River genetic group.
An Arctic grayling population residing in the Sunnyslope irrigation
canal in Teton County, Montana, is thought to be derived from an
introduction into Pishkin Reservoir (Kaya 1990, p. 41; see Figure 1
above) and is not easily assigned to either of the two genetic groups
suggested by Campton. These fish appear to be genetic outliers relative
to the two other native genetic groups of Arctic grayling (Leary 1990,
p. 8; Campton 2006, p. 7).
Overall, both Campton and Leary observe that: (a) Fluvial Arctic
grayling from the Big Hole River are genetically different from native
adfluvial Arctic grayling in Red Rock Lakes based on observed
differences in allozyme allele frequencies even if the genetic
divergence between these populations appears to be low (average Nei's
genetic distance of the cluster containing these populations equals
0.0132 (Leary 1990, pp. 1,8)); (b) the existing genetic data do not
strongly support the hypothesis that the fluvial form of Arctic
grayling in the upper Missouri River represents a unique genetic
lineage, because it is genetically similar to adfluvial populations in
Miner Lake and in the Madison River (Leary 2005, pp. 3-4; Campton 2006,
p. 12); and (c) the low allozyme variability in upper Missouri River
Arctic grayling samples results in a weak dataset for resolving
ancestries among recently diverged populations (Leary 2005, pp. 3-4;
Campton 2006, p. 10). The Service views Campton's and Leary's
conclusions about the ancestral relationships among Arctic grayling
populations in the upper Missouri River as tentative, given the
inherent limitations of the existing genetic data. However, it is the
best available scientific information at this time. Further
investigations with more variable genetic markers, such as
microsatellite DNA, may clarify genetic relationships (Campton 2006,
pp. 10, 14).
Heritable, Behavioral Differences Between Fluvial and Adfluvial Arctic
Grayling in the Upper Missouri River Basin
Arctic grayling exhibit at least two life histories in the upper
Missouri River system--a river-dwelling fluvial form and a lake-
dwelling adfluvial form. Life history variation in salmonid fishes
(trout and salmon) may or may not be related to genetic differentiation
(e.g., Fausch and Young 1995, p. 365). However, experiments designed to
determine whether behavioral
[[Page 20310]]
differences were due to genetic or environmental influences found that
the behavioral differences between fluvial and adfluvial Arctic
grayling in Montana were heritable. In tests of swimming behavior of
young-of-year Arctic grayling raised in common conditions in captivity,
progeny of fluvial Big Hole River fish behaved significantly
differently, on average, than adfluvial progeny from Red Rock Lakes and
Madison River-Ennis Reservoir populations (Kaya 1989, 1991; Kaya and
Jeanes 1995). The Big Hole River progeny exhibited a greater tendency
to hold position in flowing water (Kaya and Jeanes 1995, pp. 453-456).
Because the test fish from the Big Hole River population were progeny
of parents reared in a non-fluvial environment, retention of this
rheotactic behavior (behavior in response to flowing water) was taken
as evidence that such behavior has a genetic (heritable) basis (Kaya
and Jeanes 1995, p. 456), consistent with conclusions of previous
investigations (Kaya 1989, pp. 474, 478-479; Kaya 1991, pp. 53, 55-58).
Expression of rheotactic characteristics in Arctic grayling also
can be influenced by ontogeny, or the developmental history of an
individual (in this case, time from emergence from gravel as fry until
maturity; Kaya 1991, pp. 53, 55-57), and environmental conditions, such
as time of day (Kaya 1989, p. 56), light intensity (Kaya 1989, p. 478;
Kaya 1991, p. 56), or water temperature (Kaya 1989, p. 478). However,
the collective results are nonetheless consistent with the hypothesis
that heritable, behavioral differences in the test populations exist
between the fluvial and adfluvial populations and those associated with
lakes or reservoirs.
Adfluvial Arctic grayling repeatedly introduced into rivers have
failed to establish viable populations (Kaya 1992b, pp. 12-14).
Adaptive divergence and lack of ecological exchangeability between life
history types are among the factors that may have contributed to these
failures (Campton 2006, p. 13). However, introductions of fluvial
grayling into other rivers within the native range have not been
successful either, so success may be due to other factors (e.g.,
habitat degradation or competition with nonnative fish (Kaya 1992b, pp.
10-12, 60)). In general, life history expression in salmonid species
can be flexible, and Arctic grayling exhibit variation in migratory
behavior across the range of the species (Northcote 1997, p. 1030).
Geography may be a stronger determinant of ancestral relationships than
life history for Arctic grayling. Native Arctic grayling populations
within the upper Missouri River basin may be similar based on genetics,
because they reside in the same river basin and presumably share a
recent evolutionary ancestry (Campton 2006, p. 12), while at the same
time expressing different life histories in response to local habitat
conditions.
Previous Federal Action
The Service initiated a status review for the Montana Arctic
grayling (Thymallus arcticus montanus) through a notice of review
published on December 30, 1982 (47 FR 58454). In that notice, Montana
Arctic grayling was designated a Category 2 species, which included
taxa for which information in possession of the Service at that time
indicated that proposing to list the species as Endangered or
Threatened was possibly appropriate, but for which substantial data
were not currently available to biologically support a proposed rule
(47 FR 58454). We received a petition, dated October 2, 1991, from the
Biodiversity Legal Foundation and George Wuerthner on October 9, 1991.
The petition requested that the ``fluvial Arctic grayling'' be listed
as an endangered species throughout its historic range ``in the
conterminous United States.''
We published a notice of a 90-day finding in the January 19, 1993,
Federal Register (58 FR 4975). In that 90-day finding we found that 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. We also found that because the
Michigan population of Arctic grayling is extinct and, therefore, by
definition cannot be listed, the finding would address only the fluvial
population of the Arctic grayling in the upper Missouri River drainage.
On July 25, 1994, we published a notice of a 12-month petition
finding in the Federal Register concluding that listing the fluvial
Arctic grayling indigenous to the upper Missouri River was warranted
but precluded by other higher priority listing actions (59 FR 37738).
This finding stated that the Service viewed adfluvial Arctic grayling
as not under consideration in the Service's finding as it was believed
to be a distinct population from the fluvial Arctic grayling. This 1994
status review identified the fluvial form of Arctic grayling in the
upper Missouri River drainage as a DPS based on its geographic
isolation and behavioral distinctiveness (59 FR 37738-37741, July 25,
1994). This status review occurred prior to the finalization of the
Service and the National Marine Fisheries Service's joint DPS policy in
1996 (61 FR 4722, February 7, 1996).
Since 1994, and based on the best available information and the
assessment that we conduct during our candidate review process, we have
continued to preliminarily recognize the fluvial Arctic grayling of the
upper Missouri River as a DPS, and has maintained it as a candidate
species through the annual Candidate Notice of Review. In 2004, the
Service elevated the listing priority number of the fluvial Arctic
grayling to 3 (69 FR 24881, May 4, 2004) because the abundance of the
remnant population in the Big Hole River declined substantially and
reestablishment efforts had not yet produced self-sustaining
populations elsewhere in the upper Missouri River.
On May 31, 2003, the Center for Biological Diversity and Western
Watersheds Project (collectively plaintiffs) filed a complaint in
United States District Court in Washington, DC (1:03-cv-01110),
challenging the Service's continuing ``warranted but precluded''
determination for fluvial Arctic grayling contained in the 2002
Candidate Notice of Review (67 FR 40657, June 13, 2002). Plaintiffs
filed an amended complaint on July 22, 2004, challenging the Service's
failure to use its emergency listing authority to protect the fluvial
Arctic grayling under the Act (16 U.S.C. 1531 et seq.). The litigation
with plaintiffs was settled in August 2005. In this settlement
agreement, the Service agreed that on or before April 16, 2007, it
shall submit for publication in the Federal Register a final
determination made pursuant to the Act as to whether or not the
``Montana fluvial Arctic grayling'' is an endangered or threatened
species. During the evaluation of the petition, the Service considered
the term ``Montana fluvial Arctic grayling'' as synonymous with
``fluvial Arctic grayling of the upper Missouri River.'' In this
finding, as in the past, the fluvial form of the indigenous Arctic
grayling from the upper Missouri River drainage in Montana and Wyoming
is referred to as the fluvial Arctic grayling. This revised 12-month
finding is being published as a final listing determination in
accordance with the settlement agreement.
Distinct Vertebrate Population Segment
Pursuant to the Act, we must consider for listing any species,
subspecies, or, for vertebrates, any DPS of these taxa if there is
sufficient information to indicate that such action may be warranted.
The petition we received
[[Page 20311]]
concerns a potential DPS of fluvial Arctic grayling. Under our Policy
Regarding the Recognition of Distinct Vertebrate Population Segments
(61 FR 4722, February 7, 1996) (known as the DPS Policy), three
elements are considered in a decision regarding the status of a
possible DPS as endangered or threatened under the Act. These factors
are applied similarly for additions to the Lists of Endangered and
Threatened Wildlife and Plants (Lists), reclassification, and removal
from the Lists. They are: (1) Discreteness of the population segment 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 (i.e., is the population
segment, when treated as if it were a species, endangered or
threatened?). Discreteness refers to the isolation of a population from
other members of the species, and we evaluate this based on specific
criteria that are also contained in the DPS Policy and are listed
below. If the population segment is determined to be discrete, then we
evaluate significance by using the available scientific information to
determine the population segment's importance to the taxon to which it
belongs. If we determine that a population segment is discrete and
significant, we subsequently evaluate it for endangered or threatened
status based on the Act's standards.
Discreteness
Under our DPS Policy, a population segment of a vertebrate species
may be considered discrete if it satisfies either one of the following
conditions: (1) It is markedly separated from other populations of the
same taxon as a consequence of physical, physiological, ecological, or
behavioral factors. Quantitative measures of genetic or morphological
discontinuity may provide evidence of this separation; or (2) It is
delimited by international governmental boundaries within which
differences in control of exploitation, management of habitat,
conservation status, or regulatory mechanisms exist that are
significant in light of section 4(a)(1)(D) of the Act.
The subject of this DPS evaluation is the fluvial Arctic grayling
of the upper Missouri River. In response to a petition, the fluvial
Arctic grayling was the subject of a status review by the Service in
1994, which identified Arctic grayling indigenous to the Big Hole and
Madison Rivers as elements of a fluvial DPS in the upper Missouri River
(59 FR 37738-37741, July 25, 1994). However, this status review
occurred prior to the finalization of the Service and the National
Marine Fisheries Service's joint DPS policy in 1996 (61 FR 4722,
February 7, 1996). Since 1994, and most recently in 2004 and 2005, the
Service reviewed the available information concerning the taxonomic
status of the species in relation to the DPS policy and again
preliminarily determined that the fluvial Arctic grayling of the upper
Missouri River was a valid DPS (Service 2004, 2005). This DPS
evaluation considers the information used in the previous assessments
as well as a solicited review (Campton 2006) and unsolicited review
(Leary 2005) of the available genetic data for Arctic grayling in
Montana.
(1) Fluvial Arctic Grayling Are Discrete as a Consequence of Physical
Features
Fluvial arctic grayling native to the upper Missouri River are
``markedly separated'' from other grayling, both those in Canada and
Alaska, and from the adfluvial form in the Missouri River drainage
because of physical and reproductive isolation. Fluvial actic grayling
are geographically disjunct and reproductively isolated from
populations inhabiting Arctic Ocean and Hudson Bay drainages in Canada
and Alaska (Scott and Crossman 1973, p. 301). Arctic grayling in the
upper Missouri River are reproductively isolated from their nearest
conspecifics by at least 800 kilometers (km) (500 miles (mi)) (Nelson
and Paetz 1991, p. 255) and have been separated from Arctic Ocean
populations for perhaps 70,000 years as a result of glacial activity
(Lynch and Vyse 1979, p. 263; Redenbach and Taylor 1999, p. 32). This
long period of reproductive isolation coupled with genetic drift and
environmental selection pressures has resulted in genetic differences
between Arctic grayling from the Missouri River and elsewhere based on
analyses of allozymes and mitochondrial DNA (Lynch and Vyse 1979, pp.
263, 268, 275; Everett and Allendorf 1985, pp. 22-23, 26; Everett 1986,
pp. 79-80; Redenbach and Taylor 1999, p. 23; reviewed by Campton 2006,
pp. 5-6; reviewed by Leary 2005, pp. 1-3).
Fluvial and adfluvial Arctic grayling within the upper Missouri
River basin are ``markedly separated'' from each other as a result of
physical features. The fluvial form was once widespread in the upper
Missouri River basin, but the adfluvial form was native only to the Red
Rocks Lakes and possible Elk Lake in the headwaters of the Beaverhead
River (Kaya 1990). Extant populations of native fluvial and adfluvial
Arctic grayling within the upper Missouri River are reproductively
isolated, and the available genetic data are consistent with the
hypothesis of two genetic groups of Arctic grayling (the Big Hole--
Madison River and Red Rock Lakes genetic groups) within the upper
Missouri River (Leary 2005, p. 3; Campton 2006, pp. 6-9, 12)
(2) Fluvial Arctic Grayling Are Not Discrete as a Consequence of
Physiological Features
We do not believe that fluvial Arctic grayling are discrete because
of unique or different physiological characteristics. Lohr et al.
(1996) examined the thermal tolerance of juvenile fluvial Arctic
grayling from the Big Hole River to elevated temperatures in laboratory
tests. However, grayling from the Big Hole River did not appear to be
more tolerant of warm stream temperatures than grayling from Alaska
(Lohr et al. 1996, p. 937).
Arctic grayling from the upper Missouri River tend to grow more
quickly than individuals from northern populations (Northcote 1995, pp.
156-157). However, experimental data are lacking that permit these
differences to be attributed to environmental versus genetic
influences.
(3) Fluvial Arctic Grayling Are Not Discrete as a Consequence of
Ecological Features
The Arctic grayling of the upper Missouri River represent the only
natural example of the taxon inhabiting an Atlantic Ocean drainage (via
the Missouri and Mississippi Rivers and Gulf of Mexico). All other wild
populations of Arctic grayling inhabit drainages of the Arctic Ocean,
Hudson Bay, or north Pacific Ocean (USFWS 2005, p. 10). However,
fluvial Arctic grayling of the upper Missouri River basin are not
discrete from adfluvial Arctic grayling of the upper Missouri River
basin as a consequence of ecological features as they exist within a
common drainage.
(4) Fluvial Arctic Grayling Are Discrete as a Consequence of Behavioral
Features
Under historical conditions within the upper Missouri River basin,
native fluvial and adfluvial populations of Arctic grayling spawned in
different locations (Vincent 1962, pp. 98-121; Kaya 1990, pp. 24-30;
Kaya 1992a, pp. 47-53). Homing behavior to natal (birth) habitats that
is typically expressed by Arctic grayling (e.g., Carl et al. 1992, p.
245) would presumably result in the reproductive isolation of
historical fluvial and adfluvial populations even if occasional
exchange was possible. In
[[Page 20312]]
addition, genetic differences between the extant fluvial population in
the Big Hole River and the native adfluvial population in Red Rock
Lakes (e.g., Everett 1986, pp. 79-30; Leary 1990, pp. 7-8) are
consistent with reproductive isolation between those populations based
on observed differences in allozyme allele frequencies.
Fluvial and adfluvial Arctic grayling do not appear to represent
distinct lineages based strictly on life histories within the upper
Missouri River system (e.g., Leary 2005, p. 3; Campton 2006, p. 12);
there are clearly some heritable differences in juvenile swimming
behavior among fluvial Arctic grayling and the native adfluvial
populations in terms of rheotactic response to flowing water (Kaya
1989, pp. 474, 478-479; Kaya 1991, pp. 53, 55-58; Kaya and Jeanes 1995,
pp. 453-456). These differences in behavior are sufficient to satisfy
the discreteness criterion of the DPS policy.
On the basis of the available information, we conclude that the
fluvial Arctic grayling of the upper Missouri River drainage is
discrete from other populations of the same taxon as a consequence of
physical and behavioral factors. Since a population segment of a
vertebrate species may be considered discrete if the first factor is
met (marked separateness), we need not address the second factor
(delimitation by an international boundary). Therefore, we considered
the potential significance of this discrete population to the remainder
of the taxon.
Significance
If a population segment is determined to be discrete, the Service
considers the available scientific evidence of its significance to the
taxon to which it belongs. Our policy states that this consideration
may include, but is not limited to, the following:
(1) Persistence of the discrete population segment in an ecological
setting unusual or unique for the taxon;
(2) Evidence that loss of the discrete population segment would
result in a significant gap in the range of the 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 historic 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 criteria to
be considered significant. Furthermore, the list of criteria is not
exhaustive; other criteria may be used, as appropriate.
(1) Fluvial Arctic Grayling Do Not Persist in an Ecological Setting
Unusual or Unique for the Taxon
As discussed above, Arctic grayling generally occur throughout
their native range in the holarctic region of Canada and Alaska to
eastern Siberia and northern Eurasia (Scott and Crossman 1973, pp. 301-
302). In our 2005 candidate assessment, we asserted that the fluvial
Arctic grayling of the upper Missouri River persist in an ecological
setting unusual or unique for the taxon as they represent the only
natural example of the taxon inhabiting an Atlantic Ocean drainage via
the Missouri and Mississippi Rivers and Gulf of Mexico. We noted that
all other wild populations of Arctic grayling inhabit drainages of the
Arctic Ocean, Hudson Bay, or north Pacific Ocean (USFWS 2005, p. 10).
However, as established above, we now note that adfluvial Arctic
grayling also persist in the upper Missouri River drainage. Our prior
finding did not take these fish into account in its discussion of
ecological setting. Because both the fluvial and adfluvial forms are
found in the upper Missouri drainage, we cannot find that the
population persists in an ecological setting unique or unusual to the
taxon as a whole.
Further, existence of the species in a different drainage, or
different rivers and lakes, from those grayling found in Canada and
Alaska is not necessarily evidence of a unique ecological setting.
Arctic grayling in the neararctic region are found in the same habitat
type as those in Montana. Grayling inhabit clear water streams, rivers,
and lakes. Riverine populations depend on large streams, deep pools of
small streams, or spring-fed reaches that are not completely frozen in
winter for overwinter survival. Populations not associated with lakes
are found in both Alaska and Montana (Hubert 1985, p. 1). For this
reason also, we find that fluvial Arctic grayling do not persist in an
ecological setting unique or unusual for the taxon.
(2) The Loss of the Fluvial Arctic Grayling Would Not Result in a
Significant Gap in the Range of the Taxon
Loss of the fluvial Arctic grayling in the upper Missouri River,
when considered in relation to grayling throughout the remainder of the
nearctic region, would mean the loss of a small percentage of the
entire range of the taxon. Due to the broad geographic range of Arctic
grayling, the gap in the range of Arctic grayling resulting from the
loss of fluvial Arctic grayling in the upper Missouri River basin would
not result in a significant gap in the range of the taxon as a whole.
In our 2005 candidate assessment, we asserted that the loss of the
fluvial Arctic grayling of the upper Missouri River would result in a
significant gap in the range of the taxon as these fish are the only
extant fluvial grayling population in the contiguous United States and
represent the southernmost extent of the species (USFWS 2005, p. 10).
However, the Ninth Circuit Court has rejected this argument as a
misconstruction of this criterion in the case of National Association
of Home Builders v. Norton, 340 F. 3d 835, 852 (9th Cir. 2003)
concerning the cactus ferruginous pygmy-owl (Glaucidium brasilianum
cactorum) (70 FR 44551, August 3, 2005). The Court found that in
designating a DPS under the DPS policy, we must find that a discrete
population is significant to the taxon as a whole, not to the United
States. Therefore, we have determined, based on the information
available to the Service, the loss of the fluvial Arctic grayling in
the upper Missouri River would not result in a significant gap in the
range of the species on the basis of the significance of the Montana
population to the species as a whole.
(3) Fluvial Arctic Grayling Do Not Represent the Only Surviving Natural
Occurrence of the Taxon
This criterion from the DPS policy does not apply to the fluvial
Arctic grayling in the upper Missouri River because it is clearly not a
population segment representing the only surviving natural occurrence
of a taxon that may be more abundant elsewhere as an introduced
population outside its historic range. Consequently, this population of
grayling is not significant according to this standard.
(4) Fluvial Arctic Grayling in the Missouri River Drainage Do Not
Differ Markedly in Genetic Characteristics From Adfluvial Populations
in the Missouri River Drainage
As noted above, analyses of allozymes and mitochondrial DNA show
genetic divergence between Arctic grayling in the upper Missouri River
and Arctic grayling in Canada and Alaska (Lynch and Vyse 1979, pp. 263,
268, 275; Everett and Allendorf 1985, pp. 22-23, 26; Everett 1986, pp.
79-80; Redenbach and Taylor 1999, p. 23; reviewed by Campton 2006, pp.
5-6; reviewed by Leary 2005, pp. 1-3) and appear to be most closely
related evolutionarily to populations in northeastern
[[Page 20313]]
Saskatchewan, Canada (Stamford and Taylor 2004, p. 1538).
In addition, fluvial Arctic grayling from the Big Hole River are
genetically different from native adfluvial Arctic grayling in Red Rock
Lakes based on observed differences in allozyme allele frequencies
(Campton 2006, p. 6). However, the relative genetic difference between
these two groups within the upper Missouri River basin is less than
that between upper Missouri River Arctic grayling and sample
populations from Alaska and Canada (Leary 1990, pp. 1, 7-8).
Resolving ancestries among recently diverged upper Missouri River
Arctic grayling populations is difficult due to the low allozyme
variability among samples (Leary 2005, pp. 3-4; Campton 2006, p. 10).
In this case, although allozyme data from 39 loci are available from
these populations, only 2 of the loci analyzed were generally variable
among them (Everett 1986; Leary 1990; Leary 2005, p. 3). Information
from only two loci may cause chance similarities or differences and
require cautious interpretation (Leary 2005, p. 3).
Likewise, the paucity of genetic variation detected by Redenbach
and Taylor (1999, p. 27) in their restriction enzyme analysis of mtDNA
of upper Missouri River basin Arctic grayling precludes making any
inferences about genetic similarities or differences among the upper
Missouri River populations sampled except that they all appear to share
a common maternal lineage (Leary 2005, p. 4). The level of genetic
divergence observed among populations within the upper Missouri River
is consistent with what would be expected for populations within a
geographic area that share a recent ancestry (Campton 2006, p. 12).
Discerning genetic divergence among Arctic grayling populations is
further complicated by the extensive hatchery propagation and
transplantation of stocks, as discussed above (Everett 1986, p. 40).
The Service does not regard the introduced, lake-dwelling grayling to
be part of the indigenous upper Missouri River fluvial Arctic grayling
population (59 FR 37739, July 25, 1994). However, widespread stocking
of hatchery-reared Arctic grayling in the Big Hole River system and
other locations (e.g., Everett 1986, pp. 4, 16; Kaya 1990, pp. 31, 75-
80) makes it possible that some fish are introduced populations or that
the existing populations are a mixture of native and introduced Arctic
grayling.
We find that, based on the genetic information currently available,
the fluvial Arctic grayling of the upper Missouri River drainage do not
differ markedly from adfluvial populations of the species in their
genetic characteristics such that they should be considered
biologically or ecologically significant based simply on genetic
characteristics. Biological and ecological significance under the DPS
policy is always considered in light of Congressional guidance (see
Senate Report 151, 96th Congress, 1st Session) that the authority to
list DPSs be used ``sparingly'' while encouraging the conservation of
genetic diversity.
Conclusion on DPS
Under section 3 of the Act and our implementing regulations at 50
CFR 424.02, a ``species'' is defined to include any species or
subspecies of fish, wildlife, or plant, and any distinct population
segment of any vertebrate species which interbreeds when mature. Our
implementing regulations provide further guidance on determining
whether a particular taxon or population is a species or subspecies for
the purposes of the Act: ``The Secretary shall rely on standard
taxonomic distinctions and the biological expertise of the Department
and the scientific community concerning the relevant taxonomic group''
(50 CFR 424.11). As noted above, Arctic grayling in the upper Missouri
River basin have been classified into separate species and subspecies,
but these designations are not widely accepted. Therefore, we do not
consider the subject of this petition to constitute a distinct species
or subspecies.
The 1994 status review identified the fluvial form of Arctic
grayling in the upper Missouri River drainage as a DPS based on its
geographic isolation and behavioral distinctiveness (59 FR 37738, July
25, 1994). On the basis of the best available information, we continue
to conclude that the fluvial Arctic grayling of the upper Missouri
River drainage is ``markedly separated'' from all other populations of
the same taxon as a consequence of physical and behavioral factors.
Consequently, the Service concludes that the petitioned entity is
discrete according to the 1996 DPS policy. However, on the basis of the
four significance criteria in the 1996 DPS Policy, the Service is
unable to conclude at this time that the petitioned entity is
significant. Therefore, we find that the fluvial Arctic grayling of the
upper Missouri River does not qualify as a distinct population segment
under the Act.
Significant Portion of the Range
Pursuant to the Act and our implementing regulations, a species may
warrant listing if it is threatened or endangered in a significant
portion of its range. However, the petition did not request that we
determine whether the grayling was threatened or endangered in a
significant portion of its range. Rather, it asked that we list the
fluvial Arctic grayling in the U.S. as an endangered species.
Consistent with the petition, our previous petition findings have
uniformly addressed possible listing in the context of whether the
fluvial Arctic grayling in Montana constitutes a DPS, and therefore a
``species'' under the Act. As discussed above, we have now determined
that the fluvial Arctic grayling is not a DPS. Thus, we have disposed
of the question raised by the petition: we have no obligation under the
Act to address the separate question of whether the fluvial Arctic
grayling in Montana constitutes a significant portion of the range of
some of the entire grayling species, or some valid but currently
undefined DPS. If the Service determines in the future that the
grayling is threatened or endangered in a significant portion of its
range, we will add the species to the candidate list and propose its
listing. However, that would be a future action. Because the petition
and our prior finding were with respect to a DPS, and we have found
that there is not a valid DPS, we do not need to address significant
portion of the range at this time.
Finding
On the basis of the discussion presented in this document, we find
that the fluvial Arctic grayling of the upper Missouri River does not
qualify as a distinct population segment. As a result, we find that the
petition to list the fluvial Arctic grayling of the upper Missouri
River is not warranted. Based on this determination, we withdraw the
fluvial Arctic grayling of the upper Missouri River from the candidate
list. Although no further action will result from this finding, we
request that you submit new information concerning the taxonomy,
biology, ecology, and status of the Arctic grayling of the upper
Missouri River system to the Montana Field Office (see ADDRESSES below)
whenever it becomes available. We will accept additional information
and comments from all concerned governmental agencies, the scientific
community, industry, or any other interested party concerning this
finding; and will reconsider this determination in the event of new
information as appropriate. The Service continues to strongly encourage
cooperative conservation and restoration of fluvial Arctic grayling in
the upper Missouri River.
[[Page 20314]]
References
A complete list of all references cited herein is available upon
request from the Montana Field Office, U.S. Fish and Wildlife Service
(see ADDRESSES).
Author
The authors of this finding are biologists in Region 6 of the U.S.
Fish and Wildlife Service.
Authority
The authority for this action is the Endangered Species Act of
1973, as amended (16 U.S.C. 1531 et seq.).
Dated: April 13, 2007.
H. Dale Hall,
Director, Fish and Wildlife Service.
[FR Doc. E7-7484 Filed 4-23-07; 8:45 am]
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