Endangered and Threatened Wildlife and Plants; 12-Month Finding on a Petition To List the Coaster Brook Trout as Endangered, 23376-23388 [E9-11527]
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Federal Register / Vol. 74, No. 95 / Tuesday, May 19, 2009 / Proposed Rules
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Background
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[FWS–R3–ES–2008–0030; 92210–1111–
0000–FY09–B3]
Endangered and Threatened Wildlife
and Plants; 12-Month Finding on a
Petition To List the Coaster Brook
Trout as Endangered
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AGENCY: Fish and Wildlife Service,
Interior.
ACTION: Notice of 12-month petition
finding.
SUMMARY: We, the U.S. Fish and
Wildlife Service (Service), announce a
12-month finding on a petition to list
the coaster brook trout (Salvelinus
fontinalis) as endangered under the
Endangered Species Act of 1973, as
amended (Act). The petition also asked
that critical habitat be designated for the
species. After review of all available
scientific and commercial information,
we find that the coaster brook trout is
not a listable entity under the Act, and
therefore, listing is not warranted. We
ask the public to continue to submit to
us any new information that becomes
available concerning the taxonomy,
biology, ecology, and status of coaster
brook trout and to support cooperative
conservation of coaster brook trout
within its historical range in the Great
Lakes.
DATES: The finding announced in this
document was made on May 19, 2009.
ADDRESSES: This finding is available on
the Internet at https://
www.regulations.gov at Docket Number
[FWS–R3–ES–2008–0030]. Supporting
documentation for this finding is
available for inspection, by
appointment, during normal business
hours at the U.S. Fish and Wildlife
Service, Region 3 Fish and Wildlife
Service Regional Office, 1 Federal Drive,
Bishop Henry Whipple Federal
Building, Fort Snelling, MN 55111.
Please submit any new information,
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materials, comments, or questions
concerning this finding to the above
address, Attention: Coaster brook trout.
FOR FURTHER INFORMATION CONTACT:
Jessica Hogrefe, Region 3 Fish and
Wildlife Service Regional Office (see
ADDRESSES) (telephone 612–713–5346;
facsimile 612–713–5292). Persons who
use a telecommunications device for the
deaf (TDD) may call the Federal
Information Relay Service (FIRS) at
800–877–8339.
SUPPLEMENTARY INFORMATION:
Section 4(b)(3)(B) of the Act (16
U.S.C. 1531 et seq.) requires that, for
any petition to revise the Lists of
Endangered and Threatened Wildlife
and Plants that contains substantial
scientific and commercial information
that listing may be warranted, we make
a finding within 12 months of the date
of our receipt of the petition on whether
the petitioned action is: (a) Not
warranted, (b) warranted, or (c)
warranted, but the immediate proposal
of a regulation implementing the
petitioned action is precluded by other
pending proposals to determine whether
species are threatened or endangered,
and expeditious progress is being made
to add or remove qualified species from
the List of Endangered and Threatened
Species. Section 4(b)(3)(C) of the Act
requires that we treat a petition for
which the requested action is found to
be warranted but precluded as though
resubmitted on the date of such finding,
that is, requiring that we make a
subsequent finding within 12 months.
Such 12-month findings must be
published in the Federal Register. This
notice constitutes our 12-month finding
for the petition to list the U.S.
population of coaster brook trout.
Previous Federal Action
The Sierra Club Mackinac Chapter,
Huron Mountain Club, and Marvin J.
Roberson filed a petition, dated
February 22, 2006, with the Secretary of
the Interior to list as endangered the
‘‘naturally spawning anadromous (lakerun) coaster brook trout throughout its
known historic range in the
conterminous United States’’ and to
designate critical habitat under the Act.
The petition clearly identified itself as
such and included the requisite
identification information for the
petitioners, as required in 50 CFR
424.14(a). On behalf of the petitioners,
Peter Kryn Dykema, Secretary of the
Huron Mountain Club, submitted
supplemental information, dated May
23, 2006, in support of the original
petition. This supplemental information
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provided further information on the
species’ status and biology, particularly
for brook trout in the Salmon Trout
River.
On September 13, 2007, we received
a 60-day notice of intent to sue over the
Service’s failure to determine, within 1
year of receiving the petition, whether
the coaster brook trout warrants listing.
Under section 4 of the Act, the Service
is to make a finding, to the maximum
extent practicable within 90 days of
receiving a petition, that it does or does
not present substantial scientific or
commercial information indicating that
the petitioned action may be warranted.
Further, the Act requires that, within 12
months of receiving a petition found to
present substantial information, the
Service must determine whether the
petitioned action is warranted. A
complaint was filed in U.S. District
Court in the District of Columbia on
December 17, 2007, for failure to make
a timely finding (Sierra Club, et al. v.
Kempthorne, No. 1:07–cv–02261 (D.D.C.
December 17, 2007)). The Service
reached a negotiated settlement with the
plaintiffs to submit the 90-day finding to
the Federal Register by March 15, 2008.
We published a ‘‘substantial’’ 90-day
finding March 20, 2008. The negotiated
settlement further required the Service
to publish the 12-month finding in the
Federal Register by December 15, 2008.
The deadline for the 12-month finding
was extended to April 15, 2009, by
mutual consent. On April 15, 2009, we
filed an unopposed motion to extend
the deadline for the coaster brook trout
12-month finding to May 12, 2009.
Species Information
Species Description
Brook trout (Salvelinus fontinalis),
also called brook char or speckled trout,
is one of three species in the genus
Salvelinus (chars) native to north and
eastern North America; the others being
lake trout (S. namaycush) and Arctic
char (S. alpinus). The chars are a subgroup of fishes in the salmon and trout
subfamily (Salmoninae) that is distinct
from the ‘‘true’’ trout and salmon subgroups.
The brook trout throughout its range
in eastern North America exhibits
considerable variation in growth rate,
color, and other features, but generally
can be distinguished from other char
and trout species by its olive-green to
dark brown back with a light yellowbrown vermiculate pattern, sides with
large yellow-brown spots and blue halos
surrounding small, sporadic red and
orange spots. Pectoral, pelvic, anal, and
lower caudal fin have leading edges of
white bordered by black with the
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remainder predominantly reddish to
orange. Sea-run brook trout become
silver with purple iridescence and show
red spots on the sides (Scott and
Crossman 1973, p. 208).
Distribution
The historical range of native brook
trout extends along Hudson Bay in
Canada across the Provinces of
Manitoba, Ontario and Quebec, to
Newfoundland and Labrador and south
to Nova Scotia and New Brunswick in
Canada; and from eastern Iowa through
northern Illinois, northern Ohio, and the
Great Lakes drainage (Minnesota,
Michigan, Wisconsin), through the New
England States (New York, New
Hampshire, Vermont, Maine,
Massachusetts, Pennsylvania, New
Jersey), large New England rivers (such
as the Hudson River and Connecticut
River), and through the Appalachian
Mountains in Maryland, Virginia, West
Virginia, North Carolina, South
Carolina, Tennessee, south to Georgia
(MacCrimmon and Campbell 1969, pp.
1700–1702; MacCrimmon et al. 1971, p.
452; Scott and Crossman 1973, pp. 209–
210; Power 1980, p. 142). Naturalized
populations of brook trout were
established as early as the late 1800s
beyond the historical native range by
introductions to waters in western
North America, South America, Eurasia,
Africa, and New Zealand (MacCrimmon
and Campbell 1969, p. 1699, pp. 1703–
1717). The current range of native brook
trout still extends through Canada and
down to Georgia in the U.S., but in
many locations, populations have been
completely extirpated or have
contracted within this range towards
upper stream reaches, higher altitudes,
or headwaters (EBJV 2006, p. 2).
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Distribution of Brook Trout in the Great
Lakes
According to Bailey and Smith (1981,
p. 1549) and MacCrimmon and
Campbell (1969, p. 1701), brook trout
are native to the lakes and tributaries of
Lakes Superior, Huron, Michigan, and
the tributaries of Lakes Erie and Ontario.
Brook trout are not believed to have
been present in Minnesota streams
above barrier falls to Lake Superior
(Smith and Moyle 1944, p. 119) or
throughout most of the lower peninsula
of Michigan (MIDNR 2008a, pp. 1–2;
MacCrimmon and Campbell 1969, p.
1704).
Habitat Requirements
Brook trout require clear, cold, welloxygenated water to thrive. They are
generally found in water ranging
between 41–68° Fahrenheit (5–20°
Celsius), with their likely preferred
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temperature falling near the middle of
this range (Power 1980, p. 172). Thermal
requirements within this range vary by
life cycle phase and season (Scott and
Crossman 1973, p. 211; Blanchfield and
Ridgway 1997, p. 750; Baril and Magnan
2002, pp. 177–178).
The brook trout spawns in late
summer or autumn, the date varying
with latitude and temperature.
Spawning takes place most often over
gravel beds but may be successfully
accomplished over a variety of
substrates if there is spring upwelling or
a moderate current (Scott and Crossman
1973, p. 210). Power (1980, p. 151)
describes rangewide brook trout
spawning, which occurs in the fall,
when day length and temperature are
decreasing. In northerly regions and at
high elevations, brook trout may spawn
as early as late August and spawning
may be delayed until December in
southern areas. As is typical for
salmonids, females prepare redds
(hollows scooped out for spawning) in
suitable gravel substrate. The female
then deposits her eggs in the redd where
they are fertilized by a male. After
spawning there is no further parental
involvement with the young. The redd
protects the eggs and allows an adequate
exchange of dissolved gases and other
materials during development.
Brook trout are carnivorous, feeding
opportunistically upon a variety of prey,
such as worms, leeches, crustaceans,
aquatic insects, terrestrial insects,
spiders, mollusks, and fish (Scott and
Crossman 1973, p. 212). Anadromous
(migrating from salt water to spawn in
fresh water) forms vary their feeding
behavior and prey items based on their
age and the environment, marine or
riverine, they are occupying (Newman
and Dubois 1997, p. 9). Brook trout also
show diverse foraging behaviors; some
individuals may be sedentary, eating
crustaceans from the lower portion of
the water column, whereas others in the
same system may be more active and eat
insects from the upper portion of the
water column (McLaughlin et al. 1999,
p. 386). This resource polymorphism
may play a supplementary role in the
extensive adaptive radiation (evolution
of ecological variability within a rapidly
multiplying lineage; Smith and
´
Skulason 1996) observed in this species.
Genetics of Brook Trout
A large amount of genetic variation for
brook trout is distributed among populations
(large Fst values). This pattern is heavily
influenced by the diverse ecological and lifehistory characteristics of brook trout
populations (population connectivity or
isolation, philopatric tendency). This pattern
of highly differentiated populations of brook
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trout is found at small and large geographic
scales. Population genetic structuring is
common in brook trout throughout its range
(Angers et al. 1999, pp. 1049–1050). Like
many salmonids, brook trout tend to have a
hierarchical population structure resulting
from the hierarchical design of the networks
of streams and lake or coastal areas in which
they live, and a complicated life cycle that
leads to strong local adaptations. Taxonomic
resolution can be even more complicated at
the lake level when lakes include sympatric
(occupying the same or overlapping
geographic area without interbreeding) but
genetically divergent brook trout populations
such as in Lake Mistassini in Canada (Fraser
and Bernatchez 2008, p. 1197). This degree
of genetic divergence that forms among
populations is reflective of the reproductive
connections (isolation) among the
populations across the range of the taxon.
Six distinct genetic mitochondrial
(mtDNA) clades have been identified
throughout the range of brook trout in
eastern North America (Danzmann et al.
1998, p. 1307). These mtDNA clades
reflect historical isolation in glacial
refugia or long periods of isolation in
nonglacial areas in the southern part of
the species’ range. The Wisconsin
glacial advance which covered portions
of Canada covered all five Great Lakes
15,000 years ago (Bailey and Smith
1981, p. 1543). As these glaciers
receded, brook trout recolonized the
lakes from the Mississippi and Atlantic
refugia (Danzmann et al. 1998, pp. 1308,
1312). Given this pattern of glaciation,
genetic diversity is greatest at the
southern portion of the species’ range
and gradually decreases northward
(Danzmann et al. 1998, pp. 1310–1311).
As the most geographically isolated (for
tens of thousands of years), brook trout
in the southern part of the species’ range
(along the Appalachian Mountains
south to Georgia) are the most diverse,
containing all six mtDNA clades. The
Great Lakes contains three of the six
mtDNA clades. Throughout the northern
portion of their range in Canada, brook
trout are the least genetically diverse,
with only a single mtDNA clade present.
Within each of these lineages, there is
evidence to suggest that selection is
driving rapid phenotypic divergence in
some populations.
Results based on microsatellite DNA
variation identified nine distinct genetic
assemblages of brook trout in the U.S.
(King 2009, unpub. data). Assemblages
from the nonglacial southern part of the
species’ range (along the Appalachian
Mountains from Pennsylvania to
Georgia) in the U.S. are the most
genetically divergent, and this
divergence among the assemblages
generally decreases as the range
progresses northward.
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Genetics of Brook Trout in the Great
Lakes
Populations from Lake Superior and
tributaries to Lake Erie form two of the
nine genetic assemblages of brook trout
in the U.S. The Lake Erie populations
are the most divergent assemblage from
the northern part of the species’ range.
Lake Superior populations are similar in
the degree of genetic divergence to the
remaining northern assemblages
grouping with the average genetic
distance between brook trout
populations in the U.S. Samples from
the rest of the Great Lakes were not
available for analysis. Although brook
trout in the Great Lakes do not contain
any wholly unique mtDNA clades, they
do contain a large amount of the genetic
variation in a confined portion of the
range (Danzmann et al. 1998, pp. 1310–
1311).
Native populations of brook trout in
Lake Superior in most cases have
retained their native genetic
characteristics despite the stocking of
hatchery fish from sources outside and
within the Lake Superior basin. In Lake
Superior, the intensity and purpose of
stocking has varied over time and space.
For example, Minnesota tributaries to
Lake Superior have been stocked with
hatchery strains that originated from
outside of the Great Lakes Basin to
provide fishing opportunities above fish
passage barriers (Wilson et al. 2008, p.
1312). Until the early 1990s, most of the
stocked fish in Lake Superior were
domesticated strains from outside the
Great Lakes basin (Schreiner et al. 2008,
p. 1357), although many stocking events
were undocumented and records of
early stocking events are incomplete
(Wilson et al. 2008, p. 1312). These
stocking efforts were not targeted at
rehabilitation and from that perspective,
results were poor. The stocked fish were
not behaviorally or evolutionarily
adapted to the environment in which
they were planted, criteria known to
limit survival and reproductive success
(Schreiner et al. 2008, p. 1357).
Burnham-Curtis (2001, p. 2) concluded
that hatchery fish have had little
reproductive success in Lake Superior
streams based on her examination of 36
tributaries to Lake Superior and 9
hatchery stocks outplanted into the lake.
However, the genetic methods used by
Burnham-Curtis provided low power to
detect genetic introgression of hatchery
fish into native populations (Wilson et
al. 2008, p. 1312). A recent study by
D’Amelio and Wilson (2008, p. 1215)
used genetic methods with high power
to detect genetic introgression of
hatchery fish into natural populations.
This study documented only low levels
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of genetic introgression of Lake Nipigon
hatchery fish into native populations of
brook trout from six tributaries to Lake
Superior’s Nipigon Bay (D’Amelio and
Wilson 2008, p. 1222), despite decades
of stocking. A study by Scribner et al.
(2006, pp. 3–4) examined nine brook
trout populations from Lake Superior
tributaries on the south shore of
Michigan and four hatchery strains
outplanted into those tributaries. This
study used similar methods to D’Amelio
and Wilson (2008). Scribner et al. (2006,
p. 8) concluded that hatchery stocking
appears to have minimal if any impact
of on brook trout.
Brook Trout Life-History Diversity
An individual’s ability to produce
multiple phenotypes (visible or
observable characteristics) in response
to its environment is termed phenotypic
plasticity (Scheiner 1993, p. 36). Recent
studies have recognized the role of
phenotypic plasticity as a major source
of phenotypic variation in natural
populations (Price et al. 2003, p. 1438).
The brook trout exhibits remarkable
phenotypic plasticity across its natural
range. This plasticity allows it to thrive
in a variety of environments, from cold
subarctic regions, through temperate
zones and in southern refugia in eastern
North America, and in a range of places
where it has been introduced (Power
1980, p. 142). Although primarily a
stream-dwelling species, brook trout
also occupy inland lakes and coastal
waters. Because of the variety of the
freshwater, estuary, and ocean
environments, migratory plasticity is
also favored. The brook trout’s dispersal
subsequent to receding glaciation, and
separation into isolated breeding stocks
in diverse habitats subject to an array of
natural and man-made influences have
all contributed to this variability (Power
1980, p. 142).
Brook trout display considerable lifehistory variation throughout their native
range (Huckins and Baker 2008, p.
1229). Brook trout across its range
exhibit a variety of life-history types
(polymorphisms or ecotypes), including
fluvial (stream-dwelling), adfluvial
(migrating between lakes and streams),
lacustrine (lake-dwelling), and
anadromous (migrating from salt water
to spawn in fresh water) forms.
Understanding life-history diversity in a
species requires knowledge of the
evolutionary history, ecological setting,
and reproductive relationships among
ecotypes. Reproductive interactions
between ecotypes are reflected by the
magnitude and pattern of genetic
differentiation observed between lifehistory phenotypes at neutral genetic
markers. The expression of migratory
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behavior (expressed as the adfluvial and
anadromous ecotypes) by any
individual fish will be partially in direct
response to its environment. Phenotypic
expression of more than one form may
be expected in a population located in
a variable environment containing
habitats for several ecotypes. The
amount of phenotypic plasticity a
population will exhibit for the migratory
trait also has a heritable genetic basis
and will be determined by the intensity
and type of selective pressures that
population experiences (Via and Lande
1985, pp. 517–519; Theriault et al. 2008,
pp. 418–419).
Adoption of migratory adfluvial form
or stream-resident life-history form in
brook trout has been modeled under a
conditional strategy framework where
environmentally influenced threshold
traits determine which ecotype a fish
will adopt (Hendry et al. 2004, pp. 124–
125). Growth rate efficiencies, body size,
and concentration of juvenile hormone
have all been identified as potential
threshold traits (Theriault and Dodson
2003, pp. 1155–1157). Theoretical work
by Ridgway (2008, p. 1185) and Uller
(2008, pp. 436–437) also provide
information to suggest parental effects
are important to the expression of
alternate ecotypes of brook trout. These
parental effects describe an affect of the
parental phenotype on the offspring’s
phenotype such as coaster females
producing larger eggs and spawning in
different locations from stream-resident
ecotypes, influencing the habitat use
(Morinville and Rasmussen 2006, pp.
701–702) and growth rate at the juvenile
stage (Perry et al. 2005, p. 1358). These
differences in growth rate and habitat
use impact potential threshold traits.
Work on sympatric brook trout life
forms at young ages largely comes from
a few studies on anadromous
populations. Morinville and Rasmussen
(2003) studied the bioenergetics of
young brook trout exhibiting
anadromous migratory and streamresident life tactics. They found that the
anadromous migrants have higher
metabolic costs and had consumption
rates 1.4 times that of stream residents
but growth efficiencies of the
anadromous form were lower than that
of residents. Spatial utilization of
habitat differed among the life tactics as
well, with migratory individuals
occupying faster-flowing waters
compared to the resident fish which
used pool areas (p. 408). They
concluded that migrant brook trout have
noticeably different energy budgets than
resident brook trout from the same
system (p. 406). Morinville and
Rasmussen (2008) also investigated
morphological differences between life
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tactics. The authors concluded that
migrant brook trout were found to be
more streamlined (narrower and
shallower bodies) than resident brook
trout, and these differences persisted
into the marine life of the migrant fish
(pp. 175, 183). The differences were
powerful enough to derive discriminant
functions using five of the measured
traits allowing for accurate classification
of juvenile brook trout as either migrant
or resident with an overall correct
classification rate of 87 percent.
A study by Theriault et al. (2007b, p.
61) found that sympatric anadromous
and fluvial brook trout in the SainteMarguerite River in Quebec belonged to
a single gene pool. Phenotypic plasticity
is, therefore, a major force driving the
expression of these two life histories
from this population. Evolution of
phenotypic plasticity in this population
was influenced by mating systems with
most of the mating between different
morphotypes occurring between fluvial
males and anadromous females.
Additional work in this system
demonstrated significant heritability for
life-history tactic and for body size
(Theriault et al. 2007a, pp. 7–8)
indicating expression of life-history
tactic in this population can be effected
by natural or artificial selection.
Life-History Diversity in Great Lakes
Brook Trout
Fish that complete their life cycle
exclusively in tributaries to the Great
Lakes exhibit the fluvial life history and
are defined as stream residents.
‘‘Coaster’’ (the subject of the petition) is
a regional term for a life-history variant
of brook trout in the Great Lakes
(Burnham-Curtis 2001, p. 2; Wilson et
al. 2008, p. 1) which use lake waters of
the Great Lakes for all or a portion of its
life cycle (Becker 1983, p. 320). The
coaster form can be further divided into
an adfluvial ecotype that migrates from
the stream to the lake and back into
tributaries to spawn and a lacustrine
ecotype that completes its life cycle
entirely within the lake (Huckins et al.
2008, p. 1323). In the Great Lakes
region, spawning usually occurs from
mid-September through mid-November.
Distinct life histories associated with
the coaster and stream-resident types
result in different physical,
demographic, and ecological
characteristics for the forms (Huckins et
al. 2008, p. 1337; Huckins and Baker
2008, p. 1241; Ridgway 2008, p. 1185).
Specifically, coasters tend to live longer
than stream residents (5–8 years versus
less than 5 years), reach maturation later
(females at 2–4 years versus 1–2 years),
attain larger length and weight as adults
(12–25 inches and 0.75–8 pounds (30–
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64 centimeters (cm) and 341–3632
grams (g)) versus (5–15 inches (13–38
cm) and (less than 1 pound (<454 g), be
more fecund (1500–3000 eggs per
female versus 100–1500 eggs per
female), and move greater distances (up
to 19–217 miles (30–350 kilometers
(km)) versus less than 19 miles (30 km))
(Scott and Crossman 1973, pp. 208, 210,
211; Power 1980, p. 157; Becker 1983,
pp. 318, 320; Ritchie and Black 1988,
pp. 19, 50, 51; Quinlan 1999, pp. 11, 12,
14, 16, 17, 20; Swainson 2001, pp. 40,
41, 60, 64; WIDNR and USFWS 2005, p.
16; Huckins and Baker 2008, pp. 1239,
1241; Huckins et al. 2008, pp. 1328,
1329, 1337; Mucha and Mackereth 2008,
p. 1210; Schram 2008a, pers. comm.;
Chase 2008, pers. comm.).
Coasters have been historically
documented in Lakes Superior, Huron,
and Michigan brook trout populations
(Bailey and Smith 1981, p. 1549;
Dehring and Krueger 1985, p. 1;
Enterline 2000, p. 1; MIDNR 2008a, pp.
1–2). However, Lake Superior is the
only Great Lake with extant coaster
forms of brook trout, and all available
literature is from this area. Coasters in
the Great Lakes are found in Canada and
the U.S. in substantially fewer locations
than they were historically (Newman et
al. 2003, p. 39). Populations in the Great
Lakes basin with these life-history forms
are documented within Canada in
tributaries to Nipigon and Black Bays,
the Nipigon River, Lake Nipigon and the
Pancake River in the eastern part of
Lake Superior (Newman et al. 2003, p.
39; Chase and Swainson 2009, pers.
comm.). Within the U.S. portion of the
Great Lakes basin, populations that
express the coaster form occur in Isle
Royale National Park in Tobin Harbor,
Big and Little Siskiwit Rivers, and
Washington Creek as well as on the
south shore of Lake Superior in the
Salmon Trout River (Newman et al.
2003, p. 39).
As previously stated, brook trout
populations within the upper Great
Lakes exhibit fluvial, adfluvial, and
lacustrine life-history forms, coasters
comprising the latter two forms.
Populations of brook trout in Lake
Superior likely function as types of
metapopulations, with the coaster life
forms serving as dispersers (D’Amelio
and Wilson 2008, p. 1222; Sloss et al.
2008, p. 1249). The viability of a
metapopulation is strongly contingent
upon maintaining dispersal among
populations. Although brook trout
exhibit spawning site fidelity,
individuals exhibiting the adfluvial life
forms in Lake Superior have also been
shown to stray or disperse among
streams (D’Amelio and Wilson 2008, p.
1222; Mucha and Mackereth, p. 1211).
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The long-term persistence of a
metapopulation requires a balance
between local extinction and
recolonization of constituent
populations (see Hanski 1998 for a
review of metapopulations). Dispersing
individuals offset local population
extinction by providing a means for
recolonization (Brown and KodricBrown 1977, p. 448; Reeves et al. 1995,
p. 340). Dispersing individuals also
provide for gene flow among discrete
populations, countering losses of
genetic fitness while still allowing the
development and distribution of unique
adaptive traits (Ingvarsson 2001, p. 63;
Tallmon et al. 2004, p. 494). Thus, the
coaster life-history forms are important
to the long-term viability of brook trout
populations throughout Lake Superior.
Genetic studies of stream-resident
(fluvial life form) brook trout show
substantial genetic structuring among
populations in Michigan, Wisconsin,
Minnesota, and Canada characterized by
distinct regional groupings or
metapopulations (Burnham-Curtis 1996,
pp. 10–11; Burnham-Curtis 2001, p. 10;
Sloss et al. 2008, p. 1249; Wilson et al.
2008, p. 1312; Scribner et al. 2008, p. 9).
In studies aimed at determining genetic
differences between the coaster
polymorphism and stream-resident fish
occupying tributaries connected to the
lake, molecular genetic work in Lake
Superior indicates that coasters and
stream-resident brook trout occupying
tributaries to the first barrier are parts of
the same population (D’Amelio and
Wilson. 2008, p. 1221; Scribner et al.
2008, p. 9; Stott 2008, p. 5). Work
investigating the genetic differences of
various tributaries to the lake found
distinct differences among populations
of brook trout in each tributary to Lake
Superior (Burnham-Curtis 1996, p. 10;
Burnham-Curtis 2000, p. 7; BurnhamCurtis 2001, p. 10; D’Amelio and Wilson
2008, p. 1222; Sloss et al. 2008, p. 1249;
Scribner et al. 2008, p. 9). Within Lake
Superior, regional genetic differences
are evident between brook trout
populations in Nipigon Bay, Isle Royale,
and Lake Nipigon-Grand Portage
(Wilson et al. 2008, p. 1313). Adfluvial
brook trout are thought to be the
mechanism providing genetic
communication among these regional
aggregations and straying of a coaster
was documented in Nipigon Bay and at
Isle Royale (D’Amelio et al. 2008, p.
1347; Stott 2008, p. 4). Sloss et al. (2008)
investigated genetic differentiation
among four Wisconsin populations of
stream-resident brook trout. His work
found significant differentiation among
populations to the point the authors
observed that for these populations,
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there appears to be a near complete lack
of gene flow among them resulting in
genetic drift (Sloss et al. 2008, p. 1249).
None of these isolated populations are
thought to currently have adfluvial
ecotypes as part of the population. This
observation is consistent with the
contemporary lack of an adfluvial form
that historically provided the regional
genetic connection for the three
metapopulations previously mentioned.
As characterized in the entire brook
trout species, phenotypic plasticity and
adaptive radiation (Schluter 2000, p. 1)
appear to represent the continuum of
evolutionary processes underlying the
expression of life-history variation in
populations of brook trout in Lake
Superior (Ardren 2008, pp. 1–2). As
stated above, plastic responses allow
individuals to obtain high fitness in new
environments. Alternatively, adaptive
genetic differentiation among
populations may provide evolutionary
advantages. First, there are fitness costs
to being highly plastic. For example,
plastic genotypes need to maintain
sensory and developmental pathways in
order to induce plastic responses that
are not required by nonplastic
genotypes (Relyea 2002, pp. 272–273).
Secondly, if the plastic response to a
new environment is insufficient and
directional selection favors an extreme
phenotype, there will be genetic
evolution of the trait (adaptive
radiation). Therefore, if a population of
brook trout experiences divergent
selection in stable environments, we
would expect the ecotypes to evolve
genetic differences and nonplastic forms
because the cost of maintaining the
phenotypic plasticity would be too high.
Findings in the Salmon Trout River
indicate phenotypic plasticity plays a
major role in the expression of the
adfluvial and fluvial ecotypes while
information from Isle Royale indicates
adaptive radiation has occurred
separating adfluvial and lacustrine
coaster ecotypes. Migratory plasticity
could be favored in situations where
adfluvial and stream-resident brook
trout co-occur because the environments
they occupy are highly variable
(Huckins et al. 2008, p. 1324; Ridgway
2008, pp. 1186–1187). The alternating
selection patterns associated with these
diverse and variable environments
create a fitness advantage for plastic
genotypes over nonplastic genotypes. In
addition, the metapopulation structure
mediated by coaster brook trout
(D’Amelio and Wilson 2008, p. 1222;
Ridgway 2008, p. 1181) favors plasticity
over adaptive genetic differences among
populations because dispersal among
populations increases environmental
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heterogeneity and favors an increase in
trait reaction norm (the pattern of
visible characteristics produced by a
given genetic makeup of an organism
under different environmental
conditions; Sultan and Spencer 2002, p.
281). Alternatively, the adfluvial and
lacustrine ecotypes on Isle Royale are
physically isolated and in this situation,
adaptive radiation would be favored
over the evolution of phenotypic
plasticity (Price 2003, pp. 1437–1438).
If phenotypic plasticity is the source
of differences observed between streamresident and brook trout, then these
ecotypes are expressed in a single
population and represent the extremes
of the reaction norm for migratory
behavior. Scribner et al. (2008, p. 10)
did not observe genetic differences
between sympatric adfluvial brook trout
and presumed stream-resident ecotypes
in the Salmon Trout River on the south
shore of Lake Superior. Analysis of
microsatellite DNA provided high
statistical power to detect genetic
differences between ecotypes. In fact,
the authors did observe highly
significant genetic differences between
brook trout sampled above and below
the impassable waterfall in this system.
In addition, when collections from the
Salmon Trout River were compared
with native brook trout populations
sampled from 10 other nearby
tributaries, the lowest pairwise measure
of genetic distinction was observed
between the resident and adfluvial
ecotypes sampled below the waterfall in
the Salmon Trout River. D’Amelio and
Wilson (2008, p. 1221) used similar
methods to document that adfluvial
brook trout in the Nipigon Bay were not
genetically distinct from presumed
resident brook trout sampled from
tributaries to the bay. These findings in
the Salmon Trout River and the Nipigon
Bay area indicate phenotypic plasticity
likely plays a major role in the
expression of the adfluvial and fluvial
ecotypes.
Theriault et al. (2008, pp. 417–419)
used an eco-genetic model to
demonstrate that intensive harvest of
anadromous fish reduces the probability
of migration in brook trout over the
course of 100 years. This study provides
a basic framework for understanding
how fisheries-induced selection
(mortality from fishing) influences the
evolution of alternate life-history tactics
that are expressed by phenotypic
plasticity. For example, directional
selection imposed by fishing-induced
mortality on coaster brook trout confers
high fitness to the survivors of the
fishery but not necessarily with respect
to natural selection. There is also
uncertainty regarding the rate of
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recovery for expression of the adfluvial
form after fishing selection is reduced or
eliminated because there is not
automatically equal directional
selection in the opposite direction for
expression of the adfluvial form. In the
case of the coaster, habitat degradation
and competition from nonnative salmon
may exclude brook trout from habitats
that would allow juvenile brook trout to
achieve growth rates necessary to
express the adfluvial coaster ecotype
(Huckins et al. 2008, pp. 1337–1339).
Additionally, metapopulation structure
mediated by coaster brook trout
(D’Amelio et al. 2008, p. 1348) favors
plasticity over adaptive genetic
differences among populations (Sultan
and Spencer 2002, p. 281). Loss of
coasters in most populations in Lake
Superior has reduced migration among
populations (Sloss et al. 2008, p. 1249)
resulting in a reduction in
environmental heterogeneity favoring a
decrease in the reaction norm of traits.
These studies demonstrate that humaninduced selective forces can alter the
reaction norm for a population which
can result in the loss of plasticity
needed to express the coaster lifehistory forms.
Brook trout experts contend that if
environmental conditions are suitable
(i.e., threats are abated), the adfluvial
life form of brook trout populations in
Lake Superior can be readily
reconstituted from purely resident stock
(USFWS 2009, p. 8); this is believed
unlikely for other salmonids (e.g.,
Oncorhynchus mykiss). This assertion is
predicated on three premises. First,
adult brook trout of one ecotype may
produce offspring of the other ecotype.
For example, two resident fish could
breed and produce offspring that exhibit
both the adfluvial and fluvial lifehistory strategies. Further, streamresident and adfluvial ecotypes from the
same population interbreed. This means
that within a stream, individuals that
exhibit the resident and adfluvial forms
reside within and are drawn from the
same population. Second, the chars
(genus Salvelinus), including brook
trout, show greater phenotypic plasticity
than most other salmonids. Adfluvial
brook trout do not require substantial
physiological changes (for example,
smoltification) to successfully migrate
and survive in the lake environment.
Thus, the fitness costs to maintain the
genetic code for plasticity are likely less
relative to saltwater-dwelling
salmonids. Hence, it is reasonable to
expect a brook trout population will
maintain the ability (genetic code) to
express the full array of life forms over
time. Third, life-history strategy for
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brook trout is strongly controlled by
environmental conditions or triggers. As
such, the experts believe that, provided
the necessary environmental conditions
or triggers exist, life forms can be
expressed even if temporally lost from
a population.
Current Population Status of Brook
Trout
The current range of native brook
trout remains generally unchanged,
extending through much of eastern
North America, from eastern Canada,
south through the Great Lakes and
northeast to Georgia in the U.S.
However, populations throughout this
range have experienced significant
declines. The current range of native
brook trout started diminishing over the
past 200 years as a result of ecosystem
disruption following European
settlement of North America (Newman
and DuBois 1997). Habitat destruction
by forestry, agricultural practices,
industrial water use, dams, and
pollution were responsible for this
decline (Power 1980, p. 141). Brook
trout were once present in nearly every
coldwater stream and river in the
eastern U.S. and Canada, but
populations began to disappear as early
agriculture, timber, and textile practices
and industries cleared the region’s
protective forests and degraded the
streams with sediment and pollution
(Power 1980, p. 141; EBJV 2006, p. 1).
Throughout much of their natural
range, remaining stream populations
have retreated into extreme headwater,
high elevation, or upstream reaches
(EBJV 2006, p. 2). In the eastern U.S.,
healthy stream populations of brook
trout (wild brook trout occupying 90–
100 percent of their historical habitat)
exist in only 5 percent of subwatersheds
(EBJV 2006, p. 2). Anadromous stocks
along the U.S. coast and in many
Canadian rivers have been decimated by
dams and estuarine pollution (Power
1980, p. 195). In the southern portion of
its range (southern Appalachian
Mountains), brook trout populations
have declined by 75 percent, persisting
now only in isolated headwater reaches
(EBJV 2006, p. 6).
Various threats are persistent across
the brook trout range. Most of them
involve habitat loss and degradation,
such as poor land management, high
water temperature, sedimentation
(roads), urbanization, degraded riparian
habitat, stream fragmentation (roads),
dam inundation/fragmentation, and
forestry practices (EBJV 2006, pp. 3, 5).
Poor land management associated with
agriculture (such as clearing streamside
vegetation, over-grazing sensitive areas,
ineffectively managing nutrients, and
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ditching small streams) ranks as the
most widely distributed impact to brook
trout across the eastern U.S. (EBJV 2006,
p. 2). Climate change presents a
significant threat to brook trout, with
some southern portions predicted to
lose between 53–97 percent of their
brook trout habitat due to high water
temperatures (Flebbe 2006, p. 1379).
While some uncertainty remains about
the exact temperature increase that will
result from climate change, the present
range of brook trout is predicted to
shrink, particularly in the southern
Appalachians (Hudy et al. 2005, p. 5).
Nonnative species are now present
throughout most of the range (Parsons
1973, p. 5). Interactions with these
nonnatives are considered to be among
the most significant biological threats to
brook trout rangewide (Peck 2001, p.13;
Hudy et al. 2005, p. 3; EBJV 2006, pp.
2–3, 5). Brown trout have been shown
to displace or reduce stream
populations of brook trout throughout
their natural range (Nyman 1970, p. 348;
Fausch and White 1981, p. 1226; Waters
1983, p. 144). Encroachment by rainbow
trout has also been documented in the
contraction of the range of native brook
trout across their native range (Kelly et
al., 1980, pp. 9–10; Power 1980, p. 195;
Larson and Moore 1985, p. 200). Species
such as small mouth bass and yellow
perch are considered to be significant
competitors with lake-dwelling brook
trout (EBJV 2006, pp. 22, 28, 34).
Current Population Status of Brook
Trout in the Upper Great Lakes
Brook trout populations throughout
the upper Great Lakes region are
relatively common and geographically
widespread, although distribution and
abundance is much reduced from
historical levels (Power 1980, p. 195;
Becker 1983, pp. 321–322; WIDNR and
USFWS 2005, p. 17). Dramatic declines
in abundance and distribution of both
coaster and stream-resident ecotypes of
brook trout occurred in the upper Great
Lakes from the 1850s to mid-1900s
(Goodier 1982, pp. 110, 112; Ritchie and
Black 1988, p. 15; Newman and Dubois
1997, pp. 4–6; Enterline 2000, p. 1;
WIDNR and USFWS 2005, pp. 17–18;
Schreiner et al. 2008, p. 1305; Schreiner
et al. 2008, p. 1351; Huckins et al. 2008,
p. 1322).
There are presently at least 200
streams with documented brook trout
populations in the upper Great Lakes
(Moore and Bream 1965, p. 19; Goodier
1982, p. 110; Enterline 2000, p. 30;
Newman et al. 2003, pp. 31–37; Quinlan
2004, unpub. data; Bassett 2009, unpub.
data; Ward 2007, p. 16; Schram 2008b,
pers. comm.; Scott 2008, pers. comm.;
Chase 2009, pers. comm.; OMNR 2009,
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unpub. data). The current specific status
of most of these populations is not
known, but they are described by the
Michigan, Minnesota, and Wisconsin
natural resource agencies as stable and
self-sustaining in the upper Great Lakes
(Holtz 2008, p. 2; MIDNR 2008a, p. 49;
Schreiner and Ebbers 2008, pers.
comm.).
In coldwater tributaries to the upper
Great Lakes, brook trout were
historically distributed from the river
mouth upstream to the headwaters or to
impassible barriers (Smith and Moyle
1944, p. 119; Moore and Braem 1965, p.
19; Goodier 1982, p. 111; Becker 1983,
p. 321; WIDNR and USFWS 2005). The
brook trout numbers in these stream
reaches once numbered in the hundreds
to thousands (Huckins and Baker 2008,
p. 1231). A 30-year data set from
Wisconsin tributaries shows that, in
streams historically occupied solely by
brook trout, brook trout have contracted
into upstream sections and are now
nearly absent in lower reaches (WIDNR
2008, unpub. data). Brook trout
abundance has declined despite the
persistence of suitable conditions for
brook trout and high numbers of
juvenile nonnative salmonids (WIDNR
2008, unpub. data). In Wisconsin
tributaries to Lake Superior, the
distribution of stream-resident brook
trout populations has declined by nearly
50 percent from historical levels
(WIDNR and USFWS 2005, p. 17).
Historically, 119 tributaries to Lake
Superior and purportedly 6 Lake Huron
streams supported populations of brook
trout with coaster ecotypes (Newman et
al. 2003, pp. 31–38; Enterline 2000, p.
30). Once abundant and widespread
throughout the northern portions of the
Great Lakes, populations of brook trout
that still exhibit the coaster ecotypes are
presently limited to a few locations
(Dehring and Krueger 1985, p. 1; Bailey
and Smith 1981, p. 1549; Goodyear et al.
1982, pp. 63–65; Enterline 2000, p. 30;
Newman et al. 2003, p. 39; Schreiner et
al. 2008, p. 1351; Mucha and Mackereth
2008, p. 1). Although self-sustaining
populations of stream-resident brook
trout are currently present in 56 of 58
U.S. streams and in all 61 Canadian
streams identified in the Brook Trout
Rehabilitation Plan for Lake Superior as
historically supporting populations with
coaster ecotypes (Newman et al. 2003,
pp. 31–37; Quinlan 2008, unpub. data;
Schreiner 2008, pers. comm.; Schram
2008c, pers. comm.; Scott 2008, pers.
comm.; Chase 2009, pers. comm.), only
18 populations with coaster ecotypes
still persist there (15 stream-spawning–
adfluvial, and 3 lake-spawning–
lacustrine) (Goodyear 1982, pp. 63–65;
Quinlan 1999, p. 19; Ritchie and Black
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1988, p. 15; Swainson 2001, p. 41;
Newman et al. 2003, pp. 28–39;
Enterline 2000, p. 30; Chase 2009, pers.
comm.).
Over the last decade, the presence of
coaster brook trout has been confirmed
in other locations within the upper
Great Lakes. Surveys, and in some cases
genetic analysis, have confirmed the
presence of brook trout with coaster
ecotypes in the following locations;
Minnesota tributaries to Lake Superior
(Newman et al. 1999, p. 2; BurnhamCurtis 2000, p. 4; Pranckus and
Ostazeski 2003, p. 5; Ward 2007, p. 16),
three Michigan tributaries to Lake
Superior (Stimmel 2006, p. 56; MIDNR
2008a, p. 2; Leonard 2009, pers. comm.),
along the shoreline of the Red Cliff
Indian Reservation, Wisconsin (Stott
and Quinlan 2008, p. 21), and in Little
Todd Harbor and Rock Harbor, Isle
Royale (Gorman et al. 2008, p. 1257).
The origin of these fish is unknown and
natural reproduction of fish exhibiting
the coaster ecotype has not been
confirmed, therefore these locations are
not identified as supporting selfsustaining populations. However, they
have potential to be self-sustaining
populations, as outlined by Schreiner et
al. (2008).
Abundance of individuals in
populations exhibiting the coaster
ecotypes is stable or increasing in
several regions of Lake Superior. In the
Salmon Trout River, Michigan,
abundance as determined by video
surveillance increased from 118 to 243
in the period from 2004 to 2006 (MIDNR
2008a, p. 6). In the Nipigon River, angler
catch per hour has increased from the
late 1980s to the present, while harvest
has decreased substantially (Houle
2004, p. 13). In South Bay, Lake
Nipigon, estimates of spawner
abundance continue to increase and
currently number about 600 fish—up
from fewer than 100 in the recent past,
but still fewer than the estimated 2,500
present in the mid-1900s (Swainson
2009, pers. comm.). In Tobin Harbor,
Isle Royale National Park, Michigan,
estimates of adult brook trout from
1996, 2001, and 2008 has remained
around 200–250 fish (USFWS
unpublished data). Relative abundance
based on shoreline electrofishing index
surveys in Tobin Harbor from 1997 to
2008 has fluctuated from 0.3 per hour to
16.7 per hour (USFWS 2008, unpub.
data).
There are reintroduction stocking
efforts ongoing in several streams on the
Grand Portage Indian Reservation
(Newman and Johnson 1996, p. 4), Red
Cliff Indian Reservation, Keweenaw Bay
Indian Community Reservation
(Donofrio 2002, p. 1), and in Whittlesey
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Creek, Wisconsin (USFWS and WIDNR
2003, p. 5). Supplementation stocking
occurred in Siskiwit Bay, Isle Royale,
from 1999 to 2005. Data collected to
date indicates limited success with
these efforts (Newman et al. 1999, p. 2;
Quinlan 2008, pers. comm.; Stott and
Quinlan 2008, p. 22). Reintroduction
efforts in Michigan have recently been
terminated in the Gratiot, Little Carp,
Hurricane, and Mosquito Rivers and
Sevenmile Creek (Scott 2007, pers.
comm.; Loope 2007, pers. comm.).
Threats to brook trout across its native
range are also acting on brook trout
within the upper Great Lakes. A primary
impact is the presence of introduced
fishes (e.g., non-native salmonids).
Introduced salmonids have competitive
and predatory impacts on brook trout,
although the precise mechanisms may
not be fully understood and the
magnitude of impact may vary by
species, population size, and
environmental conditions. The decline
or loss of the migratory coaster form has
diminished connectivity among
populations that once operated as
metapopulations. Populations that occur
in such isolated patches can be lost,
increasing the possibility of extirpation.
As a species, brook trout are known to
be highly susceptible to exploitation by
anglers (Newman and Dubois 1996, p. 3;
Newman et al. 2003, p. 11; Huckins et
al. 2008, p. 1322). Overharvest was a
primary cause of the decline of Great
Lakes brook trout populations by the
early 1900s, especially the coaster
ecotype, and continues to threaten some
populations within the region (Newman
and Dubois 1996, p. 1; Huckins et al.
2008, p. 1322; Schreiner et al. 2008, p.
1356). Climate change also presents a
threat to upper Great Lakes brook trout,
through increased water temperatures,
leading to increased presence of
nonnative competitors and predators
along with a decrease in habitat
suitability. Although the enormous
coldwater reservoir within the lake
environment represents a potential
refuge for Great Lakes brook trout,
predicted impacts in both stream and
lake environments still represent a
potential threat to their long-term
viability.
Defining a Species Under the Act
Section 3(16) of the Act defines
‘‘species’’ to include ‘‘any species or
subspecies of fish and wildlife or plants,
and any distinct vertebrate population
segment of fish or wildlife that
interbreeds when mature’’ (16 U.S.C.
1532 (16)). Our implementing
regulations at 50 CFR 424.02 provide
further guidance for determining
whether a particular taxon or
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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
previously discussed, coaster brook
trout are classified as Salvelinus
fontinalis, the same as other brook trout,
and as such we do not consider the
coaster form of the brook trout to
constitute a distinct species or
subspecies. Since the coaster brook trout
is not a distinct species or subspecies,
we then evaluated whether the coaster
brook trout is a distinct vertebrate
population segment to determine
whether it would constitute a listable
entity under the Act.
To interpret and implement the
distinct vertebrate population segment
(DPS) provisions of the Act and
Congressional guidance, the Service and
the National Marine Fisheries Service
(now the National Oceanic and
Atmospheric Administration—
Fisheries), published the Policy
Regarding the Recognition of Distinct
Vertebrate Population Segments (DPS
Policy) in the Federal Register on
February 7, 1996 (61 FR 4722). Under
the DPS Policy, three elements are
considered in the decision regarding the
establishment and classification of a
population of a vertebrate species as a
possible DPS. These are applied
similarly for additions to and removals
from the List of Endangered and
Threatened Wildlife and Plants. These
elements are (1) the discreteness of a
population in relation to the remainder
of the species to which it belongs, (2)
the significance of the population
segment to the species to which it
belongs, and (3) the population
segment’s conservation status in relation
to the Act’s standards for listing,
delisting, or reclassification.
Distinct Vertebrate Population Segment
Analysis
In accordance with our DPS Policy,
this section details our analysis of the
first two elements used to assess
whether a vertebrate population
segment under consideration for listing
may qualify as a DPS. These elements
are (1) the population segment’s
discreteness from the remainder of the
species to which it belongs and (2) the
significance of the population segment
to the species to which it belongs.
Discreteness refers to the ability to
circumscribe a population segment from
other members of the taxon based on
either (1) physical, physiological,
ecological, or behavioral factors or (2)
international boundaries that result in
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significant differences in control of
exploitation, habitat management,
conservation status, or regulatory
mechanisms in light of section 4(a)(1)(B)
of the Act.
Under our DPS Policy, if we have
determined that a vertebrate population
segment is discrete, we consider its
biological and ecological significance to
the larger taxon to which it belongs 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. To
evaluate whether a discrete vertebrate
population may be significant to the
taxon to which it belongs, we consider
the best available scientific evidence.
This evaluation may include, but is not
limited to: (1) Evidence of the
persistence of the discrete population
segment in an ecological setting that is
unusual or unique for the taxon; (2)
evidence that loss of the population
segment would result in a significant
gap in the range of the taxon; (3)
evidence that the population segment
represents the only surviving natural
occurrence of a taxon that may be more
abundant elsewhere as an introduced
population outside its historical range;
and (4) evidence that the discrete
population segment differs markedly in
its genetic characteristics from other
populations of the species.
The first step in our DPS analysis was
to identify population segments of the
brook trout to evaluate. The petition
asked us to (1) ‘‘list as ‘endangered’ the
naturally spawning anadromous (lakerun) Coaster Brook Trout (Salvelinus
fontinalis) throughout its known
historic range in the conterminous
United States’’ (including designation of
critical habitat) and (2) ‘‘determine
whether the Salmon Trout River (STR)
coaster is a DPS’’ and (3) ‘‘whether the
south shore of Lake Superior population
of coasters (which are known to breed
today only in the STR) is ‘endangered.’ ’’
Although brook trout in the Great Lakes
exhibit three life-history forms (fluvial,
adfluvial, and lacustrine), the petition
specifically focused on the coaster, or
adfluvial and lacustrine, forms.
To address the entity identified in the
first petition request (coaster brook trout
throughout their historical range in the
U.S.), we identified two approaches to
analyzing a potential population
segment: (1) Describe and analyze an
upper Great Lakes ‘‘all brook trout’’
population segment, which includes all
brook trout life forms—fluvial,
adfluvial, and lacustrine ecotypes,
inclusive of coaster brook trout—present
throughout the documented historical
range of brook trout in the Great Lakes
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basin, and (2) describe and analyze an
upper Great Lakes ‘‘coaster-only’’
population segment, which includes
only the coaster forms—adfluvial and
lacustrine ecotypes—of brook trout
throughout the documented historical
range of brook trout in the Great Lakes
basin.
We find that neither of the population
segments analyzed constitute a valid
DPS, and therefore the first petitioned
entity, coaster brook trout throughout
their historical range in the U.S., is not
a valid DPS. To address the second and
third petition requests, we focused on
the brook trout population in the
Salmon Trout River and evaluated
whether it qualified as a DPS per our
policy. We find that the brook trout
population in the Salmon Trout River
also does not constitute a valid DPS.
The remainder of this section details the
evaluation of these population segments
as DPSs per our 1996 DPS Policy.
Upper Great Lakes All Brook Trout
Population Segment
This population segment
encompasses the range of brook trout
populations within the Great Lakes
basin that currently or historically
occupied both the tributary and lake
environments (including streamresident, adfluvial, and lacustrine
ecotypes of brook trout). Although
technically not one of the ‘‘Great
Lakes,’’ we include Lake Nipigon in
Canada in this population because it is
part of the Great Lakes drainage. The
best available information indicates the
known historical range of brook trout
within the basin included all of Lake
Superior and its drainage (including
Lake Nipigon), and the northern
portions of Lakes Michigan and
Huron—specifically, that portion of
Lake Michigan north of a line from the
Sheboygan River, Wisconsin to Grand
Traverse Bay, Michigan, and that
portion of Lake Huron north of Thunder
Bay, Michigan, eastward to include
Manitoulin Island to the 81°30′
longitudinal demarcation and west of
81°30′ longitude (MacCrimmon and
Campbell 1969, p. 1701; Dehring and
Krueger 1985, p. 1; Enterline 2000, pp.
29–30).
23383
population segment are physically
isolated from other populations of brook
trout as the result of the physical
separation between the drainage of the
Great Lakes basin and neighboring
drainages. Consequently, brook trout in
the Great Lakes basin meet the
discreteness criterion of being markedly
separate from other members of the
brook trout taxon.
International Border
We presently do not find that the
brook trout in the Upper Great lakes on
either side of the international United
States border with Canada are discrete
due to differences in control of
exploitation, management of habitat,
conservation status, or regulatory
mechanisms that are significant in light
of section 4(a)(1)(D) of the Act.
Conclusion for Discreteness
In conclusion, we determine that the
Upper Great Lakes brook trout
population segment, as defined here, is
discrete from the remainder of the brook
trout taxon. This discreteness arises
from the population segment’s physical
isolation from the remainder of the
taxon. Therefore, we will now consider
the potential significance of this discrete
population segment to the remainder of
the taxon.
Significance
We have determined that the
population of brook trout in the Upper
Great Lakes meets the discreteness
elements of the DPS policy, and as such,
we will now evaluate whether this
specific population is significant to the
taxon as a whole (i.e., native brook trout
in eastern North America). A discrete
population is considered significant
under the DPS policy if it meets one of
four of the elements identified in the
policy under significance or can
otherwise be reasonably justified as
being significant.
We discuss further below our
evaluation of the significance of the
population of brook trout in the Upper
Great Lakes relative to the taxon as a
whole.
Marked Separation
Evidence of the Persistence of the
Discrete Population Segment in an
Ecological Setting That Is Unusual or
Unique for the Taxon
As previously described, the Upper
Great Lakes brook trout population
segment we have evaluated
encompasses the range of brook trout
populations that currently or
historically occupied both the tributary
and lake environments within the Great
Lakes basin. Brook trout within this
On the basis of an evaluation of the
best available scientific information, we
have determined that the habitat for
brook trout in the Upper Great Lakes
does not represent an ecological setting
that is unusual or unique for the native
brook trout relative to the habitat
available to it throughout the entire
Discreteness
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taxon’s range in eastern North America.
A summary of our evaluation is below.
Brook trout exhibiting differing lifehistory forms occupy a variety of
ecosystems from subarctic regions of the
Hudson Bay coast, to temperate areas
bordering and east of the Great Lakes,
and southern coldwater habitats in the
Appalachian Mountains of Tennessee
and Georgia (Power 1980, p. 142). They
have been successfully naturalized in
western North America, South America,
Eurasia, Africa, and New Zealand
(MacCrimmon and Campbell 1969, p.
1699, pp. 1703–1717). Within their large
native range in eastern North America,
brook trout habitat includes coastal
areas and various-sized lakes, streams,
and rivers at varying altitudes. Most
populations inhabit coldwater streams,
but lake-dwelling and lake-spawning
(lacustrine form) populations also occur
throughout the range, in spring-fed
ponds, small- to medium-sized lakes,
and a few large, oligotrophic (containing
relatively little plant life or nutrients,
but rich in dissolved oxygen) lakes.
Anadromous populations (‘‘salters’’) of
brook trout use marine habitats in
Hudson Bay and along the Atlantic
coast.
The upper Great Lakes represent a
complex ecological setting for brook
trout. The very large size of the Great
Lakes watershed creates an environment
that more closely resembles oceanic
physical conditions (available to the
anadromous forms of brook trout) than
conditions in smaller lakes (available to
other forms of brook trout). With
approximately 1,500 tributaries and
almost 2,800 miles (4,506 km) of
shoreline, Lake Superior also provides
brook trout access to a very large
freshwater habitat network. Although
the Great Lakes are the largest
freshwater water bodies occupied by
brook trout, there are thousands of lakes
in its range including large postglacial
lakes further north in Canada that
contain populations of the adfluvial and
lacustrine forms (e.g., Fraser and
Bernatchez 2008, p. 1193).
If predicted rising water temperatures
in response to climate change are
realized over the entire range of brook
trout, the distributions of brook trout
populations would probably shift
toward cooler waters at higher latitudes
and altitudes (Meisner 1990b, p. 1068;
Magnuson et al. 1997, p. 859; Kling et
al. 2003, pp. 53–54). The greatest effects
would likely begin in populations
located at the margins of the taxon’s
hydrologic and geographic distributions
(Meisner et al. 1990a, p. 282, Kling et al.
2003, p. 54). Although the upper Great
Lakes have already experienced some
impacts of climate change (see Kling et
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al. 2003, pp. 14–16) and will not be
immune to future impacts (see Kling et
al. 2003, pp. 21–25), they may provide
substantial coldwater habitat for brook
trout in the future. However, brook trout
have abundant coldwater habitat
available in the northern latitudes of its
range, and habitat in northern North
America which is presently too cold
may develop into appropriate brook
trout habitat under a warming scenario.
We will further evaluate the extent that
this may be the case in the range-wide
assessment of native brook trout that we
plan to conduct (see Finding section).
Although the upper Great Lakes
represent a diverse and complex
ecological setting which may offer
potential coldwater habitat for brook
trout, we must evaluate the breadth of
ecological diversity of brook trout
habitat rangewide in our assessment of
this population segment’s significance
to the rest of the taxon. First, available
information indicates that the large area
and wide geographical range of brook
trout habitats, which vary in latitude
and altitude and water form, contain a
vast diversity of habitats for brook trout.
The ecological setting of the upper Great
Lakes is a small portion of the brook
trout range, and based on available
information, its relative significance to
the brook trout species is limited.
Second, although we expect that the
Great Lakes may offer substantial
coldwater habitat, there are other large,
deep, oligotrophic lakes, and numerous
lakes and streams at higher latitudes
that may buffer the species from
potential climate change impacts. Given
the available information on the
diversity and extent of ecological
settings of brook trout in the rest of its
range, we conclude at this time that the
upper Great Lakes is a not unique or
unusual setting of significance for the
native brook trout in eastern North
America.
Evidence That Loss of the Population
Segment Would Result in a Significant
Gap in the Range of the Taxon
Loss of brook trout, including any or
all life forms, in the upper Great Lakes,
when considered in relation to brook
trout throughout the remainder of the
species’ range in eastern North America,
would mean the loss of a small
geographic portion (approximately ten
percent) of the entire range of the taxon.
Further, the number of streams with
populations in the upper Great Lakes
(about 200) are a small proportion of the
amount of streams and lakes with brook
trout populations in the rest of the
native range in eastern North America.
Due to the broad geographic range of
brook trout, the wide diversity of
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habitats available to it, and its plasticity,
and the fact that the upper Great Lakes
are at the western periphery of its
natural range, we find that the gap in
the range resulting from the loss of
brook trout in the upper Great Lakes
would not be significant.
Evidence That the Population Segment
Represents the Only Surviving Natural
Occurrence of a Taxon That May Be
More Abundant Elsewhere as an
Introduced Population Outside Its
Historical Range
This criterion from the DPS policy
does not apply to the brook trout in the
upper Great Lakes because it is not a
population segment representing the
only surviving natural occurrence of the
taxon that may be more abundant
elsewhere as an introduced population
outside its historical range.
Consequently, this population of brook
trout does not meet the significance
element of this factor.
Evidence That the Discrete Population
Segment Differs Markedly in Its Genetic
Characteristics From Other Populations
of the Species
A large amount of rangewide genetic
variation for brook trout is distributed
among brook trout populations (large
Fst values, values in a fixation index
which describe the degree of population
differentiation based on genetic
polymorphisms). This pattern is heavily
influenced by the ecological and lifehistory characteristics of brook trout
populations (population connectivity or
isolation, philopatric tendency).
We find that, based on the genetic
information currently available
(outlined under the Brook Trout
Genetics section above), the brook trout
in the upper Great Lakes, including all
life forms, do not differ markedly from
other populations of the species in their
genetic characteristics (such as
exhibiting unique alleles or a proportion
of genetic variability beyond the norm
of distribution) such that they should be
considered biologically or ecologically
significant based simply on genetic
characteristics. They do not show any
more genetic distinctiveness in
comparison to the remainder of the
taxon than other populations
demonstrate. With the additional
consideration that the authority to list
DPSs be used ‘‘sparingly,’’ we conclude
that this population segment of brook
trout does not meet the significance
element of this factor.
DPS Conclusion—Upper Great Lakes All
Brook Trout Population Segment
On the basis of the best available
information, we conclude that the all-
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brook-trout population segment in the
Upper Great Lakes is discrete due to
marked separation as a consequence of
physical, ecological, physiological, or
behavioral factors according to the 1996
DPS Policy. However, on the basis of an
evaluation of brook trout in the Great
Lakes relative to the four significance
elements of the 1996 DPS Policy, we
conclude that this discrete population
segment is not significant to the taxon
to which it belongs, and therefore, does
not qualify as a DPS under 1996 policy.
As such, we find that population of
brook trout in the Great Lakes basin is
not a listable entity under the Act.
Upper Great Lakes Coaster-Only Brook
Trout Population Segment
This population segment
encompasses the historical range of
brook trout populations in the Great
Lakes basin exhibiting the coaster
ecotypes, which includes northern
portions of the Lakes Michigan and
Huron and all of Lake Superior,
including Lake Nipigon (see
Discreteness analysis for the Upper
Great Lakes All Brook Trout Population
Segment below for more detailed range
description).
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Discreteness
Hubbs and Lagler (1949, p. 44) and
Becker (1983, p. 320) described coasters
as brook trout that spend a portion of
their life cycle in the Great Lakes.
Coaster brook trout have long been
recognized by local and scientific
communities (Newman and Dubois
1997, p. 4).
Marked Separation
As described previously, coasters are
adfluvial and lacustrine life forms of
brook trout that occupy the nearshore
zone of the Great Lakes. Coasters, being
a subset of brook trout within the Great
Lakes basin, are markedly separate from
all other brook trout outside of the Great
Lakes Basin as the result of the physical
separation between the drainage of the
Great Lakes basin and neighboring
drainages. Thus, brook trout within this
population segment are markedly
separate from other members of the
brook trout taxon outside the Great
Lakes basin because they are physically
isolated.
Isolation also exists within the Great
Lakes basin, among brook trout
populations in Lakes Huron, Michigan,
Erie, and Ontario. The best available
information indicates that adfluvial
brook trout likely did not historically
occupy lake waters of southern Lakes
Michigan and Huron (boundary as
previously defined in this section) or
Lakes Erie and Ontario (MacCrimmon
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and Campbell 1969, p. 1700; Bailey and
Smith 1981, p. 1549; Dehring and
Krueger 1985, p. 1; MIDNR 2008a, pp.
2–3). Brook trout found within these
lake areas in the last 100 years are likely
the result of stocking as no known
adfluvial, migratory or lake dwelling
populations exist. The reason that brook
trout never occupied these lake areas is
unknown; we suspect that unidentified
environmental conditions preclude
brook trout use of these habitats.
Regardless, without brook trout use of
the lake environment, natural dispersal
between stream populations cannot
occur. This absence of adfluvial and
lacustrine ecotypes in these populations
effectively restricts populations with
coaster brook trout forms to the
distribution previously defined, namely
the watershed and lake habitats of all of
Lake Superior, and the northern
portions of Lakes Michigan and Huron.
Within the Great Lakes basin, coasters
are ecologically, behaviorally, and
physiologically discrete from streamresident brook trout. Coasters are
markedly separate from resident brook
trout in their lake-dwelling and
adfluvial behavior (Hubbs and Lagler
1949, p. 44; Becker 1983, p. 320;
Huckins and Baker 2008, p. 1229;
Schreiner et al. 2008, p. 1350). Lakedwelling coasters spend their entire life
within the lake environment (Huckins et
al. 2008, p. 1323; Schreiner et al. 2008,
p. 1350); adfluvial coasters move
between streams and the lake (Huckins
et al. 2008, p. 1323). Stream-resident
brook trout remain within the river
system. These differences mark an
ecological (i.e., lake versus stream
habitat) and a behavioral (i.e.,
migratory) separation between the two
forms.
Coaster ecotypes and stream-resident
ecotypes of brook trout also differ
physiologically in adult size, longevity,
age at maturity, and fecundity. As stated
in the Species Description section
above, adult coasters range in size from
12 to 25 in (30 to 64 cm), and commonly
reach lengths of 16 in (41 cm) (Ritchie
and Black 1988, pp. 50–51; Quinlan
1999, p. 17; Huckins and Baker 2008, p.
1239; Huckins et al. 2008, p. 1337). The
body mass of adult coasters typically
ranges from 0.75 to 8 pounds (341 to
3632 g) (Quinlan 1999, p. 16; Swainson
2001, p. 60; Huckins and Baker 2008, p.
1239; WIDNR and USFWS 2005, p. 16)
with a maximum measurement of 14.5
pounds (6577 g) (Scott and Crossman
1973, p. 211). Adult resident brook trout
typically range in size from 5 to 15 in
(13 to 38 cm) (Scott and Crossman 1979,
p. 208; Becker 1983, pp. 318, 320;
WIDNR and USFWS 2005, p. 16;
Schram 2008a pers. comm.) and usually
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23385
weigh less than a pound (<454 g)
(WIDNR and USFWS 2005, p. 16). Most
female coasters do not reach maturity
until they are 2 to 4 years old and 12
to 15 in. (30 to 38 cm) in length (Ritchie
and Black 1998, p. 19; Quinlan 1999, p.
11; Huckins and Baker 2008, p. 1241;
Huckins et al. 2008, p. 1329), and live
5 to 8 years (Quinlan 1999, p. 11;
Huckins et al. 2008, p. 1328). Whereas
most female stream-resident brook trout
mature by age 1 or 2 (Becker 1983, p.
318), and typically live to age 3 and
rarely reach ages of 4 or 5 years (Scott
and Crossman 1973, p. 211, Becker
1983, p. 318). Coaster females produce
around 1,500 to 3,000 eggs (Quinlan
1999, p. 20; Swainson 2001, p. 41),
while stream-resident brook trout
fecundity ranges from 100 to 1,500 eggs
per female (Scott and Crossman 1973, p.
210; Power 1980, p. 157; Becker 1983,
p. 318).
We recognize that many of the
ecological, physiological, and
behavioral characteristics discussed
here are influenced to varying extents
by environmental factors. For example,
fish exhibit indeterminate growth,
where adults can reach larger sizes in
larger habitats with more favorable
growth conditions or greater prey
availability, but may be more
diminutive under less favorable habitat
conditions (Huckins et al. 2008, p.
1323). To this effect, many physiological
characteristics of coasters would be
expected to differ from their streamresident counterparts, with coasters
being larger than residents, simply
because coasters access the more
productive lake environments. In
addition, many of the characteristics we
evaluate are interrelated, with one
characteristic influencing or
determining one or more of the other
characteristics. For example, fecundity
is largely a function of the size and
condition of the fish. Also, prey
selection will be influenced by the prey
availability in different habitat types.
We rely on all the characteristics taken
together to describe the phenotypic
characteristics of each type. Regardless
of the source of the phenotypic
characteristics of the types, be they
controlled by genetic heritability,
environmental influences, or both, they
accumulate to form a description of
each form and that defines either their
similarity or separation.
We further recognize that upper Great
Lakes brook trout display a continuum
of traits in most of the characteristics
described. However, the range of
overlap is small in comparison to the
broader range of difference between the
two forms, with the majority of adult
coasters and stream-residents clearly
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occupying nonoverlapping portions of
the continuum. Further, at the end of
the continuum of traits, coasters are
markedly separate in their use of Great
Lakes habitat. As we stated in adopting
the DPS Policy in 1996, ‘‘logic demands
a distinct population recognized under
the Act be circumscribed in some way
that distinguishes it from other
representatives of its species. The
standard established for discreteness is
simply an attempt to allow an entity
given DPS status under the Act to be
adequately defined and described’’ (61
FR 4721, at 4724; February 7, 1996). In
the case of brook trout in the Great
Lakes, there is a group that can be
clearly distinguished by a variety of
characteristics, particularly its use of the
Great Lakes habitat, which leads to or
results from marked separation in the
other characteristics.
Despite the apparent reproductive
exchange and genetic similarity between
stream-resident forms and coaster forms
of brook trout, the life forms remain
markedly separated physiologically,
ecologically, and behaviorally. The DPS
Policy states that ‘‘the standard adopted
[for discreteness] does not require
absolute separation of a DPS from other
members of its species, because this can
rarely be demonstrated in nature for any
population of organisms * * * [T]he
standard adopted allows for some
limited interchange among population
segments considered to be discrete, so
that loss of an interstitial population
could well have consequences for gene
flow and demographic stability of a
species as a whole’’ (61 FR 4722;
February 7, 1996). Coasters are a group
of organisms that can be distinguished
from stream-resident brook trout by a
variety of characteristics, particularly its
migratory life strategy and use of the
Great Lakes.
Thus, given marked separation in
physical, physiological, ecological, and
behavioral factors, we conclude that the
coaster-only population segment is
discrete from Great Lakes streamresident brook trout. Further, as stated
above, given its marked separation from
all other brook trout outside of the Great
Lakes Basin as the result of the physical
separation between the drainage of the
Great Lakes basin and neighboring
drainages, the coaster-only population
segment is discrete from brook trout
outside the Great Lakes basin.
Consequently, we find that the coasteronly population satisfies the element of
marked separation under the 1996 DPS
Policy, and is therefore considered to be
a discrete population per our policy.
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International Border
We presently do not find that this
population segment of the brook trout
on either side of the international
United States border with Canada is
discrete due to differences in control of
exploitation, management of habitat,
conservation status, or regulatory
mechanisms that are significant in light
of section 4(a)(1)(D) of the Act.
Significance
We must next evaluate whether the
coaster brook trout population segment
is significant to the larger brook trout
taxon. We find that, although we
determined that coaster brook trout are
a discrete population segment, they cooccur with and are a subset of the same
population as other brook trout types
(stream residents) in the upper Great
Lakes (see Species Information section
above). Review of the best available
scientific information does not suggest
that the coaster and resident life forms
in these populations are genetically
distinct from each other, indicating that
they are part of one breeding population
(D’Amelio and Wilson 2008, p. 1221;
Scribner et al. 2008, p. 10). Thus,
similar to our Upper Great Lakes All
Brook Trout population segment, the
loss of coasters would not create a
significant gap in the range of the taxon,
they are not the only remaining natural
occurrence of the taxon, and they do not
show significant genetic distinctiveness
in comparison to the remainder of the
taxon. In addition, coasters occupy a
smaller portion of the same ecological
setting as other brook trout in the upper
Great Lakes. Although, as discussed
above, coasters may be important to the
long-term viability of brook trout
populations throughout Lake Superior,
the relevant question is whether
coasters are significant to the taxon as
a whole, here, all native brook trout.
Given this, the significance analysis
documented for the all brook trout
population segment (see Upper Great
Lakes All Brook Trout DPS section
above) also applies to the coaster-only
population segment, and we similarly
conclude that the coaster-only
population segment does not meet the
significance elements of the DPS Policy.
DPS Conclusion—Coaster-Only
Population Segment
On the basis of the best available
information, we conclude that the
coaster-only population segment in the
Upper Great Lakes is discrete due to
marked separation as a consequence of
physical, ecological, physiological, or
behavioral factors according to the 1996
DPS policy. However, on the basis of the
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four significance elements in the 1996
DPS Policy, we conclude that this
discrete population segment is not
significant to the rest of the taxon, and
therefore, does not qualify as a valid
DPS under our 1996 DPS Policy. As
such, we find that the coaster-only
population in the upper Great Lakes is
not a listable entity under the Act.
Salmon Trout River/South Shore Lake
Superior Brook Trout Population
Segment
This section evaluates whether the
Salmon Trout River-South Shore Lake
Superior brook trout population
segment qualifies as a DPS. Since the
Salmon Trout River contains the only
known brook trout population with
naturally reproducing coaster on the
South Shore of Lake Superior, we
addressed these two petition requests in
one analysis.
Discreteness
Markedly Separate
The brook trout population segment
that occupies the Salmon Trout River is
markedly separate from other members
of the brook trout taxon because it is
genetically or reproductively isolated.
This physical isolation is supported by
recent evidence from Scribner et al.
(2008, pp. 12–13), which found no
genetic evidence of Salmon Trout River
fish in neighboring streams, indicating
that Salmon Trout River coasters are not
a source of gene flow among streams.
International Border
Since the Salmon Trout River
population segment does not cross an
international border, this basis for
finding discreteness is not applicable.
In conclusion, the Salmon Trout River
brook trout population segment, as
defined here, meets the element for
discreteness under our 1996 DPS Policy
and is considered discrete from the
remainder of the brook trout taxon. This
discreteness arises from the population
segment’s genetic or reproductive
isolation from the remainder of the
taxon which is supported by evidence of
genetic discontinuity.
Significance
Evidence of the Persistence of the
Discrete Population Segment in an
Ecological Setting That Is Unique for the
Taxon
The ecological setting for the Salmon
Trout River discrete population segment
is similar to that of other brook trout
populations throughout the upper Great
Lakes region. We are unaware of any
features that make the Salmon Trout
River unique or unusual in terms of
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brook trout habitat. There is nothing
about the ecological setting that is
unique or unusual for the species,
particularly in light of the other
occurrences within Lake Superior.
Consequently, this population of brook
trout does not meet the significance
element of this factor.
Evidence That Loss of the Population
Segment Would Result in a Significant
Gap in the Range of the Taxon
This criterion from the DPS policy
does not apply to the Salmon Trout
River discrete population segment
because this population is one of
thousands of brook trout populations
existing throughout the range of the
taxon and its loss would represent an
extremely small portion of the range.
Consequently, this population of brook
trout does not meet the significance
element of this factor.
populations is reflective of the
reproductive connections (isolation)
among the populations across the range
of the taxon.
We are unaware of any information
indicating that this population segment
differs from the species in its genetic
characteristics (such as exhibiting
unique alleles or a proportion of genetic
variability beyond the norm of
distribution) such that it should be
considered biologically or ecologically
significant to the taxon based on genetic
characteristics. Consequently, this
population of brook trout does not meet
the significance element of this factor.
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Evidence That the Population Segment
Represents the Only Surviving Natural
Occurrence of a Taxon That May Be
More Abundant Elsewhere as an
Introduced Population Outside Its
Historical Range
This criterion from the DPS policy
does not apply to the Salmon Trout
River discrete population segment
because it is not a population segment
representing the only surviving natural
occurrence of the taxon that may be
more abundant elsewhere as an
introduced population outside its
historical range. Consequently, this
population of brook trout does not meet
the significance element of this factor.
DPS Conclusion—Salmon Trout River/
South Shore Lake Superior Population
Segment
On the basis of the best available
information, we conclude that the
Salmon Trout River brook trout
population segment is ‘‘markedly
separated’’ from all other populations of
the same taxon as a consequence of
physical factors, supported by genetic
evidence. Consequently, the Service
concludes that the petitioned entity is
discrete according to the 1996 DPS
Policy. However, on the basis of an
evaluation of the four significance
elements of the 1996 DPS Policy, we
conclude that this discrete population
segment is not significant to the species
to which it belongs. Therefore, we find
that the Salmon Trout River brook trout
population does not qualify as a DPS
under our DPS Policy and is
consequently not a listable entity under
the Act.
Evidence That the Discrete Population
Segment Differs Markedly in Its Genetic
Characteristics From Other Populations
of the Species
Scribner et al. (2008, p. 9) indicates
that Lake Superior brook trout
populations, including the Salmon
Trout River, are highly genetically
structured with low levels of gene flow
among populations. The Salmon Trout
River contains two genetically distinct
populations that are separated by
impassable waterfalls (Scribner et al.
2008, p. 10). Both populations in the
Salmon Trout River were equally
genetically diverged from the other
populations included in the study
(Scribner et al. 2008, p. 7). This pattern
of population genetic structuring is
common in brook trout throughout the
species’ range because, like many
salmonids, this species likely exhibits
some degree of spawning site fidelity
(Angers et al. 1999, p. 1044; D’Amelio
et al. 2008, pp. 1347–1348; Mucha and
Mackereth 2008, p. 1211). This degree of
genetic divergence that forms among
Significant Portion of the Range
Analysis
The Act defines an endangered
species as one ‘‘in danger of extinction
throughout all or a significant portion of
its range,’’ and a threatened species as
one ‘‘likely to become an endangered
species within the foreseeable future
throughout all or a significant portion of
its range.’’ Having determined that the
northern Great Lakes population
segment of brook trout and the Salmon
Trout River/South Shore Lake Superior
populations of the coaster brook trout
do not meet the elements of our 1996
DPS Policy as being valid DPSs, we then
assessed whether the upper Great Lakes
brook trout is a significant portion of the
range (SPR) of the native brook trout
where the species is in danger of
extinction or likely to become so in the
foreseeable future.
On March 16, 2007, a formal opinion
was issued by the Solicitor of the
Department of the Interior, ‘‘The
Meaning of ‘In Danger of Extinction
Throughout All or a Significant Portion
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23387
of Its Range’ ’’ (DOI 2007). We have
summarized our interpretation of that
opinion and the underlying statutory
language below. A portion of a species’
range is significant if it is part of the
current range of the species and is
important to the conservation of the
species because it contributes
meaningfully to the representation,
resiliency, or redundancy of the species.
The contribution must be at a level such
that its loss would result in a decrease
in the ability of the species to persist.
The first step in determining whether
a species is endangered in an SPR is to
identify any portions of the range of the
species that warrant further
consideration. The range of a species
can theoretically be divided into
portions in an infinite number of ways.
However, there is no purpose to
analyzing portions of the range that are
not reasonably likely to be significant
and threatened or endangered. To
identify those portions that warrant
further consideration, we determine
whether there is substantial information
indicating that (i) the portions may be
significant and (ii) the species may be in
danger of extinction there. In practice, a
key part of this analysis is whether the
threats are geographically concentrated
in some way. If the threats to the species
are essentially uniform throughout its
range, no portion is likely to warrant
further consideration. Moreover, if any
concentration of threats applies only to
portions of the range that are
unimportant to the conservation of the
species, such portions will not warrant
further consideration.
The petition specified two portions of
the range of brook trout: (1) The
historical range of coaster brook trout in
the contiguous U.S., namely the upper
Great Lakes, and (2) the Salmon Trout
River/South Shore Lake Superior. In our
SPR analysis, we assessed threats to
brook trout in these portions in
comparison to threats acting on other
portions of the range. Information on
threats within the upper Great Lakes
region included primarily habitat
degradation, overutilization, nonnative
fishes, and loss of connectivity and lifehistory diversity. We had comparatively
less detailed information on the threats
acting throughout the rest of the range.
The best information available to us
regarding other portions of the brook
trout range was found in analyses
completed for the Eastern Brook Trout
Joint Venture (see Hudy et al. 2005, TU
2006). Given the information available
to us on threats to brook trout across its
range, we conclude that threats to this
species were similar throughout its
range, that the conservation status of the
species is similar throughout its range,
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Federal Register / Vol. 74, No. 95 / Tuesday, May 19, 2009 / Proposed Rules
and that there is no area within the
range of the upper Great Lakes and the
Salmon Trout River/South Shore Lake
Superior portions of the coaster brook
trout where potential threats to this
species are significantly concentrated or
are substantially greater than in other
portions of the range. We found no
evidence that more threats were
geographically concentrated within the
upper Great Lakes than in any other part
of the range; according to the findings
of Hudy et al. (2005), it seems that
threats may be greater in portions of the
Northeastern U.S. populations than in
the Great Lakes.
Therefore, we find that the brook trout
is not threatened or endangered solely
in any significant portion of its range
within the upper Great Lakes. As stated
in the Finding section below, we plan
to initiate a range-wide assessment of
the native brook trout that will enable
us to better understand the status of the
native brook trout across the range of
species, including a determination of
whether the threats to the species,
which are not concentrated in the upper
Great Lakes, warrant listing the native
brook trout rangewide.
erowe on PROD1PC63 with PROPOSALS-1
Finding
In making this finding, we considered
information provided by the petitioners,
as well as other information available to
us concerning coaster brook trout. We
have carefully assessed the best
scientific and commercial information
available regarding the status of and
threats to coaster brook trout in the
upper Great Lakes. We reviewed the
petition, and available published and
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unpublished scientific and commercial
information. We also consulted with
Federal and State land managers, along
with recognized experts in conservation
and population genetics and brook trout
and salmonid biology. This 12-month
finding reflects and incorporates
information that we received from the
public following our 90-day finding or
that we obtained through consultation,
literature research, and field visits.
On the basis of this review, we have
determined that the coaster brook trout
in the upper Great Lakes does not meet
the elements of our 1996 DPS Policy as
being a valid DPS. We also find that the
coaster brook trout is not an SPR of the
native brook trout and does not warrant
further consideration as such under the
Act. Therefore, we find that the coaster
brook trout is not a listable entity under
the Act, and that listing is not
warranted.
Although we find that population
segments analyzed above are not listable
entities, we found enough information
concerning the diversity, habitats,
population structure, threats, and trends
of the native brook trout in its entire
range to initiate a range-wide
assessment that will enable us to better
understand the status of the native
brook trout across the range of species.
Completing a range-wide assessment
will allow us to better evaluate if any
population would meet the elements of
the DPS policy or constitute an SPR of
the taxon. We will also continue to
assess the status of and threats to both
the upper Great Lakes and Salmon Trout
River/South Shore Lake Superior
populations of the coaster brook trout.
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We request that you submit any new
information concerning the taxonomy,
biology, ecology, and status of the brook
trout in its entire native range. Send this
information to the Region 3 Fish and
Wildlife Service Regional Office (see
ADDRESSES section) 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 with new
information as appropriate. The Service
continues to strongly support the
cooperative conservation and
restoration of the coaster brook trout in
the upper Great Lakes.
References
A comprehensive list of the
referenced materials is available upon
request (see ADDRESSES section above).
Author
The primary authors of this document
are staff located at the Region 3 Fish and
Wildlife Service Regional Office (see
ADDRESSES).
Authority
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Stephen Guertin,
Acting Deputy Director, U.S. Fish and Wildlife
Service.
[FR Doc. E9–11527 Filed 5–18–09; 8:45 am]
BILLING CODE 4310–55–P
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Agencies
[Federal Register Volume 74, Number 95 (Tuesday, May 19, 2009)]
[Proposed Rules]
[Pages 23376-23388]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: E9-11527]
=======================================================================
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[FWS-R3-ES-2008-0030; 92210-1111-0000-FY09-B3]
Endangered and Threatened Wildlife and Plants; 12-Month Finding
on a Petition To List the Coaster Brook Trout as Endangered
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of 12-month petition finding.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), announce a
12-month finding on a petition to list the coaster brook trout
(Salvelinus fontinalis) as endangered under the Endangered Species Act
of 1973, as amended (Act). The petition also asked that critical
habitat be designated for the species. After review of all available
scientific and commercial information, we find that the coaster brook
trout is not a listable entity under the Act, and therefore, listing is
not warranted. We ask the public to continue to submit to us any new
information that becomes available concerning the taxonomy, biology,
ecology, and status of coaster brook trout and to support cooperative
conservation of coaster brook trout within its historical range in the
Great Lakes.
DATES: The finding announced in this document was made on May 19, 2009.
ADDRESSES: This finding is available on the Internet at https://www.regulations.gov at Docket Number [FWS-R3-ES-2008-0030]. Supporting
documentation for this finding is available for inspection, by
appointment, during normal business hours at the U.S. Fish and Wildlife
Service, Region 3 Fish and Wildlife Service Regional Office, 1 Federal
Drive, Bishop Henry Whipple Federal Building, Fort Snelling, MN 55111.
Please submit any new information, materials, comments, or questions
concerning this finding to the above address, Attention: Coaster brook
trout.
FOR FURTHER INFORMATION CONTACT: Jessica Hogrefe, Region 3 Fish and
Wildlife Service Regional Office (see ADDRESSES) (telephone 612-713-
5346; facsimile 612-713-5292). Persons who use a telecommunications
device for the deaf (TDD) may call the Federal Information Relay
Service (FIRS) at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Background
Section 4(b)(3)(B) of the Act (16 U.S.C. 1531 et seq.) requires
that, for any petition to revise the Lists of Endangered and Threatened
Wildlife and Plants that contains substantial scientific and commercial
information that listing may be warranted, we make a finding within 12
months of the date of our receipt of the petition on whether the
petitioned action is: (a) Not warranted, (b) warranted, or (c)
warranted, but the immediate proposal of a regulation implementing the
petitioned action is precluded by other pending proposals to determine
whether species are threatened or endangered, and expeditious progress
is being made to add or remove qualified species from the List of
Endangered and Threatened Species. Section 4(b)(3)(C) of the Act
requires that we treat a petition for which the requested action is
found to be warranted but precluded as though resubmitted on the date
of such finding, that is, requiring that we make a subsequent finding
within 12 months. Such 12-month findings must be published in the
Federal Register. This notice constitutes our 12-month finding for the
petition to list the U.S. population of coaster brook trout.
Previous Federal Action
The Sierra Club Mackinac Chapter, Huron Mountain Club, and Marvin
J. Roberson filed a petition, dated February 22, 2006, with the
Secretary of the Interior to list as endangered the ``naturally
spawning anadromous (lake-run) coaster brook trout throughout its known
historic range in the conterminous United States'' and to designate
critical habitat under the Act. The petition clearly identified itself
as such and included the requisite identification information for the
petitioners, as required in 50 CFR 424.14(a). On behalf of the
petitioners, Peter Kryn Dykema, Secretary of the Huron Mountain Club,
submitted supplemental information, dated May 23, 2006, in support of
the original petition. This supplemental information provided further
information on the species' status and biology, particularly for brook
trout in the Salmon Trout River.
On September 13, 2007, we received a 60-day notice of intent to sue
over the Service's failure to determine, within 1 year of receiving the
petition, whether the coaster brook trout warrants listing. Under
section 4 of the Act, the Service is to make a finding, to the maximum
extent practicable within 90 days of receiving a petition, that it does
or does not present substantial scientific or commercial information
indicating that the petitioned action may be warranted. Further, the
Act requires that, within 12 months of receiving a petition found to
present substantial information, the Service must determine whether the
petitioned action is warranted. A complaint was filed in U.S. District
Court in the District of Columbia on December 17, 2007, for failure to
make a timely finding (Sierra Club, et al. v. Kempthorne, No. 1:07-cv-
02261 (D.D.C. December 17, 2007)). The Service reached a negotiated
settlement with the plaintiffs to submit the 90-day finding to the
Federal Register by March 15, 2008. We published a ``substantial'' 90-
day finding March 20, 2008. The negotiated settlement further required
the Service to publish the 12-month finding in the Federal Register by
December 15, 2008. The deadline for the 12-month finding was extended
to April 15, 2009, by mutual consent. On April 15, 2009, we filed an
unopposed motion to extend the deadline for the coaster brook trout 12-
month finding to May 12, 2009.
Species Information
Species Description
Brook trout (Salvelinus fontinalis), also called brook char or
speckled trout, is one of three species in the genus Salvelinus (chars)
native to north and eastern North America; the others being lake trout
(S. namaycush) and Arctic char (S. alpinus). The chars are a sub-group
of fishes in the salmon and trout subfamily (Salmoninae) that is
distinct from the ``true'' trout and salmon sub-groups.
The brook trout throughout its range in eastern North America
exhibits considerable variation in growth rate, color, and other
features, but generally can be distinguished from other char and trout
species by its olive-green to dark brown back with a light yellow-brown
vermiculate pattern, sides with large yellow-brown spots and blue halos
surrounding small, sporadic red and orange spots. Pectoral, pelvic,
anal, and lower caudal fin have leading edges of white bordered by
black with the
[[Page 23377]]
remainder predominantly reddish to orange. Sea-run brook trout become
silver with purple iridescence and show red spots on the sides (Scott
and Crossman 1973, p. 208).
Distribution
The historical range of native brook trout extends along Hudson Bay
in Canada across the Provinces of Manitoba, Ontario and Quebec, to
Newfoundland and Labrador and south to Nova Scotia and New Brunswick in
Canada; and from eastern Iowa through northern Illinois, northern Ohio,
and the Great Lakes drainage (Minnesota, Michigan, Wisconsin), through
the New England States (New York, New Hampshire, Vermont, Maine,
Massachusetts, Pennsylvania, New Jersey), large New England rivers
(such as the Hudson River and Connecticut River), and through the
Appalachian Mountains in Maryland, Virginia, West Virginia, North
Carolina, South Carolina, Tennessee, south to Georgia (MacCrimmon and
Campbell 1969, pp. 1700-1702; MacCrimmon et al. 1971, p. 452; Scott and
Crossman 1973, pp. 209-210; Power 1980, p. 142). Naturalized
populations of brook trout were established as early as the late 1800s
beyond the historical native range by introductions to waters in
western North America, South America, Eurasia, Africa, and New Zealand
(MacCrimmon and Campbell 1969, p. 1699, pp. 1703-1717). The current
range of native brook trout still extends through Canada and down to
Georgia in the U.S., but in many locations, populations have been
completely extirpated or have contracted within this range towards
upper stream reaches, higher altitudes, or headwaters (EBJV 2006, p.
2).
Distribution of Brook Trout in the Great Lakes
According to Bailey and Smith (1981, p. 1549) and MacCrimmon and
Campbell (1969, p. 1701), brook trout are native to the lakes and
tributaries of Lakes Superior, Huron, Michigan, and the tributaries of
Lakes Erie and Ontario. Brook trout are not believed to have been
present in Minnesota streams above barrier falls to Lake Superior
(Smith and Moyle 1944, p. 119) or throughout most of the lower
peninsula of Michigan (MIDNR 2008a, pp. 1-2; MacCrimmon and Campbell
1969, p. 1704).
Habitat Requirements
Brook trout require clear, cold, well-oxygenated water to thrive.
They are generally found in water ranging between 41-68[deg] Fahrenheit
(5-20[deg] Celsius), with their likely preferred temperature falling
near the middle of this range (Power 1980, p. 172). Thermal
requirements within this range vary by life cycle phase and season
(Scott and Crossman 1973, p. 211; Blanchfield and Ridgway 1997, p. 750;
Baril and Magnan 2002, pp. 177-178).
The brook trout spawns in late summer or autumn, the date varying
with latitude and temperature. Spawning takes place most often over
gravel beds but may be successfully accomplished over a variety of
substrates if there is spring upwelling or a moderate current (Scott
and Crossman 1973, p. 210). Power (1980, p. 151) describes rangewide
brook trout spawning, which occurs in the fall, when day length and
temperature are decreasing. In northerly regions and at high
elevations, brook trout may spawn as early as late August and spawning
may be delayed until December in southern areas. As is typical for
salmonids, females prepare redds (hollows scooped out for spawning) in
suitable gravel substrate. The female then deposits her eggs in the
redd where they are fertilized by a male. After spawning there is no
further parental involvement with the young. The redd protects the eggs
and allows an adequate exchange of dissolved gases and other materials
during development.
Brook trout are carnivorous, feeding opportunistically upon a
variety of prey, such as worms, leeches, crustaceans, aquatic insects,
terrestrial insects, spiders, mollusks, and fish (Scott and Crossman
1973, p. 212). Anadromous (migrating from salt water to spawn in fresh
water) forms vary their feeding behavior and prey items based on their
age and the environment, marine or riverine, they are occupying (Newman
and Dubois 1997, p. 9). Brook trout also show diverse foraging
behaviors; some individuals may be sedentary, eating crustaceans from
the lower portion of the water column, whereas others in the same
system may be more active and eat insects from the upper portion of the
water column (McLaughlin et al. 1999, p. 386). This resource
polymorphism may play a supplementary role in the extensive adaptive
radiation (evolution of ecological variability within a rapidly
multiplying lineage; Smith and Sk[uacute]lason 1996) observed in this
species.
Genetics of Brook Trout
A large amount of genetic variation for brook trout is
distributed among populations (large Fst values). This pattern is
heavily influenced by the diverse ecological and life-history
characteristics of brook trout populations (population connectivity
or isolation, philopatric tendency). This pattern of highly
differentiated populations of brook trout is found at small and
large geographic scales. Population genetic structuring is common in
brook trout throughout its range (Angers et al. 1999, pp. 1049-
1050). Like many salmonids, brook trout tend to have a hierarchical
population structure resulting from the hierarchical design of the
networks of streams and lake or coastal areas in which they live,
and a complicated life cycle that leads to strong local adaptations.
Taxonomic resolution can be even more complicated at the lake level
when lakes include sympatric (occupying the same or overlapping
geographic area without interbreeding) but genetically divergent
brook trout populations such as in Lake Mistassini in Canada (Fraser
and Bernatchez 2008, p. 1197). This degree of genetic divergence
that forms among populations is reflective of the reproductive
connections (isolation) among the populations across the range of
the taxon.
Six distinct genetic mitochondrial (mtDNA) clades have been
identified throughout the range of brook trout in eastern North America
(Danzmann et al. 1998, p. 1307). These mtDNA clades reflect historical
isolation in glacial refugia or long periods of isolation in nonglacial
areas in the southern part of the species' range. The Wisconsin glacial
advance which covered portions of Canada covered all five Great Lakes
15,000 years ago (Bailey and Smith 1981, p. 1543). As these glaciers
receded, brook trout recolonized the lakes from the Mississippi and
Atlantic refugia (Danzmann et al. 1998, pp. 1308, 1312). Given this
pattern of glaciation, genetic diversity is greatest at the southern
portion of the species' range and gradually decreases northward
(Danzmann et al. 1998, pp. 1310-1311). As the most geographically
isolated (for tens of thousands of years), brook trout in the southern
part of the species' range (along the Appalachian Mountains south to
Georgia) are the most diverse, containing all six mtDNA clades. The
Great Lakes contains three of the six mtDNA clades. Throughout the
northern portion of their range in Canada, brook trout are the least
genetically diverse, with only a single mtDNA clade present. Within
each of these lineages, there is evidence to suggest that selection is
driving rapid phenotypic divergence in some populations.
Results based on microsatellite DNA variation identified nine
distinct genetic assemblages of brook trout in the U.S. (King 2009,
unpub. data). Assemblages from the nonglacial southern part of the
species' range (along the Appalachian Mountains from Pennsylvania to
Georgia) in the U.S. are the most genetically divergent, and this
divergence among the assemblages generally decreases as the range
progresses northward.
[[Page 23378]]
Genetics of Brook Trout in the Great Lakes
Populations from Lake Superior and tributaries to Lake Erie form
two of the nine genetic assemblages of brook trout in the U.S. The Lake
Erie populations are the most divergent assemblage from the northern
part of the species' range. Lake Superior populations are similar in
the degree of genetic divergence to the remaining northern assemblages
grouping with the average genetic distance between brook trout
populations in the U.S. Samples from the rest of the Great Lakes were
not available for analysis. Although brook trout in the Great Lakes do
not contain any wholly unique mtDNA clades, they do contain a large
amount of the genetic variation in a confined portion of the range
(Danzmann et al. 1998, pp. 1310-1311).
Native populations of brook trout in Lake Superior in most cases
have retained their native genetic characteristics despite the stocking
of hatchery fish from sources outside and within the Lake Superior
basin. In Lake Superior, the intensity and purpose of stocking has
varied over time and space. For example, Minnesota tributaries to Lake
Superior have been stocked with hatchery strains that originated from
outside of the Great Lakes Basin to provide fishing opportunities above
fish passage barriers (Wilson et al. 2008, p. 1312). Until the early
1990s, most of the stocked fish in Lake Superior were domesticated
strains from outside the Great Lakes basin (Schreiner et al. 2008, p.
1357), although many stocking events were undocumented and records of
early stocking events are incomplete (Wilson et al. 2008, p. 1312).
These stocking efforts were not targeted at rehabilitation and from
that perspective, results were poor. The stocked fish were not
behaviorally or evolutionarily adapted to the environment in which they
were planted, criteria known to limit survival and reproductive success
(Schreiner et al. 2008, p. 1357). Burnham-Curtis (2001, p. 2) concluded
that hatchery fish have had little reproductive success in Lake
Superior streams based on her examination of 36 tributaries to Lake
Superior and 9 hatchery stocks outplanted into the lake. However, the
genetic methods used by Burnham-Curtis provided low power to detect
genetic introgression of hatchery fish into native populations (Wilson
et al. 2008, p. 1312). A recent study by D'Amelio and Wilson (2008, p.
1215) used genetic methods with high power to detect genetic
introgression of hatchery fish into natural populations. This study
documented only low levels of genetic introgression of Lake Nipigon
hatchery fish into native populations of brook trout from six
tributaries to Lake Superior's Nipigon Bay (D'Amelio and Wilson 2008,
p. 1222), despite decades of stocking. A study by Scribner et al.
(2006, pp. 3-4) examined nine brook trout populations from Lake
Superior tributaries on the south shore of Michigan and four hatchery
strains outplanted into those tributaries. This study used similar
methods to D'Amelio and Wilson (2008). Scribner et al. (2006, p. 8)
concluded that hatchery stocking appears to have minimal if any impact
of on brook trout.
Brook Trout Life-History Diversity
An individual's ability to produce multiple phenotypes (visible or
observable characteristics) in response to its environment is termed
phenotypic plasticity (Scheiner 1993, p. 36). Recent studies have
recognized the role of phenotypic plasticity as a major source of
phenotypic variation in natural populations (Price et al. 2003, p.
1438). The brook trout exhibits remarkable phenotypic plasticity across
its natural range. This plasticity allows it to thrive in a variety of
environments, from cold subarctic regions, through temperate zones and
in southern refugia in eastern North America, and in a range of places
where it has been introduced (Power 1980, p. 142). Although primarily a
stream-dwelling species, brook trout also occupy inland lakes and
coastal waters. Because of the variety of the freshwater, estuary, and
ocean environments, migratory plasticity is also favored. The brook
trout's dispersal subsequent to receding glaciation, and separation
into isolated breeding stocks in diverse habitats subject to an array
of natural and man-made influences have all contributed to this
variability (Power 1980, p. 142).
Brook trout display considerable life-history variation throughout
their native range (Huckins and Baker 2008, p. 1229). Brook trout
across its range exhibit a variety of life-history types (polymorphisms
or ecotypes), including fluvial (stream-dwelling), adfluvial (migrating
between lakes and streams), lacustrine (lake-dwelling), and anadromous
(migrating from salt water to spawn in fresh water) forms.
Understanding life-history diversity in a species requires knowledge of
the evolutionary history, ecological setting, and reproductive
relationships among ecotypes. Reproductive interactions between
ecotypes are reflected by the magnitude and pattern of genetic
differentiation observed between life-history phenotypes at neutral
genetic markers. The expression of migratory behavior (expressed as the
adfluvial and anadromous ecotypes) by any individual fish will be
partially in direct response to its environment. Phenotypic expression
of more than one form may be expected in a population located in a
variable environment containing habitats for several ecotypes. The
amount of phenotypic plasticity a population will exhibit for the
migratory trait also has a heritable genetic basis and will be
determined by the intensity and type of selective pressures that
population experiences (Via and Lande 1985, pp. 517-519; Theriault et
al. 2008, pp. 418-419).
Adoption of migratory adfluvial form or stream-resident life-
history form in brook trout has been modeled under a conditional
strategy framework where environmentally influenced threshold traits
determine which ecotype a fish will adopt (Hendry et al. 2004, pp. 124-
125). Growth rate efficiencies, body size, and concentration of
juvenile hormone have all been identified as potential threshold traits
(Theriault and Dodson 2003, pp. 1155-1157). Theoretical work by Ridgway
(2008, p. 1185) and Uller (2008, pp. 436-437) also provide information
to suggest parental effects are important to the expression of
alternate ecotypes of brook trout. These parental effects describe an
affect of the parental phenotype on the offspring's phenotype such as
coaster females producing larger eggs and spawning in different
locations from stream-resident ecotypes, influencing the habitat use
(Morinville and Rasmussen 2006, pp. 701-702) and growth rate at the
juvenile stage (Perry et al. 2005, p. 1358). These differences in
growth rate and habitat use impact potential threshold traits.
Work on sympatric brook trout life forms at young ages largely
comes from a few studies on anadromous populations. Morinville and
Rasmussen (2003) studied the bioenergetics of young brook trout
exhibiting anadromous migratory and stream-resident life tactics. They
found that the anadromous migrants have higher metabolic costs and had
consumption rates 1.4 times that of stream residents but growth
efficiencies of the anadromous form were lower than that of residents.
Spatial utilization of habitat differed among the life tactics as well,
with migratory individuals occupying faster-flowing waters compared to
the resident fish which used pool areas (p. 408). They concluded that
migrant brook trout have noticeably different energy budgets than
resident brook trout from the same system (p. 406). Morinville and
Rasmussen (2008) also investigated morphological differences between
life
[[Page 23379]]
tactics. The authors concluded that migrant brook trout were found to
be more streamlined (narrower and shallower bodies) than resident brook
trout, and these differences persisted into the marine life of the
migrant fish (pp. 175, 183). The differences were powerful enough to
derive discriminant functions using five of the measured traits
allowing for accurate classification of juvenile brook trout as either
migrant or resident with an overall correct classification rate of 87
percent.
A study by Theriault et al. (2007b, p. 61) found that sympatric
anadromous and fluvial brook trout in the Sainte-Marguerite River in
Quebec belonged to a single gene pool. Phenotypic plasticity is,
therefore, a major force driving the expression of these two life
histories from this population. Evolution of phenotypic plasticity in
this population was influenced by mating systems with most of the
mating between different morphotypes occurring between fluvial males
and anadromous females. Additional work in this system demonstrated
significant heritability for life-history tactic and for body size
(Theriault et al. 2007a, pp. 7-8) indicating expression of life-history
tactic in this population can be effected by natural or artificial
selection.
Life-History Diversity in Great Lakes Brook Trout
Fish that complete their life cycle exclusively in tributaries to
the Great Lakes exhibit the fluvial life history and are defined as
stream residents. ``Coaster'' (the subject of the petition) is a
regional term for a life-history variant of brook trout in the Great
Lakes (Burnham-Curtis 2001, p. 2; Wilson et al. 2008, p. 1) which use
lake waters of the Great Lakes for all or a portion of its life cycle
(Becker 1983, p. 320). The coaster form can be further divided into an
adfluvial ecotype that migrates from the stream to the lake and back
into tributaries to spawn and a lacustrine ecotype that completes its
life cycle entirely within the lake (Huckins et al. 2008, p. 1323). In
the Great Lakes region, spawning usually occurs from mid-September
through mid-November. Distinct life histories associated with the
coaster and stream-resident types result in different physical,
demographic, and ecological characteristics for the forms (Huckins et
al. 2008, p. 1337; Huckins and Baker 2008, p. 1241; Ridgway 2008, p.
1185). Specifically, coasters tend to live longer than stream residents
(5-8 years versus less than 5 years), reach maturation later (females
at 2-4 years versus 1-2 years), attain larger length and weight as
adults (12-25 inches and 0.75-8 pounds (30-64 centimeters (cm) and 341-
3632 grams (g)) versus (5-15 inches (13-38 cm) and (less than 1 pound
(<454 g), be more fecund (1500-3000 eggs per female versus 100-1500
eggs per female), and move greater distances (up to 19-217 miles (30-
350 kilometers (km)) versus less than 19 miles (30 km)) (Scott and
Crossman 1973, pp. 208, 210, 211; Power 1980, p. 157; Becker 1983, pp.
318, 320; Ritchie and Black 1988, pp. 19, 50, 51; Quinlan 1999, pp. 11,
12, 14, 16, 17, 20; Swainson 2001, pp. 40, 41, 60, 64; WIDNR and USFWS
2005, p. 16; Huckins and Baker 2008, pp. 1239, 1241; Huckins et al.
2008, pp. 1328, 1329, 1337; Mucha and Mackereth 2008, p. 1210; Schram
2008a, pers. comm.; Chase 2008, pers. comm.).
Coasters have been historically documented in Lakes Superior,
Huron, and Michigan brook trout populations (Bailey and Smith 1981, p.
1549; Dehring and Krueger 1985, p. 1; Enterline 2000, p. 1; MIDNR
2008a, pp. 1-2). However, Lake Superior is the only Great Lake with
extant coaster forms of brook trout, and all available literature is
from this area. Coasters in the Great Lakes are found in Canada and the
U.S. in substantially fewer locations than they were historically
(Newman et al. 2003, p. 39). Populations in the Great Lakes basin with
these life-history forms are documented within Canada in tributaries to
Nipigon and Black Bays, the Nipigon River, Lake Nipigon and the Pancake
River in the eastern part of Lake Superior (Newman et al. 2003, p. 39;
Chase and Swainson 2009, pers. comm.). Within the U.S. portion of the
Great Lakes basin, populations that express the coaster form occur in
Isle Royale National Park in Tobin Harbor, Big and Little Siskiwit
Rivers, and Washington Creek as well as on the south shore of Lake
Superior in the Salmon Trout River (Newman et al. 2003, p. 39).
As previously stated, brook trout populations within the upper
Great Lakes exhibit fluvial, adfluvial, and lacustrine life-history
forms, coasters comprising the latter two forms. Populations of brook
trout in Lake Superior likely function as types of metapopulations,
with the coaster life forms serving as dispersers (D'Amelio and Wilson
2008, p. 1222; Sloss et al. 2008, p. 1249). The viability of a
metapopulation is strongly contingent upon maintaining dispersal among
populations. Although brook trout exhibit spawning site fidelity,
individuals exhibiting the adfluvial life forms in Lake Superior have
also been shown to stray or disperse among streams (D'Amelio and Wilson
2008, p. 1222; Mucha and Mackereth, p. 1211). The long-term persistence
of a metapopulation requires a balance between local extinction and
recolonization of constituent populations (see Hanski 1998 for a review
of metapopulations). Dispersing individuals offset local population
extinction by providing a means for recolonization (Brown and Kodric-
Brown 1977, p. 448; Reeves et al. 1995, p. 340). Dispersing individuals
also provide for gene flow among discrete populations, countering
losses of genetic fitness while still allowing the development and
distribution of unique adaptive traits (Ingvarsson 2001, p. 63; Tallmon
et al. 2004, p. 494). Thus, the coaster life-history forms are
important to the long-term viability of brook trout populations
throughout Lake Superior.
Genetic studies of stream-resident (fluvial life form) brook trout
show substantial genetic structuring among populations in Michigan,
Wisconsin, Minnesota, and Canada characterized by distinct regional
groupings or metapopulations (Burnham-Curtis 1996, pp. 10-11; Burnham-
Curtis 2001, p. 10; Sloss et al. 2008, p. 1249; Wilson et al. 2008, p.
1312; Scribner et al. 2008, p. 9). In studies aimed at determining
genetic differences between the coaster polymorphism and stream-
resident fish occupying tributaries connected to the lake, molecular
genetic work in Lake Superior indicates that coasters and stream-
resident brook trout occupying tributaries to the first barrier are
parts of the same population (D'Amelio and Wilson. 2008, p. 1221;
Scribner et al. 2008, p. 9; Stott 2008, p. 5). Work investigating the
genetic differences of various tributaries to the lake found distinct
differences among populations of brook trout in each tributary to Lake
Superior (Burnham-Curtis 1996, p. 10; Burnham-Curtis 2000, p. 7;
Burnham-Curtis 2001, p. 10; D'Amelio and Wilson 2008, p. 1222; Sloss et
al. 2008, p. 1249; Scribner et al. 2008, p. 9). Within Lake Superior,
regional genetic differences are evident between brook trout
populations in Nipigon Bay, Isle Royale, and Lake Nipigon-Grand Portage
(Wilson et al. 2008, p. 1313). Adfluvial brook trout are thought to be
the mechanism providing genetic communication among these regional
aggregations and straying of a coaster was documented in Nipigon Bay
and at Isle Royale (D'Amelio et al. 2008, p. 1347; Stott 2008, p. 4).
Sloss et al. (2008) investigated genetic differentiation among four
Wisconsin populations of stream-resident brook trout. His work found
significant differentiation among populations to the point the authors
observed that for these populations,
[[Page 23380]]
there appears to be a near complete lack of gene flow among them
resulting in genetic drift (Sloss et al. 2008, p. 1249). None of these
isolated populations are thought to currently have adfluvial ecotypes
as part of the population. This observation is consistent with the
contemporary lack of an adfluvial form that historically provided the
regional genetic connection for the three metapopulations previously
mentioned.
As characterized in the entire brook trout species, phenotypic
plasticity and adaptive radiation (Schluter 2000, p. 1) appear to
represent the continuum of evolutionary processes underlying the
expression of life-history variation in populations of brook trout in
Lake Superior (Ardren 2008, pp. 1-2). As stated above, plastic
responses allow individuals to obtain high fitness in new environments.
Alternatively, adaptive genetic differentiation among populations may
provide evolutionary advantages. First, there are fitness costs to
being highly plastic. For example, plastic genotypes need to maintain
sensory and developmental pathways in order to induce plastic responses
that are not required by nonplastic genotypes (Relyea 2002, pp. 272-
273). Secondly, if the plastic response to a new environment is
insufficient and directional selection favors an extreme phenotype,
there will be genetic evolution of the trait (adaptive radiation).
Therefore, if a population of brook trout experiences divergent
selection in stable environments, we would expect the ecotypes to
evolve genetic differences and nonplastic forms because the cost of
maintaining the phenotypic plasticity would be too high. Findings in
the Salmon Trout River indicate phenotypic plasticity plays a major
role in the expression of the adfluvial and fluvial ecotypes while
information from Isle Royale indicates adaptive radiation has occurred
separating adfluvial and lacustrine coaster ecotypes. Migratory
plasticity could be favored in situations where adfluvial and stream-
resident brook trout co-occur because the environments they occupy are
highly variable (Huckins et al. 2008, p. 1324; Ridgway 2008, pp. 1186-
1187). The alternating selection patterns associated with these diverse
and variable environments create a fitness advantage for plastic
genotypes over nonplastic genotypes. In addition, the metapopulation
structure mediated by coaster brook trout (D'Amelio and Wilson 2008, p.
1222; Ridgway 2008, p. 1181) favors plasticity over adaptive genetic
differences among populations because dispersal among populations
increases environmental heterogeneity and favors an increase in trait
reaction norm (the pattern of visible characteristics produced by a
given genetic makeup of an organism under different environmental
conditions; Sultan and Spencer 2002, p. 281). Alternatively, the
adfluvial and lacustrine ecotypes on Isle Royale are physically
isolated and in this situation, adaptive radiation would be favored
over the evolution of phenotypic plasticity (Price 2003, pp. 1437-
1438).
If phenotypic plasticity is the source of differences observed
between stream-resident and brook trout, then these ecotypes are
expressed in a single population and represent the extremes of the
reaction norm for migratory behavior. Scribner et al. (2008, p. 10) did
not observe genetic differences between sympatric adfluvial brook trout
and presumed stream-resident ecotypes in the Salmon Trout River on the
south shore of Lake Superior. Analysis of microsatellite DNA provided
high statistical power to detect genetic differences between ecotypes.
In fact, the authors did observe highly significant genetic differences
between brook trout sampled above and below the impassable waterfall in
this system. In addition, when collections from the Salmon Trout River
were compared with native brook trout populations sampled from 10 other
nearby tributaries, the lowest pairwise measure of genetic distinction
was observed between the resident and adfluvial ecotypes sampled below
the waterfall in the Salmon Trout River. D'Amelio and Wilson (2008, p.
1221) used similar methods to document that adfluvial brook trout in
the Nipigon Bay were not genetically distinct from presumed resident
brook trout sampled from tributaries to the bay. These findings in the
Salmon Trout River and the Nipigon Bay area indicate phenotypic
plasticity likely plays a major role in the expression of the adfluvial
and fluvial ecotypes.
Theriault et al. (2008, pp. 417-419) used an eco-genetic model to
demonstrate that intensive harvest of anadromous fish reduces the
probability of migration in brook trout over the course of 100 years.
This study provides a basic framework for understanding how fisheries-
induced selection (mortality from fishing) influences the evolution of
alternate life-history tactics that are expressed by phenotypic
plasticity. For example, directional selection imposed by fishing-
induced mortality on coaster brook trout confers high fitness to the
survivors of the fishery but not necessarily with respect to natural
selection. There is also uncertainty regarding the rate of recovery for
expression of the adfluvial form after fishing selection is reduced or
eliminated because there is not automatically equal directional
selection in the opposite direction for expression of the adfluvial
form. In the case of the coaster, habitat degradation and competition
from nonnative salmon may exclude brook trout from habitats that would
allow juvenile brook trout to achieve growth rates necessary to express
the adfluvial coaster ecotype (Huckins et al. 2008, pp. 1337-1339).
Additionally, metapopulation structure mediated by coaster brook trout
(D'Amelio et al. 2008, p. 1348) favors plasticity over adaptive genetic
differences among populations (Sultan and Spencer 2002, p. 281). Loss
of coasters in most populations in Lake Superior has reduced migration
among populations (Sloss et al. 2008, p. 1249) resulting in a reduction
in environmental heterogeneity favoring a decrease in the reaction norm
of traits. These studies demonstrate that human-induced selective
forces can alter the reaction norm for a population which can result in
the loss of plasticity needed to express the coaster life-history
forms.
Brook trout experts contend that if environmental conditions are
suitable (i.e., threats are abated), the adfluvial life form of brook
trout populations in Lake Superior can be readily reconstituted from
purely resident stock (USFWS 2009, p. 8); this is believed unlikely for
other salmonids (e.g., Oncorhynchus mykiss). This assertion is
predicated on three premises. First, adult brook trout of one ecotype
may produce offspring of the other ecotype. For example, two resident
fish could breed and produce offspring that exhibit both the adfluvial
and fluvial life-history strategies. Further, stream-resident and
adfluvial ecotypes from the same population interbreed. This means that
within a stream, individuals that exhibit the resident and adfluvial
forms reside within and are drawn from the same population. Second, the
chars (genus Salvelinus), including brook trout, show greater
phenotypic plasticity than most other salmonids. Adfluvial brook trout
do not require substantial physiological changes (for example,
smoltification) to successfully migrate and survive in the lake
environment. Thus, the fitness costs to maintain the genetic code for
plasticity are likely less relative to saltwater-dwelling salmonids.
Hence, it is reasonable to expect a brook trout population will
maintain the ability (genetic code) to express the full array of life
forms over time. Third, life-history strategy for
[[Page 23381]]
brook trout is strongly controlled by environmental conditions or
triggers. As such, the experts believe that, provided the necessary
environmental conditions or triggers exist, life forms can be expressed
even if temporally lost from a population.
Current Population Status of Brook Trout
The current range of native brook trout remains generally
unchanged, extending through much of eastern North America, from
eastern Canada, south through the Great Lakes and northeast to Georgia
in the U.S. However, populations throughout this range have experienced
significant declines. The current range of native brook trout started
diminishing over the past 200 years as a result of ecosystem disruption
following European settlement of North America (Newman and DuBois
1997). Habitat destruction by forestry, agricultural practices,
industrial water use, dams, and pollution were responsible for this
decline (Power 1980, p. 141). Brook trout were once present in nearly
every coldwater stream and river in the eastern U.S. and Canada, but
populations began to disappear as early agriculture, timber, and
textile practices and industries cleared the region's protective
forests and degraded the streams with sediment and pollution (Power
1980, p. 141; EBJV 2006, p. 1).
Throughout much of their natural range, remaining stream
populations have retreated into extreme headwater, high elevation, or
upstream reaches (EBJV 2006, p. 2). In the eastern U.S., healthy stream
populations of brook trout (wild brook trout occupying 90-100 percent
of their historical habitat) exist in only 5 percent of subwatersheds
(EBJV 2006, p. 2). Anadromous stocks along the U.S. coast and in many
Canadian rivers have been decimated by dams and estuarine pollution
(Power 1980, p. 195). In the southern portion of its range (southern
Appalachian Mountains), brook trout populations have declined by 75
percent, persisting now only in isolated headwater reaches (EBJV 2006,
p. 6).
Various threats are persistent across the brook trout range. Most
of them involve habitat loss and degradation, such as poor land
management, high water temperature, sedimentation (roads),
urbanization, degraded riparian habitat, stream fragmentation (roads),
dam inundation/fragmentation, and forestry practices (EBJV 2006, pp. 3,
5). Poor land management associated with agriculture (such as clearing
streamside vegetation, over-grazing sensitive areas, ineffectively
managing nutrients, and ditching small streams) ranks as the most
widely distributed impact to brook trout across the eastern U.S. (EBJV
2006, p. 2). Climate change presents a significant threat to brook
trout, with some southern portions predicted to lose between 53-97
percent of their brook trout habitat due to high water temperatures
(Flebbe 2006, p. 1379). While some uncertainty remains about the exact
temperature increase that will result from climate change, the present
range of brook trout is predicted to shrink, particularly in the
southern Appalachians (Hudy et al. 2005, p. 5). Nonnative species are
now present throughout most of the range (Parsons 1973, p. 5).
Interactions with these nonnatives are considered to be among the most
significant biological threats to brook trout rangewide (Peck 2001,
p.13; Hudy et al. 2005, p. 3; EBJV 2006, pp. 2-3, 5). Brown trout have
been shown to displace or reduce stream populations of brook trout
throughout their natural range (Nyman 1970, p. 348; Fausch and White
1981, p. 1226; Waters 1983, p. 144). Encroachment by rainbow trout has
also been documented in the contraction of the range of native brook
trout across their native range (Kelly et al., 1980, pp. 9-10; Power
1980, p. 195; Larson and Moore 1985, p. 200). Species such as small
mouth bass and yellow perch are considered to be significant
competitors with lake-dwelling brook trout (EBJV 2006, pp. 22, 28, 34).
Current Population Status of Brook Trout in the Upper Great Lakes
Brook trout populations throughout the upper Great Lakes region are
relatively common and geographically widespread, although distribution
and abundance is much reduced from historical levels (Power 1980, p.
195; Becker 1983, pp. 321-322; WIDNR and USFWS 2005, p. 17). Dramatic
declines in abundance and distribution of both coaster and stream-
resident ecotypes of brook trout occurred in the upper Great Lakes from
the 1850s to mid-1900s (Goodier 1982, pp. 110, 112; Ritchie and Black
1988, p. 15; Newman and Dubois 1997, pp. 4-6; Enterline 2000, p. 1;
WIDNR and USFWS 2005, pp. 17-18; Schreiner et al. 2008, p. 1305;
Schreiner et al. 2008, p. 1351; Huckins et al. 2008, p. 1322).
There are presently at least 200 streams with documented brook
trout populations in the upper Great Lakes (Moore and Bream 1965, p.
19; Goodier 1982, p. 110; Enterline 2000, p. 30; Newman et al. 2003,
pp. 31-37; Quinlan 2004, unpub. data; Bassett 2009, unpub. data; Ward
2007, p. 16; Schram 2008b, pers. comm.; Scott 2008, pers. comm.; Chase
2009, pers. comm.; OMNR 2009, unpub. data). The current specific status
of most of these populations is not known, but they are described by
the Michigan, Minnesota, and Wisconsin natural resource agencies as
stable and self-sustaining in the upper Great Lakes (Holtz 2008, p. 2;
MIDNR 2008a, p. 49; Schreiner and Ebbers 2008, pers. comm.).
In coldwater tributaries to the upper Great Lakes, brook trout were
historically distributed from the river mouth upstream to the
headwaters or to impassible barriers (Smith and Moyle 1944, p. 119;
Moore and Braem 1965, p. 19; Goodier 1982, p. 111; Becker 1983, p. 321;
WIDNR and USFWS 2005). The brook trout numbers in these stream reaches
once numbered in the hundreds to thousands (Huckins and Baker 2008, p.
1231). A 30-year data set from Wisconsin tributaries shows that, in
streams historically occupied solely by brook trout, brook trout have
contracted into upstream sections and are now nearly absent in lower
reaches (WIDNR 2008, unpub. data). Brook trout abundance has declined
despite the persistence of suitable conditions for brook trout and high
numbers of juvenile nonnative salmonids (WIDNR 2008, unpub. data). In
Wisconsin tributaries to Lake Superior, the distribution of stream-
resident brook trout populations has declined by nearly 50 percent from
historical levels (WIDNR and USFWS 2005, p. 17).
Historically, 119 tributaries to Lake Superior and purportedly 6
Lake Huron streams supported populations of brook trout with coaster
ecotypes (Newman et al. 2003, pp. 31-38; Enterline 2000, p. 30). Once
abundant and widespread throughout the northern portions of the Great
Lakes, populations of brook trout that still exhibit the coaster
ecotypes are presently limited to a few locations (Dehring and Krueger
1985, p. 1; Bailey and Smith 1981, p. 1549; Goodyear et al. 1982, pp.
63-65; Enterline 2000, p. 30; Newman et al. 2003, p. 39; Schreiner et
al. 2008, p. 1351; Mucha and Mackereth 2008, p. 1). Although self-
sustaining populations of stream-resident brook trout are currently
present in 56 of 58 U.S. streams and in all 61 Canadian streams
identified in the Brook Trout Rehabilitation Plan for Lake Superior as
historically supporting populations with coaster ecotypes (Newman et
al. 2003, pp. 31-37; Quinlan 2008, unpub. data; Schreiner 2008, pers.
comm.; Schram 2008c, pers. comm.; Scott 2008, pers. comm.; Chase 2009,
pers. comm.), only 18 populations with coaster ecotypes still persist
there (15 stream-spawning-adfluvial, and 3 lake-spawning-lacustrine)
(Goodyear 1982, pp. 63-65; Quinlan 1999, p. 19; Ritchie and Black
[[Page 23382]]
1988, p. 15; Swainson 2001, p. 41; Newman et al. 2003, pp. 28-39;
Enterline 2000, p. 30; Chase 2009, pers. comm.).
Over the last decade, the presence of coaster brook trout has been
confirmed in other locations within the upper Great Lakes. Surveys, and
in some cases genetic analysis, have confirmed the presence of brook
trout with coaster ecotypes in the following locations; Minnesota
tributaries to Lake Superior (Newman et al. 1999, p. 2; Burnham-Curtis
2000, p. 4; Pranckus and Ostazeski 2003, p. 5; Ward 2007, p. 16), three
Michigan tributaries to Lake Superior (Stimmel 2006, p. 56; MIDNR
2008a, p. 2; Leonard 2009, pers. comm.), along the shoreline of the Red
Cliff Indian Reservation, Wisconsin (Stott and Quinlan 2008, p. 21),
and in Little Todd Harbor and Rock Harbor, Isle Royale (Gorman et al.
2008, p. 1257). The origin of these fish is unknown and natural
reproduction of fish exhibiting the coaster ecotype has not been
confirmed, therefore these locations are not identified as supporting
self-sustaining populations. However, they have potential to be self-
sustaining populations, as outlined by Schreiner et al. (2008).
Abundance of individuals in populations exhibiting the coaster
ecotypes is stable or increasing in several regions of Lake Superior.
In the Salmon Trout River, Michigan, abundance as determined by video
surveillance increased from 118 to 243 in the period from 2004 to 2006
(MIDNR 2008a, p. 6). In the Nipigon River, angler catch per hour has
increased from the late 1980s to the present, while harvest has
decreased substantially (Houle 2004, p. 13). In South Bay, Lake
Nipigon, estimates of spawner abundance continue to increase and
currently number about 600 fish--up from fewer than 100 in the recent
past, but still fewer than the estimated 2,500 present in the mid-1900s
(Swainson 2009, pers. comm.). In Tobin Harbor, Isle Royale National
Park, Michigan, estimates of adult brook trout from 1996, 2001, and
2008 has remained around 200-250 fish (USFWS unpublished data).
Relative abundance based on shoreline electrofishing index surveys in
Tobin Harbor from 1997 to 2008 has fluctuated from 0.3 per hour to 16.7
per hour (USFWS 2008, unpub. data).
There are reintroduction stocking efforts ongoing in several
streams on the Grand Portage Indian Reservation (Newman and Johnson
1996, p. 4), Red Cliff Indian Reservation, Keweenaw Bay Indian
Community Reservation (Donofrio 2002, p. 1), and in Whittlesey Creek,
Wisconsin (USFWS and WIDNR 2003, p. 5). Supplementation stocking
occurred in Siskiwit Bay, Isle Royale, from 1999 to 2005. Data
collected to date indicates limited success with these efforts (Newman
et al. 1999, p. 2; Quinlan 2008, pers. comm.; Stott and Quinlan 2008,
p. 22). Reintroduction efforts in Michigan have recently been
terminated in the Gratiot, Little Carp, Hurricane, and Mosquito Rivers
and Sevenmile Creek (Scott 2007, pers. comm.; Loope 2007, pers. comm.).
Threats to brook trout across its native range are also acting on
brook trout within the upper Great Lakes. A primary impact is the
presence of introduced fishes (e.g., non-native salmonids). Introduced
salmonids have competitive and predatory impacts on brook trout,
although the precise mechanisms may not be fully understood and the
magnitude of impact may vary by species, population size, and
environmental conditions. The decline or loss of the migratory coaster
form has diminished connectivity among populations that once operated
as metapopulations. Populations that occur in such isolated patches can
be lost, increasing the possibility of extirpation. As a species, brook
trout are known to be highly susceptible to exploitation by anglers
(Newman and Dubois 1996, p. 3; Newman et al. 2003, p. 11; Huckins et
al. 2008, p. 1322). Overharvest was a primary cause of the decline of
Great Lakes brook trout populations by the early 1900s, especially the
coaster ecotype, and continues to threaten some populations within the
region (Newman and Dubois 1996, p. 1; Huckins et al. 2008, p. 1322;
Schreiner et al. 2008, p. 1356). Climate change also presents a threat
to upper Great Lakes brook trout, through increased water temperatures,
leading to increased presence of nonnative competitors and predators
along with a decrease in habitat suitability. Although the enormous
coldwater reservoir within the lake environment represents a potential
refuge for Great Lakes brook trout, predicted impacts in both stream
and lake environments still represent a potential threat to their long-
term viability.
Defining a Species Under the Act
Section 3(16) of the Act defines ``species'' to include ``any
species or subspecies of fish and wildlife or plants, and any distinct
vertebrate population segment of fish or wildlife that interbreeds when
mature'' (16 U.S.C. 1532 (16)). Our implementing regulations at 50 CFR
424.02 provide further guidance for 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
previously discussed, coaster brook trout are classified as Salvelinus
fontinalis, the same as other brook trout, and as such we do not
consider the coaster form of the brook trout to constitute a distinct
species or subspecies. Since the coaster brook trout is not a distinct
species or subspecies, we then evaluated whether the coaster brook
trout is a distinct vertebrate population segment to determine whether
it would constitute a listable entity under the Act.
To interpret and implement the distinct vertebrate population
segment (DPS) provisions of the Act and Congressional guidance, the
Service and the National Marine Fisheries Service (now the National
Oceanic and Atmospheric Administration--Fisheries), published the
Policy Regarding the Recognition of Distinct Vertebrate Population
Segments (DPS Policy) in the Federal Register on February 7, 1996 (61
FR 4722). Under the DPS Policy, three elements are considered in the
decision regarding the establishment and classification of a population
of a vertebrate species as a possible DPS. These are applied similarly
for additions to and removals from the List of Endangered and
Threatened Wildlife and Plants. These elements are (1) the discreteness
of a population in relation to the remainder of the species to which it
belongs, (2) the significance of the population segment to the species
to which it belongs, and (3) the population segment's conservation
status in relation to the Act's standards for listing, delisting, or
reclassification.
Distinct Vertebrate Population Segment Analysis
In accordance with our DPS Policy, this section details our
analysis of the first two elements used to assess whether a vertebrate
population segment under consideration for listing may qualify as a
DPS. These elements are (1) the population segment's discreteness from
the remainder of the species to which it belongs and (2) the
significance of the population segment to the species to which it
belongs. Discreteness refers to the ability to circumscribe a
population segment from other members of the taxon based on either (1)
physical, physiological, ecological, or behavioral factors or (2)
international boundaries that result in
[[Page 23383]]
significant differences in control of exploitation, habitat management,
conservation status, or regulatory mechanisms in light of section
4(a)(1)(B) of the Act.
Under our DPS Policy, if we have determined that a vertebrate
population segment is discrete, we consider its biological and
ecological significance to the larger taxon to which it belongs 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. To evaluate
whether a discrete vertebrate population may be significant to the
taxon to which it belongs, we consider the best available scientific
evidence. This evaluation may include, but is not limited to: (1)
Evidence of the persistence of the discrete population segment in an
ecological setting that is unusual or unique for the taxon; (2)
evidence that loss of the population segment would result in a
significant gap in the range of the taxon; (3) evidence that the
population segment represents the only surviving natural occurrence of
a taxon that may be more abundant elsewhere as an introduced population
outside its historical range; and (4) evidence that the discrete
population segment differs markedly in its genetic characteristics from
other populations of the species.
The first step in our DPS analysis was to identify population
segments of the brook trout to evaluate. The petition asked us to (1)
``list as `endangered' the naturally spawning anadromous (lake-run)
Coaster Brook Trout (Salvelinus fontinalis) throughout its known
historic range in the conterminous United States'' (including
designation of critical habitat) and (2) ``determine whether the Salmon
Trout River (STR) coaster is a DPS'' and (3) ``whether the south shore
of Lake Superior population of coasters (which are known to breed today
only in the STR) is `endangered.' '' Although brook trout in the Great
Lakes exhibit three life-history forms (fluvial, adfluvial, and
lacustrine), the petition specifically focused on the coaster, or
adfluvial and lacustrine, forms.
To address the entity identified in the first petition request
(coaster brook trout throughout their historical range in the U.S.), we
identified two approaches to analyzing a potential population segment:
(1) Describe and analyze an upper Great Lakes ``all brook trout''
population segment, which includes all brook trout life forms--fluvial,
adfluvial, and lacustrine ecotypes, inclusive of coaster brook trout--
present throughout the documented historical range of brook trout in
the Great Lakes basin, and (2) describe and analyze an upper Great
Lakes ``coaster-only'' population segment, which includes only the
coaster forms--adfluvial and lacustrine ecotypes--of brook trout
throughout the documented historical range of brook trout in the Great
Lakes basin.
We find that neither of the population segments analyzed constitute
a valid DPS, and therefore the first petitioned entity, coaster brook
trout throughout their historical range in the U.S., is not a valid
DPS. To address the second and third petition requests, we focused on
the brook trout population in the Salmon Trout River and evaluated
whether it qualified as a DPS per our policy. We find that the brook
trout population in the Salmon Trout River also does not constitute a
valid DPS. The remainder of this section details the evaluation of
these population segments as DPSs per our 1996 DPS Policy.
Upper Great Lakes All Brook Trout Population Segment
This population segment encompasses the range of brook trout
populations within the Great Lakes basin that currently or historically
occupied both the tributary and lake environments (including stream-
resident, adfluvial, and lacustrine ecotypes of brook trout). Although
technically not one of the ``Great Lakes,'' we include Lake Nipigon in
Canada in this population because it is part of the Great Lakes
drainage. The best available information indicates the known historical
range of brook trout within the basin included all of Lake Superior and
its drainage (including Lake Nipigon), and the northern portions of
Lakes Michigan and Huron--specifically, that portion of Lake Michigan
north of a line from the Sheboygan River, Wisconsin to Grand Traverse
Bay, Michigan, and that portion of Lake Huron north of Thunder Bay,
Michigan, eastward to include Manitoulin Island to the 81[deg]30'
longitudinal demarcation and west of 81[deg]30' longitude (MacCrimmon
and Campbell 1969, p. 1701; Dehring and Krueger 1985, p. 1; Enterline
2000, pp. 29-30).
Discreteness
Marked Separation
As previously described, the Upper Great Lakes brook trout
population segment we have evaluated encompasses the range of brook
trout populations that currently or historically occupied both the
tributary and lake environments within the Great Lakes basin. Brook
trout within this population segment are physically isolated from other
populations of brook trout as the result of the physical separation
between the drainage of the Great Lakes basin and neighboring
drainages. Consequently, brook trout in the Great Lakes basin meet the
discreteness criterion of being markedly separate from other members of
the brook trout taxon.
International Border
We presently do not find that the brook trout in the Upper Great
lakes on either side of the international United States border with
Canada are discrete due to differences in control of exploitation,
management of habitat, conservation status, or regulatory mechanisms
that are significant in light of section 4(a)(1)(D) of the Act.
Conclusion for Discreteness
In conclusion, we determine that the Upper Great Lakes brook trout
population segment, as defined here, is discrete from the remainder of
the brook trout taxon. This discreteness arises from the population
segment's physical isolation from the remainder of the taxon.
Therefore, we will now consider the potential significance of this
discrete population segment to the remainder of the taxon.
Significance
We have determined that the population of brook trout in the Upper
Great Lakes meets the discreteness elements of the DPS policy, and as
such, we will now evaluate whether this specific population is
significant to the taxon as a whole (i.e., native brook trout in
eastern North America). A discrete population is considered significant
under the DPS policy if it meets one of four of the elements identified
in the policy under significance or can otherwise be reasonably
justified as being significant.
We discuss further below our evaluation of the significance of the
population of brook trout in the Upper Great Lakes relative to the
taxon as a whole.
Evidence of the Persistence of the Discrete Population Segment in an
Ecological Setting That Is Unusual or Unique for the Taxon
On the basis of an evaluation of the best available scientific
information, we have determined that the habitat for brook trout in the
Upper Great Lakes does not represent an ecological setting that is
unusual or unique for the native brook trout relative to the habitat
available to it throughout the entire
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taxon's range in eastern North America. A summary of our evaluation is
below.
Brook trout exhibiting differing life-history forms occupy a
variety of ecosystems from subarctic regions of the Hudson Bay coast,
to temperate areas bordering and east of the Great Lakes, and southern
coldwater habitats in the Appalachian Mountains of Tennessee and
Georgia (Power 1980, p. 142). They have been successfully naturalized
in western North America, South America, Eurasia, Africa, and New
Zealand (MacCrimmon and Campbell 1969, p. 1699, pp. 1703-1717). Within
their large native range in eastern North America, brook trout habitat
includes coastal areas and various-sized lakes, streams, and rivers at
varying altitudes. Most populations inhabit coldwater streams, but
lake-dwelling and lake-spawning (lacustrine form) populations also
occur throughout the range, in spring-fed ponds, small- to medium-sized
lakes, and a few large, oligotrophic (containing relatively little
plant life or nutrients, but rich in dissolved oxygen) lakes.
Anadromous populations (``salters'') of brook trout use marine habitats
in Hudson Bay and along the Atlantic coast.
The upper Great Lakes represent a complex ecological setting for
brook trout. The very large size of the Great Lakes watershed creates
an environment that more closely resembles oceanic physical conditions
(available to the anadromous forms of brook trout) than conditions in
smaller lakes (available to other forms of brook trout). With
approximately 1,500 tributaries and almost 2,800 miles (4,506 km) of
shoreline, Lake Superior also provides brook trout access to a very
large freshwater habitat network. Although the Great Lakes are the
largest freshwater water bodies occupied by brook trout, there are
thousands of lakes in its range including large postglacial lakes
further north in Canada that contain populations of the adfluvial and
lacustrine forms (e.g., Fraser and Bernatchez 2008, p. 1193).
If predicted rising water temperatures in response to climate
change are realized over the entire range of brook trout, the
distributions of brook trout populations would probably shift toward
cooler waters at higher latitudes and altitudes (Meisner 1990b, p.
1068; Magnuson et al. 1997, p. 859; Kling et al. 2003, pp. 53-54). The
greatest effects would likely begin in populations located at the
margins of the taxon's hydrologic and geographic distributions (Meisner
et al. 1990a, p. 282, Kling et al. 2003, p. 54). Although the upper
Great Lakes have already experienced some impacts of climate change
(see Kling et al. 2003, pp. 14-16) and will not be immune to future
impacts (see Kling et al. 2003, pp. 21-25), they may provide
substantial coldwater habitat for brook trout in the future. However,
brook trout have abundant coldwater habitat available in the northern
latitudes of its range, and habitat in northern North America which is
presently too cold may develop into appropriate brook trout habitat
under a warming scenario. We will further evaluate the extent that this
may be the case in the range-wide assessment of native brook trout that
we plan to conduct (see Finding section).
Although the upper Great Lakes represent a diverse and complex
ecological setting which may offer potential coldwater habitat for
brook trout, we must evaluate the breadth of ecological diversity of
brook trout habitat rangewide in our assessment of this population
segment's significance to the rest of the taxon. First, available
information indicates that the large area and wide geographical range
of brook trout habitats, which vary in latitude and altitude and water
form, contain a vast diversity of habitats for brook trout. The
ecological setting of the upper Great Lakes is a small portion of the
brook trout range, and based on available information, its relative
significance to the brook trout species is limited. Second, although we
expect that the Great Lakes may offer substantial coldwater habitat,
there are other large, deep, oligotrophic lakes, and numerous lakes and
streams at higher latitudes that may buffer the species from potential
climate change impacts. Given the available information on the
diversity and extent of ecological settings of brook trout in the rest
of its range, we conclude at this time that the upper Great Lakes is a
not unique or unusual setting of significance for the native brook
trout in eastern North America.
Evidence That Loss of the Population Segment Would Result in a
Significant Gap in the Range of the Taxon
Loss of brook trout, including any or all life forms, in the upper
Great Lakes, when considered in relation to brook trout throughout the
remainder of the spe