Document ID: EPA-HQ-OW-2003-0068-0053
Agency: epa
Document Type: Supporting & Related Material
Title: 
Posted Date: 2003-09-22T04:00Z

Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
Quality
Standards
Temperature
Criteria
Draft
Discussion
Paper
and
Literature
Summary
Revised
December
2002
Publication
Number
00­
10­
070
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
Quality
Standards
Temperature
Criteria
Draft
Discussion
Paper
and
Literature
Summary
Prepared
by:
Water
Quality
Program
Washington
State
Department
of
Ecology
Watershed
Management
Section
Olympia,
Washington
98504
Revised
December
2002
Publication
Number
00­
10­
070
For
additional
copies
of
this
document
contact:

Department
of
Ecology
Publications
Distribution
Center
P.
O.
Box
47600
Olympia,
WA
98504­
7600
Telephone:
(
360)
407­
7472
If
you
have
special
accommodation
needs
or
require
this
document
in
an
alternative
format,
please
call
Mark
Hicks
at
(
360)
407­
6477.
The
TTY
number
is
711
or
1­
800­
833­
6388.
Email
can
be
sent
to
mhic461@
ecy.
wa.
gov.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
i
Washington's
Surface
Water
quality
Standards
Table
of
Contents
Part
I:
Background
and
Project
History
.....................................................................................................
1
1.
Background
............................................................................................................................
1
2.
Current
Temperature
Requirements
.......................................................................................
1
3.
Organization
of
this
Review
Document
.................................................................................
2
Part
II:
The
Effect
of
Temperature
on
the
...................................................................................................
3
Freshwater
Aquatic
Life
of
Washington
.........................................................................................
3
1.
The
Goal
of
this
Technical
Review........................................................................................
3
2.
Summary
of
Thermal
Requirements
and
Recommended
Threshold
Values..........................
3
a)
Native
Char
(
Bull
Trout
and
Dolly
Varden)
......................................................................
4
b)
Salmon
and
Trout
..............................................................................................................
4
c)
Warm
Water
Aquatic
Life..................................................................................................
5
d)
Special
Provisions
to
Prevent
Acute
Effects......................................................................
6
3.
Methodology
and
Considerations
in
Establishing
Temperature
Recommendations..............
6
a)
The
Multiple
Lines
of
Evidence
(
MLE)
Methodology......................................................
6
b)
General
Thoughts
and
Observations................................................................................
10
i)
Adjusting
Laboratory
Data
for
Application
to
Natural
Waters
........................................
10
ii)
Minimum
Temperature
Thresholds
.................................................................................
15
iii)
Protection
of
Untested
Species.......................................................................................
16
4.
Temperature
Requirements
of
Char,
Salmon,
and
Trout
Species
........................................
17
a)
Native
Char
Temperature
Requirements..........................................................................
17
i)
General
Life
History
Information:....................................................................................
17
ii)
Spawning
Requirements..................................................................................................
19
iii)
Juvenile
Rearing.............................................................................................................
23
iv)
Migratory
Adult
and
Sub­
Adult
Char
Populations.........................................................
28
v)
Lethality
to
Adults
and
Juveniles
....................................................................................
30
b)
Salmon
and
Trout
............................................................................................................
33
i)
General
Life
History
Information.....................................................................................
33
ii)
Spawning
Requirements..................................................................................................
38
iii)
Juvenile
Rearing.............................................................................................................
52
iv)
Juvenile
Winter
Holding.................................................................................................
80
v)
Adult
Migration
...............................................................................................................
81
vi)
Lethality
to
Adults
and
Juveniles
...................................................................................
91
c)
Temperature
Influenced
Fish
Diseases
..........................................................................
108
d)
Smoltification
and
Sea
Water
Adaptation
.....................................................................
115
e)
Miscellaneous
Indigenous
Species
................................................................................
120
i)
Sensitive
Amphibians
.....................................................................................................
121
ii)
Other
Sensitive
Fish
Species
.........................................................................................
123
iii)
Stream
Macroinvertebrates...........................................................................................
125
f)
Miscellaneous
Indigenous
Fish
Species
.........................................................................
128
g)
Summary
of
Temperature
Requirements
for
Indigenous
Aquatic
Life
.........................
131
i)
Cold
Water
Species
........................................................................................................
131
ii)
Warm
water
Species
......................................................................................................
133
5.
References
Reviewed
.........................................................................................................
138
Part
III:
Ambient
Temperatures
of
Washington's
Streams
and
Rivers.....................................................
175
Proposal
for
Applying
Char
Protection
.......................................................................................
183
Page
ii
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Abstract
Maintaining
proper
temperatures
in
our
natural
waterways
is
vital
to
the
long­
term
health
of
fish
and
other
aquatic
life.
In
response
to
critiques
over
the
adequacy
of
Washington's
existing
temperature
criteria
for
freshwaters,
the
Washington
State
Department
of
Ecology
performed
a
comprehensive
review
of
the
available
technical
literature
on
the
temperature
requirements
of
our
native
fish
and
aquatic
life.
This
document
summarizes
the
findings
of
that
review.

Part
I
of
this
document
provides
a
brief
history
of
the
review
effort.
Part
II
of
this
document
reviews
the
scientific
literature
and
establishes
technical
recommendations
for
protecting
the
state's
freshwater
aquatic
species.
Based
on
previous
reviews
of
the
literature
it
was
determined
that
freshwater
species
can
be
reasonably
placed
into
four
groups
having
similar
sensitivity
to
temperature.
These
four
groups,
identified
by
key
species,
serve
as
the
foundation
for
this
review:

1)
Native
Char
(
bull
trout
and
Dolly
Varden);
2)
Salmon,
Steelhead,
Cutthroat
Trout,
and
Coastal
Rainbow
Trout;
3)
Interior
non­
Anadromous
Redband
Trout
(
eastern
Washington
species);
and
4)
Warm
Water
Fish
Species
(
e.
g.,
dace,
shiner,
sucker,
etc.).

Technical
evaluations
provide
recommendations
for
temperature
thresholds
that
protect
the
key
life­
stages
of
adult
migration
and
holding,
spawning,
incubation,
juvenile
rearing,
and
smoltification.
Recommendations
are
also
made
to
avoid
significant
increases
in
the
risks
of
warm
water
fish
diseases
and
parasites.

Recommendations
are
also
presented
that
may
be
instructive
in
trying
to
avoid
detrimental
effects
from
discrete
site­
specific
human
actions
that
may
cause:

1)
Thermal
blockages
to
migration,
2)
Short­
term
(
7­
day
duration)
lethality,
and
3)
Near
instantaneous
(
1­
2
seconds)
lethality.

The
temperature
thresholds
identified
in
this
paper
are
ones
which
Ecology
has
high
confidence
represents
the
upper
threshold
temperatures
that
provide
full
protection
for
the
species.
As
temperatures
rise
above
these
values,
negative
impacts
to
the
health
of
fish
and
other
aquatic
life
will
rapidly
escalate
to
levels
detrimental
to
the
health
of
aquatic
communities.
The
metric
used
to
express
the
technical
recommendations
(
a
7­
day
average
of
the
daily
maximum
temperatures)
was
chosen
to
better
match
the
laboratory
and
field
research
results
to
an
exposure
period
that
reflects
the
risk
of
harm
to
aquatic
species.
The
result
of
this
process
is
the
recommendation
of
temperature
limits
that
better
reflect
the
thermal
requirements
and
limitations
of
the
state's
indigenous
freshwater
fish
and
aquatic
life.
There
are
still
some
notable
areas
where
more
research
would
improve
confidence
in
specific
recommendations
(
particularly
for
warm
water
species
and
for
interior
redband
trout).
It
is
clear
however,
that
implementation
of
the
temperature
thresholds
identified
in
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
iii
Washington's
Surface
Water
quality
Standards
part
II
of
this
paper,
wherever
attainable
would
eliminate
human
warming
of
water
temperature
as
a
source
of
impairment
to
our
native
aquatic
communities.

Part
III
of
this
document
examines
temperature
data
from
rivers
and
streams
around
Washington.
The
information
provided
in
this
section
can
be
used
to
assess
the
potential
implications
of
selecting
specific
temperature
values
for
use
as
state
criteria.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
1
Washington's
Surface
Water
quality
Standards
Part
I
Background
and
Project
History
1.
Background
The
Washington
State
Department
of
Ecology
administers
the
state's
surface
water
quality
standards
(
Chapter
173­
201A
WAC).
These
regulations
establish
minimum
requirements
for
the
quality
of
water
that
must
be
maintained
in
lakes,
rivers,
streams,
and
marine
waters.
This
is
done
to
ensure
that
all
the
beneficial
uses
associated
with
these
waterbodies
are
protected.
Examples
of
protected
beneficial
uses
include:
aquatic
life
and
wildlife
habitat,
fishing,
shellfish
collection,
swimming,
boating,
aesthetic
enjoyment,
and
domestic
and
industrial
water
supplies.

As
part
of
a
public
review
of
its
water
quality
standards,
the
Department
of
Ecology
convened
a
technical
work­
group
to
evaluate
the
water
quality
criteria
established
to
protect
freshwater
aquatic
communities.
One
of
the
recommendations
of
the
work­
group
was
that
Ecology
should
re­
evaluate
the
state's
existing
temperature
criteria.

2.
Current
Temperature
Requirements
The
existing
state
surface
water
quality
standards
contain
three
separate
single
daily
maximum
temperature
criteria
limits
that
can
be
applied
to
rivers:

Class
AA
­
16
°
C
Class
A
­
18
°
C
Class
B
­
21
°
C
Lake
Class
­
Temperatures
are
to
be
maintained
at
natural
levels.

Class
AA
and
Class
A
provide
two
different
levels
of
protection
for
the
same
set
of
beneficial
uses,
and
are
intended
to
protect
salmonid
spawning,
rearing,
and
migration.
Class
AA
is
predominately
applied
to
forested
upland
areas,
but
Class
A
waters
are
designated
broadly
throughout
the
state.
Class
B,
is
designed
only
to
protect
salmonid
rearing
and
migration,
and
was
not
intended
to
fully
protect
spawning.
There
are
only
a
small
number
waterbodies
in
the
state
that
have
been
assigned
the
Class
B
designation.
With
each
class,
the
criteria
are
applied
as
the
highest
single
daily
maximum
measurement
of
temperature
occurring
in
the
waterbody.
Page
2
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
3.
Organization
of
this
Review
Document
Part
I:
Provides
a
brief
background
discussion
on
the
effort
to
revise
the
state's
existing
water
quality
criteria
for
temperature
in
freshwater
systems.

Part
II:
Reviews
available
scientific
research
on
the
effects
of
temperature
on
aquatic
life,
with
a
particular
focus
on
species
occurring
in
Washington.

Part
III:
Summarizes
the
patterns
of
temperature
in
Washington's
rivers
using
available
continuous
monitoring
data.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
3
Washington's
Surface
Water
quality
Standards
Part
II
The
Effect
of
Temperature
on
the
Freshwater
Aquatic
Life
of
Washington
Prepared
by:
Mark
Hicks,
Senior
Analyst
Water
Quality
Standards
Washington
Department
of
Ecology
(
360)
407­
6477
mhic461@
ecy.
wa.
gov
1.
The
Goal
of
this
Technical
Review
The
primary
goal
of
this
paper
is
to
identify
temperature
thresholds
that
if
met
would
have
a
high
probability
of
fully
protecting
Washington's
freshwater
aquatic
communities.
Full
protection
includes
the
reasonable
prevention
of
both
lethal
and
sublethal
effects
that
may
detrimentally
affect
the
health,
fecundity,
and
the
ultimate
wellbeing
of
naturally
balanced
indigenous
populations.
This
review
uses
the
available
technical
literature
to
determine
temperature
regimes
that
will
fully
protect
the
individual
life­
history
stages
of
key
species.
This
information
is
then
combined
into
stream­
wide
temperature
recommendations
that
will
protect
entire
communities
of
aquatic
life.

The
following
summarizes
the
findings
pertaining
to
the
protection
of
our
state's
native
fish
species.
The
technical
recommendations
are
based
on
grouping
together
species
having
the
same
or
nearly
the
same
temperature
requirements.
This
both
simplifies
the
resulting
recommendations
and
acknowledges
that
these
species
commonly
occur
together
in
the
state's
streams,
rivers,
and
lakes.
This
approach
results
in
establishing
temperature
threshold
values
for
four
different
species
groupings:
1)
native
char,
2)
salmon
and
coastal
trout,
3)
non­
anadromous
interior
redband
trout,
and
4)
warm
water
fish
species.

2.
Summary
of
Thermal
Requirements
and
Recommended
Threshold
Values
Page
4
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
a)
Native
Char
(
Bull
Trout
and
Dolly
Varden)

Char
require
colder
waters
to
be
fully
supported
than
the
other
fishes
native
to
Washington.
Char
do
not
begin
spawning
until
temperatures
drop
to
7­
9
°
C,
and
subsequent
egg
incubation
is
harmed
by
continuous
daily
average
water
temperatures
above
5­
6
°
C.
The
same
waters
that
are
used
for
spawning
are
also
used
for
tributary
rearing
by
juvenile
char.
Tributary
rearing
waters
that
provide
full
support
for
char
are
best
described
by
summer
maximum
temperatures
at
or
below
13­
15
°
C
or
average
(
7­
day)
daily
maximum
temperatures
below
about
12.5­
14
°
C.

At
age
2­
3,
char
often
move
to
lower
more
main
stem
waters
to
rear.
This
may
be
a
pattern
that
more
effectively
uses
the
larger
prey
bases
of
larger
stream
and
rivers,
or
just
acts
to
disperse
the
populations
in
order
to
use
wider
resource
areas.
In
any
case,
it
is
common
to
find
char
in
waters
where
the
summer
maximum
temperatures
would
appear
unsuitable
based
on
research
focused
on
young
juveniles
and
non­
migratory
(
resident)
populations.
It
appears
that
char
will
move
into
and
out
of
these
waters
in
a
manner
that
avoids
(
or
in
response
to)
warm
water
temperatures.
The
temperatures
noted
in
association
with
the
movement
varies
but
generally
appears
to
be
above
those
identified
as
necessary
to
fully
protect
the
rearing
of
young
juveniles.
At
this
time,
information
is
not
available
to
either
identify
the
historic
patterns
in
specific
Washington
streams
or
identify
with
confidence
temperature
criteria
appropriate
for
this
migratory
life­
stage.

b)
Salmon
and
Trout
Salmon
and
trout
both
have
similar
spawning
and
incubation
requirements.
In
evaluating
the
temperature
requirements
of
salmon
and
trout
it
is
important,
however,
to
remember
that
salmon
begin
spawning
in
the
fall
as
temperatures
are
cooling
and
trout
begin
spawning
in
the
spring
as
temperatures
are
warming.
For
simplicity,
the
following
discussion
follows
the
pattern
of
fall
spawning
fish:

In
the
weeks
immediately
preceding
spawning,
temperatures
above
14­
16
°
C
can
reduce
the
health
of
the
eggs
and
sperm
in
adult
fish.
Constant
temperatures
in
the
range
of
8­
10
°
C
and
daily
maximum
temperature
below
13­
15
°
C
are
necessary
to
ensure
that
fertilized
eggs
have
high
survival
success
and
the
embryos
develop
into
healthy
emergent
fry.
Once
the
fry
emerge,
juvenile
rearing
will
be
fully
supported
in
natural
streams
where
the
average
temperatures
do
not
exceed
13­
15
°
C,
and
maximum
temperatures
do
not
exceed
approximately
17­
19
°
C.
Once
the
fish
have
reached
a
critical
size,
they
will
begin
the
process
of
changing
from
parr
to
smolts.
This
is
the
critical
physiologic
process
that
allows
these
freshwater
juveniles
to
adapt
their
systems
for
life
in
marine
waters.
Average
temperatures
greater
than
15­
16
°
C
appear
capable
of
stopping
the
process
altogether,
and
average
temperatures
below
12­
13
°
C
appear
necessary
to
fully
support
smoltification
(
particularly
for
the
spring
out­
migration
of
steelhead).
In
early
spring,
overlapping
the
period
of
out­
migration
of
many
anadromous
salmonids,
adult
smelt
will
be
moving
upstream
to
spawn.
These
fish
have
been
found
to
have
very
low
temperature
thresholds.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
5
Washington's
Surface
Water
quality
Standards
While
the
information
on
these
fish
is
somewhat
limited,
it
appears
that
spawning
success
can
be
impaired
by
maximum
temperatures
above
about
12­
13
°
C,
and
adult
mortality
has
been
noted
in
association
with
water
temperatures
as
low
as
16
°
C.
After
a
period
of
growth
in
the
ocean,
salmon
and
steelhead
will
begin
their
migrations
back
up
to
their
natal
streams
to
spawn.
Migration
can
be
impaired
by
experiencing
average
temperatures
above
about
15
°
C
and
maximum
temperatures
above
about
18­
20
°
C.
Temperatures
above
21­
22
°
C
have
frequently
blocked
the
migration
of
anadromous
fish
altogether;
particularly
where
the
fish
have
come
from
cooler
waters.
For
both
the
rearing
of
juvenile
fish
and
the
migration
of
adult
fish,
warm
water
diseases
can
reduce
survival.
Average
temperatures
above
13­
14
°
C
and
periods
of
maximum
temperatures
above
about
17­
18
°
C
have
been
found
to
increase
the
risks
of
losses
associated
with
warm
water
fish
diseases
and
parasites.
Extreme
temperatures
can
be
lethal
to
salmon
and
trout
with
exposure
periods
from
a
couple
of
seconds
to
a
couple
of
hours.
Single
daily
peak
temperature
of
24­
25
°
C
are
capable
of
killing
salmonids
that
have
not
been
well
acclimated
to
warm
waters
in
advance
of
the
exposure,
and
plumes
of
water
warmer
than
32­
33
°
C
can
be
lethal
to
passing
fish
in
only
1­
2
seconds
regardless
of
their
prior
acclimation
temperature.

The
information
reviewed
for
revising
the
state's
temperature
criteria
generally
suggests
that
non­
anadromous
interior
forms
of
rainbow
trout
(
often
described
as
the
subspecies
of
redband
trout
that
occurs
on
the
eastside
of
the
Cascade
mountains
in
Washington)
have
higher
optimal
temperature
ranges
than
other
subspecies
of
rainbow
trout.
However,
the
information
was
rather
inconsistent
on
this
point,
and
caution
is
warranted
before
allowing
any
significant
departure
from
the
temperature
ranges
determined
healthy
for
the
majority
of
the
state's
salmon
and
trout.
It
is
recommended
therefore
that
temperatures
at
the
upper
end
of
the
range
determined
to
fully
protect
salmon
and
trout
in
general
be
viewed
as
the
warmest
temperatures
that
will
protect
these
interior
redband
trout.
The
available
information
suggests
that
acutely
lethal
temperatures
are
very
similar
for
all
subspecies
and
stocks
of
trout,
and
thus
no
sound
basis
exists
for
assuming
that
these
fish
would
be
better
capable
of
resisting
short­
term
spikes
of
temperature
into
the
lethal
range.

c)
Warm
Water
Aquatic
Life
A
warm
water
fish
community
in
Washington
would
be
characterized
by
the
presence
of
redside
shiner;
tui
chub;
margined,
mottled,
or
piute
sculpin;
longnose
or
speckled
dace,
sucker,
and
northern
pikeminnow.
These
fish
are
known
to
exist
in
our
warmest
waters,
where
they
often
out­
compete
introduced
populations
of
rainbow
trout.
Insufficient
information
exists
to
develop
individual
water
quality
recommendations
for
these
species,
so
it
is
recommended
that
they
be
considered
broadly
as
a
community.
There
is
very
little
experiential
data
available
that
can
be
used
to
establish
the
fully
protective
upper
temperature
range
for
our
warm
water
species
in
general,
so
these
recommendations
may
need
to
be
revisited
periodically
to
determine
if
changes
are
warranted.
Establishing
criteria
to
protect
our
temperature
tolerant
non­
salmonid
fish
species
will
also
provide
protection
for
desirable
introduced
warm
water
sport
fish
species
such
as
bass
and
crappie.
In
general,
Washington's
indigenous
warm
water
fish
communities
can
sometimes
thrive
in
waters
that
have
summer
Page
6
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
maximum
temperatures
as
high
as
25­
27
°
C;
although,
most
appear
to
prefer
waters
below
20
°
C
d)
Special
Provisions
to
Prevent
Acute
Effects
The
previous
technical
recommendations
are
designed
to
protect
waterbodies
in
recognition
of
the
way
that
temperatures
change
in
nature
over
hours,
days,
and
weeks.
Discrete
human
actions,
such
as
point
source
discharges,
are
capable
of
altering
these
normal
temperature
regimes
and
thus
may
require
special
consideration.
For
example,
scheduled
or
emergency
bypasses
for
maintenance
at
an
industrial
facility
may
require
asking
questions
like:
a)
How
much
can
temperature
be
raised
for
a
single
day
or
week?,
or
b)
How
hot
can
a
discharge
be
within
an
authorized
mixing
zone?
These
questions
require
a
departure
from
consideration
of
what
are
healthy
stream
conditions
and
instead
must
focus
on
what
is
tolerable
for
a
discrete
location
and
a
limited
duration
of
time.
To
assist
in
answering
these
inevitable
questions,
recommendations
have
been
made
for
avoiding
acute
(
short­
term)
lethality
and
barriers
to
migration.

For
evaluating
the
effects
of
discrete
human
actions,
a
7­
day
average
of
the
daily
maximum
temperatures
greater
than
22
°
C
or
a
1­
day
maximum
greater
than
23
°
C
should
be
considered
lethal
to
cold
water
fish
species
such
as
salmonids.
Discharge
plume
temperatures
should
be
maintained
such
that
fish
could
not
be
entrained
(
based
on
plume
time
of
travel)
for
more
than
2
seconds
at
temperatures
above
33
°
C
to
avoid
creating
areas
that
will
cause
near
instantaneous
lethality.
Barriers
to
migration
should
be
assumed
to
exist
anytime
daily
maximum
water
temperatures
are
greater
than
22
°
C
and
the
adjacent
down­
stream
water
temperatures
are
3
°
C
or
more
cooler.

3.
Methodology
and
Considerations
in
Establishing
Temperature
Recommendations
This
chapter
discusses
the
methodology
used
and
some
of
the
underlying
thoughts
and
concerns
that
went
into
establishing
the
temperature
threshold
recommendations
contained
in
this
paper.

a)
The
Multiple
Lines
of
Evidence
(
MLE)
Methodology
Scientific
information
comes
in
a
wide
variety
of
forms.
These
include:

 
Laboratory
testing
where
the
temperature
is
held
constant,
 
Laboratory
testing
where
the
temperature
is
made
to
fluctuate
at
a
set
rate,
 
Controlled
field
studies
using
either
natural
or
artificial
channels,
 
Field
studies
where
environmental
variables
such
as
shade
are
altered,
and
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
7
Washington's
Surface
Water
quality
Standards
 
Field
observational
studies
where
the
patterns
of
fish
are
observed
in
the
wild.

All
of
these
sources
of
information
provide
valuable
insights,
but
it
can
be
a
challenging
task
to
try
and
compare
and
contrast
such
different
types
of
research.
This
has
led
many
researchers
to
simplify
their
approach
and
select
only
a
single
type
of
research.
This
simplification,
while
understandable,
can
result
in
a
loss
of
understanding.
More
importantly,
however,
it
can
result
in
a
lost
opportunity
to
demonstrate
how
well
all
of
these
very
different
types
of
studies
correspond
to
one
another.
The
key
to
using
a
diversity
of
information
types
is
to
convert
the
results
into
a
common
metric.

The
multiple
lines
of
evidence
(
MLE)
approach
used
in
this
paper
was
developed
as
a
means
to
use
all
of
the
available
scientific
information
to
support
sound
decision
making.
The
MLE
methodology
was
developed
to
provide
a
method
for
making
recommendations
that
are
transparent
to
the
reviewer
and
that
can
be
predictably
modified
when
new
information
becomes
available.
The
basic
approach
is
rather
simple.
All
the
scientific
information
is
sorted
first
by
the
life­
stage
(
e.
g.,
spawning,
rearing,
migration,
etc.)
or
by
some
discrete
environmental
risk
(
e.
g.,
lethality,
smoltification,
disease,
etc.).
The
information
is
then
sorted
into
different
categories
of
study
types.
The
following
provides
a
simplified
example
of
how
this
information
can
be
categorized
into
independent
lines
of
evidence
(
ILOE)
for
the
life­
stage
of
juvenile
rearing:

Study
types
(
ILOE):
Constant
temperature
laboratory
testing
of
growth
Fluctuating
temperature
laboratory
testing
of
growth
Controlled
field
studies
on
growth
Studies
on
the
distribution
and
health
status
of
natural
populations
Laboratory
studies
examining
competition
and
predation
Field
studies
examining
competition
and
predation
For
each
line
of
evidence
the
study
conclusion
are
standardized
into
a
standardized
exposure
metric
and
summarized
as
the
range
of
individual
study
results.
Depending
upon
the
line
of
evidence,
this
range
may
be
either
the
absolute
range
or
the
dominant
range
(
e.
g.,
90th
percentile
of
distribution
of
study
results).

The
standard
metric
for
this
temperature
analysis
is
a
7­
day
average
of
the
daily
maximum
temperatures
(
7DADMax).
This
metric
was
chosen
primarily
because:

1.
Sublethal
chronic
biologic
reactions
generally
take
more
than
a
week's
exposure
to
become
meaningful;
2.
Small
daily
maximum
temperature
fluctuations
beyond
some
"
healthy"
target
level
will
not
be
biologically
meaningful,
but
if
a
single
daily
maximum
metric
were
chosen
and
then
not
attained
such
fluctuations
would
have
regulatory
repercussions;
and
3.
It
is
not
as
defensible
to
use
weekly
averages
of
the
daily
average
temperatures
alone
because
fluctuations
about
the
mean
temperature
can
be
highly
variable
and
extreme
fluctuations
will
erase
or
diminish
the
benefits
of
otherwise
healthy
average
temperatures.
Page
8
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
To
make
the
conversions
to
a
standard
metric,
this
analysis
relies
upon
the
conversion
equations
provided
by
Dunham
et
al.
(
2000)
that
are
based
on
data
from
752
stream
sites
located
in
the
Western
United
States;
particularly
the
Northwest.
For
converting
the
temperature
research
results
for
protecting
bull
trout
and
Dolly
Varden
(
Washington's
most
cold
water
loving
fish
species),
it
is
assumed
that
their
habitat
will
have
very
stable
temperatures.
This
analysis
uses
the
assumption
that
summer
average
diel
fluctuations
are
less
than
2
°
C
in
char
habitat.
This
is
consistent
with
the
state's
information
showing
that
colder
streams
(
7DADMax
<
15C)
have
median
average
fluctuations
of
2.1
°
C
(
90%
between
1.1­
3.6
°
C).
It
is
also
consistent
with
the
commonly
held
belief
that
many
of
these
waters
are
kept
thermally
stable
due
to
a
higher
reliance
on
input
from
groundwater.
For
salmon
and
trout
waters,
the
conversion
is
based
on
assuming
that
the
summer
average
diel
fluctuations
are
from
4­
6
°
C.
This
is
consistent
with
the
state's
data
showing
that
warmer
streams
(
7DADMax
15­
19
°
C)
have
median
average
fluctuations
of
1.2­
5.3
°
C
(
90
percentile
range
 
median
2.6
°
C).
It
also
recognizes
that
the
waterbodies
used
by
salmon
and
trout
have
the
most
variable
temperature
regimes
overall.
The
following
table
shows
the
adjustment
calculations
that
were
used
to
convert
temperatures
to
a
common
metric.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
9
Washington's
Surface
Water
quality
Standards
Table
3.1.
Conversion
equations
for
standardizing
duration
of
exposure
scenarios.
These
are
used
to
convert
study
results
to
the
standard
metric
of
a
7­
day
average
of
the
daily
maximum
temperatures.

The
metric
adjustment
step
was
also
used
where
appropriate
to
put
bounds
on
the
potential
correct
estimate
on
each
line
of
evidence.
This
bounding
process
was
used
where
there
was
reasonable
uncertainty
about
the
duration
of
exposure
that
best
represented
the
line
of
evidence
and
the
related
biological
response.
For
example,
growth
studies
are
generally
conducted
for
a
relatively
long
period
of
time
(
e.
g.,
30­
90
days),
but
significant
changes
in
growth
between
different
test
temperatures
are
commonly
obvious
after
the
first
week
or
two.
Thus
there
is
uncertainly
whether
the
results
of
these
tests
should
be
applied
as
if
they
are
summer
average
exposures
or
weekly
average
exposures.
In
the
face
of
this
uncertainty,
conversions
are
made
for
both
possible
cases.
The
range
produced
by
using
these
two
adjustment
factors
creates
a
range
within
which
the
most
probable
correct
answer
would
be
expected
to
occur.

The
results
of
this
MLE
process
are
presented
in
tabular
form,
and
a
range
is
produced
by
independently
averaging
both
the
lower
and
upper
range
values
for
each
line
of
evidence.
This
creates
a
range
within
which
the
best
estimate
should
be
found.
The
midpoint
of
this
range
is
considered
to
be
the
overall
best
estimate,
unless
overriding
concerns
with
any
particular
line
of
evidence
suggest
another
conclusion
is
warranted.
In
such
a
case,
the
suspect
line
of
evidence
is
noted
and
either
dropped
entirely
from
the
final
range
calculation,
or
is
used
as
a
basis
for
conditioning
the
recommendation.
Table
3.2
shows
a
simplified
example
based
on
protecting
the
juvenile
rearing
life­
stage
of
salmon
and
trout.
Convert
from:
To
a
7DADMax
(
°
C)
Summer
Average
of
the
Diel
Ranges
(
°
C)
In
Char
Spawning
Habitat:
Summer
max
Subtract
0.55
0­
2
Summer
mean
Add
2.00
0­
2
Weekly
mean
(
highest)
Add
0.93
0­
2
Daily
mean
(
highest)
Add
0.62
0­
2
In
Salmon
and
Trout
Habitat:
Summer
max
Subtract
0.95
4­
6
Summer
mean
Add
4.64
4­
6
Weekly
mean
(
highest)
Add
3.18
4­
6
Daily
mean
(
highest)
Add
2.60
4­
6
Page
10
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Table
3.2.
Juvenile
rearing
of
salmon
and
trout:

Line
of
Evidence
(
LOE)
7DADMax
(
°
C)
Midpoint
Comments
Regarding
LOE
Laboratory
Growth
Studies
at
Constant
Temperatures
x­
z
y
Based
on
well
controlled
laboratory
tests.
Laboratory
Growth
Studies
at
Fluctuating
Temperatures
x­
z
y
Field
Studies
on
Growth
x­
z
y
Predation
and
Competition
x­
z
y
Ranges
Identified
as
Protective
x­
z
y
Basis
for
estimates
and
intended
metrics
unclear.
Comparing
Discrete
Test
Regimes
x­
z
y
Laboratory
Temperature
Preferences
x­
z
y
Swimming
Performance
and
Scope
for
Activity
x­
z
y
Field
Distribution
of
Healthy
Populations
x­
z
y
This
estimate
relies
on
the
general
upper
range
considered
healthy,
and
temperatures
above
which
coldwater
species
begin
to
loose
dominance
Summary
Statistics
and
Final
Estimated
Range:
Ave(
x)­
Ave(
z)
Midpoint
Based
on
previous
draft
reviews
of
the
temperature
requirements
of
Washington's
native
fish,
it
was
determined
the
differences
in
thermal
thresholds
between
species
were
generally
slight.
Only
division
into
three
species
groupings
(
guilds)
appears
warranted
(
Hicks,
1998,
2000).
Therefore,
the
multiple
lines
of
evidence
procedure
was
conducted
separately
in
this
current
review
only
for
the
guilds
of:
1)
Char,
2)
Salmon
and
trout,
and
3)
Warm
water
species.

b)
General
Thoughts
and
Observations
i)
Adjusting
Laboratory
Data
for
Application
to
Natural
Waters
Laboratory
tests
do
not
represent
the
full
range
of
conditions
that
an
organism
will
face
in
the
natural
environment.
In
most
laboratory
tests
fish
are
exposed
to
a
constant
temperature
environment,
while
in
natural
waters
the
temperature
continuously
fluctuates
during
each
day,
between
days,
and
in
seasonal
trends
of
spring
warming
and
fall
cooling.
In
natural
waters,
fish
must
actively
maintain
position
and
seek
food
and
shelter
in
the
currents
of
rivers,
succeed
in
the
face
of
inter­
and
intra­
species
competition
for
both
food
and
shelter,
avoid
predation,
and
resist
disease.
In
laboratory
studies,
however,
the
fish
are
often
in
test
chambers
without
substantial
currents,
fed
food
in
pellet
form,
treated
to
prevent
disease,
and
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
11
Washington's
Surface
Water
quality
Standards
seldom
need
to
compete
or
avoid
predation.
On
the
other
hand,
in
laboratory
tests,
fish
are
often
crowded
into
very
small
unnatural
spaces,
even
Styrofoam
cups,
forced
to
perform
using
electrical
stimulation
or
prodding,
subjected
to
laminar
artificial
flows,
and
often
fed
unusual
rations
with
large
time
intervals
of
starvation.

Because
of
the
differences
between
laboratory
conditions
and
the
environmental
conditions
that
fish
face
in
the
natural
world,
we
must
use
caution
in
how
we
apply
laboratory­
derived
data
in
setting
ambient
water
criteria.
We
must
ensure
that
the
temperature
regimes
used
in
the
laboratory
tests
are
considered
in
any
application
to
natural
streams.

Growth
Studies:

Most
of
the
research
on
optimal
growth
temperatures
is
conducted
with
water
kept
at
a
constant
temperature.
Water
quality
standards,
however,
must
apply
to
naturally
fluctuating
thermal
environments.
Since
temperature
directly
effects
the
metabolism
of
the
fish,
a
fish
kept
continuously
in
warm
water
will
experience
more
metabolic
enhancement
than
one
which
only
experiences
that
same
temperature
for
one
or
two
hours
per
day.
Thus,
constant
test
results
cannot
be
directly
applied
as
a
daily
maximum
temperature
in
a
fluctuating
stream
environment.
The
literature
examined
for
this
paper
strongly
suggests
that
constant
temperature
test
results
can
be
used
to
represent
daily
mean
temperatures
(
Hokanson
et
al.,
1977;
Clarke,
1978;
Grabowski,
1973;
Thomas
et
al.,
1986;
Hahn,
1977;
and
Dickerson,
Vinyard,
and
Weber,
1999,
and
unpublished
data,
as
cited
in
Dunham,
1999);
at
least
in
systems
with
moderate
temperature
fluctuations.
Growth
studies
may
be
conducted
for
substantially
varying
periods
of
time
(
14
to
90
days)
but
generally
encompass
time­
frames
that
would
match
reasonably
well
with
periods
of
maximum
summer
temperatures
(
20­
60
days).

For
illustrative
purposes
we
can
examine
the
findings
of
Hokanson
et
al.
(
1977).
This
study
compared
specific
growth
and
mortality
rates
of
juvenile
rainbow
trout
for
50
days
at
seven
constant
temperatures
between
8
°
C
and
22
°
C
and
six
diel
temperature
fluctuations
(
sine
curve
of
amplitude
+/­
3.8
°
C
about
mean
temperatures
from
12
°
C
to
22
°
C
Figure
3.1.
Relationship
between
growth
rates
and
mean
temperature
exposures.
Based
on
Hokanson
et
al.(
1977).

R
2
=
0
.9251
0
1
2
3
4
5
6
0
5
10
15
20
25
Page
12
Evaluating
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for
Protecting
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Life
in
Washington's
Surface
Water
quality
Standards
Plotting
the
growth
rates
against
the
mean
temperatures
for
both
the
constant
and
fluctuating
tests
produced
a
characteristic
normal
distribution
for
growth
rate
and
temperature
(
Figure
3.1
above).
Thus
a
strong
relationship
appears
between
the
daily
mean
temperatures
in
fluctuating
tests
and
those
in
the
constant
tests.
A
pattern
was
demonstrated,
however,
where
daily
mean
temperatures
in
the
fluctuating
tests
at
means
of
12.5
and
15.5
°
C
produced
greater
growth
rates
than
comparable
constant
temperatures
at
11.8
and
14.8
°
C.
At
the
constant
optimal
temperature
of
17.2
°
C
and
above,
however,
this
pattern
was
reversed.
This
pattern
led
Hokanson
et
al.
to
suggest
that
the
growth
of
rainbow
trout
appears
to
be
accelerated
at
fluctuating
treatments
when
the
mean
temperature
is
below
the
constant
temperature
optimum
for
growth
and
retarded
by
mean
fluctuating
temperatures
above
the
constant
temperature
optimum.
Hokanson
et
al.
suggested
that
water
quality
standards
(
based
on
weekly
mean
values)
should
be
set
such
that
the
average
weekly
temperature
is
below
the
constant
test
temperature
producing
maximum
growth.

Like
Hokanson
et
al.
(
1977),
other
researchers
(
Thomas
et
al.,
1986;
and
Everson,
1973)
have
found
that
fluctuating
temperature
regimes
actually
enhance
growth
over
what
is
found
at
constant
temperature
exposures;
at
least
where
the
mean
of
the
fluctuating
regime
is
at
or
below
that
of
the
constant
exposure
test
temperature
producing
optimal
growth.
The
works
by
these
authors
also
suggest
that
high
peak
temperatures
may
create
stress
which
will
harm
growth
even
though
the
daily
average
temperature
appears
optimal.
For
example,
Thomas
et
al.
(
1986)
noted
stress
conditions
occurring
in
cycles
with
daily
peak
temperatures
of
20
°
C.

Variable
Feeding
Regimes:

Growth
rates
are
related
to
both
temperature
and
food
rates.
As
temperature
goes
up,
more
food
is
necessary
to
supply
basic
physiological
needs
but
also
the
efficiency
goes
up
for
utilizing
excess
foods.
This
relationship
results
in
a
situation
where
at
maximum
feeding
rates
fish
will
grow
larger
in
warmer
water,
but
at
reduced
feeding
rates
in
the
same
warm
water
growth
rates
will
suffer.
In
cooler
waters,
maximum
growth
rates
are
achieved
at
feeding
rates
well
below
those
that
produced
maximum
growth
in
the
warmer
waters.
This
relationship
between
feeding
rates
and
temperature
means
that
laboratory
test
results
would
need
to
be
modified
to
account
for
the
feeding
regimes
present
in
natural
stream
environments
to
be
able
to
confidently
set
a
very
precise
maximal
growth
temperature.

Numerous
authors
have
demonstrated
that
food
availability
in
the
natural
environment
is
well
below
that
used
in
laboratory
growth
studies
(
Brett
et
al.,
1982;
Saski,
1966;
Nedham
and
Jones,
1959;
Wurtsbaugh
and
Davis,
1977;
Ensign
et
al.,
1990;
Bisson
and
Davis,
1976;
as
cited
in
ODFW,
1992;
Elliott
1975,
McCullough
1975,
1979,
Murphy
and
Hall
1981,
Edwards
et
al.
1979,
Vannote
et
al.
1980,
Bisson
and
Bilby
1998,
and
James
et
al.
1998;
as
cited
in
USEPA,
2001,
and
others).
As
an
example
of
the
influence
of
feeding
rates,
McMahon,
Zale,
and
Selong
(
1999)
tested
bull
trout
growth
and
found
that
at
satiation
(
maximum)
and
66%
satiation
ration
levels
growth
was
highest
at
16
°
C,
whereas
at
a
33%
satiation
ration
growth
rate
was
maximized
at
12
°
C.
Thus
in
waters
of
lower
productivity,
Evaluating
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Aquatic
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in
Page
13
Washington's
Surface
Water
quality
Standards
maximum
growth
may
occur
at
temperatures
well
below
those
that
produce
optimal
growth
at
high
rations.

While
the
basic
relationship
between
feeding
rates
and
growth
at
various
temperatures
is
well
established,
there
is
a
problem
with
trying
to
apply
an
adjustment
factor
to
laboratory
test
results.
In
the
laboratory
tests
reviewed
for
this
paper,
feeding
rates
and
regimes
varied
significantly.
In
addition,
the
nutritional
value
can
be
different
between
feeds,
the
size
and
type
of
food
along
with
its
method
of
presentation
can
influence
the
ability
of
fish
to
feed
and
consequently
grow,
and
the
specific
starting
size
of
the
test
fish
will
greatly
influence
growth
rates.
Given
that
the
feeding
regimes
and
test
conditions
were
so
highly
variable,
trying
to
make
a
standard
adjustment
to
laboratory
test
results
to
try
and
match
natural
feeding
regimes
is
problematic.
Nevertheless,
it
is
important
that
this
factor
be
recognized
when
setting
water
quality
criteria
recommendations
for
juvenile
rearing.

There
are
at
least
a
couple
of
ways
that
this
factor
can
be
accounted
for
without
having
to
develop
complex
growth
models
to
test
various
temperature
regime
scenarios.
The
first
is
to
make
a
standard
2­
4
°
C
adjustment
downward
to
the
temperature
value
determined
optimal
for
growth
at
satiation
feeding.
The
second
is
to
apply
the
criteria
to
a
relatively
brief
window
of
time
(
e.
g.,
2­
3
weeks),
even
though
the
growth
tests
often
lasted
4
weeks
or
more,
and
recognize
that
temperatures
will
need
to
be
below
this
value
most
of
the
time
in
even
the
warmest
years
to
result
in
compliance,
thus
virtually
ensuring
that
temperatures
will
produce
excellent
growth
overall.
In
assessing
the
risks
associated
with
making
such
simplifying
assumptions
it
is
important
to
recognize
that
growth
rates
diminish
on
either
side
of
the
optimal
temperature
range.
It
is
not
substantially
different
to
err
slightly
on
the
warmer
side
of
optimal
than
it
is
to
err
on
the
cooler
side
of
optimal,
barring
any
other
detrimental
biologic
responses.
By
extending
just
beyond
the
warmer
side
of
optimum
during
the
warmest
period
of
the
warmest
years
the
fish
will
actually
be
experiencing
a
greater
number
of
days
in
the
optimum
range
over
most
years.
In
the
recommendations
of
this
paper,
except
where
noted
for
testing
that
examined
reduced
feeding
rates,
temperatures
that
consistently
resulted
in
maximum
growth
at
satiation
feeding
were
used
to
set
the
threshold
value.

Incubation
Studies:

Specific
studies
were
not
found
that
compared
the
effects
of
constant
and
fluctuating
temperatures
on
incubating
fish.
However,
it
is
assumed
in
this
paper
that
that
incubating
fish
generally
respond
to
the
daily
mean
temperature.
This
seems
warranted
given
the
strong
basis
provided
through
the
use
of
the
standardized
"
temperature
unit"
calculations
in
hatchery
operations
and
in
fisheries
science
in
general.
It
is
fortunate
that
many
of
the
incubation
studies
were
conducted
with
fluctuating
water
temperatures
(
highlighted
in
the
discussions
of
individual
species).
This
provided
a
good
opportunity
to
generally
check
the
effect
of
applying
the
results
as
average
temperatures
and
to
assess
the
risks
of
allowing
higher
daily
maximum
temperatures.
Additionally,
a
natural
safety
factor
often
exists
to
protect
egg
incubation.
Since
salmonids
bury
their
eggs
in
the
gravel
of
the
stream
bed,
they
are
buffered
slightly
from
both
daily
maximum
and
daily
minimum
stream
temperatures.
The
buffering
of
the
stream
bed
gravel
can
effectively
reduce
the
daily
maximum
Page
14
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
temperature
by
at
least
1­
1.5
°
C
(
Crisp,
1990).
While
this
natural
safety
factor
is
not
accounted
for
in
laboratory
tests
it
is
also
not
dependable.
Therefore,
in
the
recommendations
of
this
paper
no
adjustment
is
made
to
account
for
this
often
occurring
buffering
effect.
It
is
useful,
however,
when
reviewing
the
potential
risks
of
the
recommendations
in
this
paper
to
recognize
that
sudden
and
unseasonable
rises
in
stream
temperature
during
incubation
will
often
not
cause
similar
temperature
exchanges
in
the
egg
pockets
situated
in
the
gravel.

Few
studies
examined
the
risk
of
short­
term
lethality
to
eggs
from
unusually
high
temperatures
for
individual
species,
so
the
few
that
were
conducted
are
used
broadly
to
suggest
limits
on
daily
maximum
temperatures
during
the
incubation
period
for
all
related
species.
Limits
for
daily
maximum
temperatures
are
also
based
on
the
results
from
controlled
laboratory
tests
where
the
temperature
regime
experienced
by
the
fish
was
started
at
varying
high
daily
maximum
temperatures
and
then
allowed
to
fluctuate
and
fall
in
concert
with
the
seasonal
changes
in
the
natural
waters
used
to
supply
water
for
the
tests.
From
these
experiments
it
can
be
observed
what
temperatures
at
the
start
of
the
incubation
tests
are
generally
associated
with
reduced
incubation
performance,
and
what
temperatures
appear
not
to
hinder
incubation.

Lethality
and
Acute
Effects:

The
water
quality
standards
must
be
applicable
to
a
broad
range
of
human
activities.
The
standards
must
protect
against
both
gradual
basin­
wide
increases
in
temperature
as
well
as
localized
shifts
in
daily
maximum
temperatures.
Rapid
or
site­
specific
changes
in
temperature
can
be
caused
by
unique
human
activities
(
such
as
industrial
cooling
waters,
process
wastewaters,
and
water
releases
from
reservoirs).
While
localized
extreme
changes
in
water
temperature
are
not
as
common
as
the
gradual
basin­
wide
changes,
they
do
exist,
and
their
regulation
through
discharge
permits
and
water
quality
certification
programs
require
careful
application
of
biologically­
based
temperature
standards.
These
localized
point
sources
of
temperature
change
are
also
those
most
capable
of
creating
short­
term
lethality
to
aquatic
life
as
they
may
discharge
water
significantly
hotter
or
colder
than
the
ambient
water
to
which
organisms
are
acclimated.

The
incipient
lethal
level
(
ILL)
is
typically
determined
by
exposing
juvenile
fish
to
constant
temperatures
and
determining
the
test
temperature
that
causes
50%
mortality
of
test
fish
within
typically
a
7­
day
exposure
period.
It
would
not
be
appropriate
to
establish
criteria
that
would
allow
50%
mortality,
however,
so
the
ILL
value
needs
to
be
adjusted
to
a
level
that
would
not
be
expected
to
cause
any
mortality.
The
National
Academy
of
Sciences
(
1972)
and
Coutant
(
1973)
recommend
subtracting
2
°
C
from
the
LT50
value
to
determine
a
safe
(
no
more
than
1%
mortality
 
an
LT1)
short­
term
temperature
limit.
Some
reasons
for
extra
caution
in
interpreting
lethality
studies
include
that:

1.
The
time
above
lethal
levels
appears
cumulative
(
DeHart,
1974,
1975,
and
Golden,
1978);
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
15
Washington's
Surface
Water
quality
Standards
2.
Adults
appear
more
sensitive
than
the
juveniles
which
are
most
commonly
tested
(
Coutant,
1970;
Becker,
1973;
Bouck
and
Chapman,
1975);
3.
Individual
stocks
possess
slightly
different
tolerance
levels
(
Beacham
and
Withler,
1991);
4.
Indirect
acute
effects
are
often
cited
to
occur
just
below
lethal
levels
(
e.
g.,
increased
predation,
feeding
cessation,
migration
blockage);
and
5.
The
range
between
no
mortality
and
high
mortality
rates
is
often
described
by
as
little
as
0.5
°
C
(
Charlon,
Barbier,
and
Bonnet,
1970).

The
selection
of
laboratory
results
for
use
in
developing
recommendations
focuses
on
test
fish
that
had
previously
been
acclimated
to
reasonably
cool
water
temperatures.
This
recognizes
that
fish
migrating
from
cooler
upstream
tributaries
or
from
marine
waters
may
not
be
fully
acclimated
to
warmer
main
stem
summer
river
temperatures.
It
also
helps
to
bridge
the
gap
in
protection
that
occurs
when
protecting
fish
from
hot
thermal
discharges
occurring
in
the
late
fall
through
early
spring
period
when
the
ambient
river
temperatures
are
often
very
cold.

Although
test
results
that
determine
lethal
threshold
temperatures
are
used
as
one
line
of
evidence
in
this
paper
for
recommending
water
temperature
thresholds
that
will
prevent
acute
(
short­
term)
effects,
the
final
recommendations
are
not
based
on
just
avoiding
lethality.
Temperatures
that
would
result
in
any
detrimental
acute
effect
such
as
causing
a
barrier
to
adult
fish
spawning
or
migration
take
precedence
in
the
recommendations.
Thus,
while
a
species
may
be
able
to
survive
in
a
laboratory
environment
at
a
given
temperature,
research
may
show
that
the
temperature
maximum
would
be
unacceptable
in
natural
waters.

ii)
Minimum
Temperature
Thresholds
One
issue
not
directly
included
in
the
technical
recommendations
at
this
time
are
recommendations
for
how
cold
a
water
body
can
be
allowed
to
be
due
to
human
actions.
It
is
well
documented
that
unseasonably
cold
waters
can
be
as
detrimental
to
aquatic
organisms
as
unseasonably
warm
waters.
Sudden
releases
of
very
cold
water
can
cause
rapid
lethality
to
unacclimated
organisms,
and
waters
substantially
colder
than
what
is
optimal
can
decrease
survival
of
eggs
and
embryos
and
detrimentally
depress
juvenile
growth
rates.
Human
activities,
such
as
the
removal
of
overhead
vegetative
canopies
will
increase
long­
wave
radiative
cooling
causing
potentially
harmful
drops
in
the
natural
winter
low
temperatures.
This
may
encourage
anchor
ice,
a
result
of
streams
freezing,
that
may
cause
additional
damage
to
fish
and
their
habitat
as
it
causes
super­
cooling
of
the
water
and
scours
the
channel
during
spring
break­
up
(
Jakober
et
al.,
1998;
Needham
and
Slater,
1944;
Needham
et
al.,
1945
Maciolek
and
Needham,
1952).
It
may
be
that
thresholds
for
low
temperature
limits
should
be
considered
at
some
point
in
the
future,
and
information
is
included
in
this
paper
to
assist
in
setting
site­
specific
limits
for
individual
activities.
However,
to
keep
the
focus
of
this
paper
on
the
area
of
greatest
concern,
that
of
high
water
temperatures,
the
issue
of
cold
water
limits
is
not
being
addressed
at
this
time.
Page
16
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
iii)
Protection
of
Untested
Species
The
technical
recommendations
made
in
this
paper
are
primarily
based
upon
data
obtained
on
the
temperature
requirements
of
our
two
native
char,
the
five
Pacific
salmon,
rainbow
and
cutthroat
trout,
smelt,
mountain
whitefish,
several
stream
breeding
amphibians,
and
numerous
insect
species.
Rigorous
scientific
testing
generally
has
not
been
done
on
most
of
the
other
aquatic
species
indigenous
to
Washington's
waters.
What
references
have
been
found
for
these
other
species
tends
to
be
of
a
more
anecdotal
nature;
noting
what
the
temperature
was
when
spawning
was
observed,
or
indicating
the
temperature
of
a
waterbody
where
a
species
was
found.
For
some
of
these
less
well
known
indigenous
species,
however,
direct
testing
of
their
temperature
optima
and
lethal
limits
has
occurred.
Based
on
what
temperature
information
has
been
found,
both
direct
and
indirect,
these
other
species
do
not
appear
to
possess
more
sensitive
temperature
requirements
than
the
species
that
have
been
used
to
make
temperature
threshold
recommendations
in
this
document.

Many
of
the
less
well
known
aquatic
species,
which
often
occur
together
as
a
community,
appear
to
thrive
in
waters
which
are
at
the
upper
margin
of
what
is
optimal
for
salmonids,
and
represent
warm
water
fish
communities
indigenous
to
Washington.
Recognition
of
these
species
has
resulted
in
the
recommendation
that
the
state
establish
a
unique
level
of
usesupport
that
can
be
assigned
to
our
naturally
warmer
waters.

The
warm
waters
that
occur
in
the
margins
and
slack­
water
areas
of
many
rivers
serve
as
important
habitat
to
many
species
that
prefer
summer
water
temperatures
somewhat
warmer
than
what
is
healthy
for
the
state's
indigenous
salmonids.
These
other
species
also
commonly
serve
as
food
fish
for
salmonids
and
are
important
to
the
overall
health
of
the
system.
Thus,
these
warmer
stream
margins
should
not
be
viewed
as
lost
or
unhealthy
habitat,
but
as
a
necessary
and
natural
adjunct
to
a
healthy
aquatic
system.
It
is
important
to
note
that
young
salmonid
fry
also
often
initially
rear
in
the
shallow
margins
of
rivers
until
reaching
a
size
that
allows
them
to
habitat
the
more
active
channel,
and
that
these
young
fry
have
been
commonly
shown
to
have
slightly
higher
temperature
preferences
than
older
fry
or
juvenile
fish.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
17
Washington's
Surface
Water
quality
Standards
4.
Temperature
Requirements
of
Char,
Salmon,
and
Trout
Species
In
this
chapter,
the
temperature
requirements
of
the
char,
salmon,
and
trout
species
indigenous
to
the
state
of
Washington
are
examined.
Whenever
the
data
allowed
doing
so,
two
temperature
values
are
provided
to
define
the
upper
end
of
the
temperature
range
that
is
fully
protective
of
each
species'
key
life
stages.
This
was
generally
done
for
the
primary
metric
recommended
which
is
a
7­
day
average
of
daily
maximum
temperatures.
The
approach
taken
recognizes
that
the
upper
fully
protective
endpoint
is
often
more
accurately
described
by
defining
a
narrow
range
of
temperature
rather
than
a
single
value.
The
final
selection
of
a
single
recommended
value
comes
only
after
the
needs
of
all
of
the
species,
their
life­
stages,
and
stressors
are
considered
together
[
see
Summary
Section
4
g)].

a)
Native
Char
Temperature
Requirements
i)
General
Life
History
Information:

Bull
trout
(
Salvelinus
confluentus)
and
Dolly
Varden
(
Salvelinus
malma)
are
the
only
two
species
of
char
native
to
Washington
(
Hass
and
McPhail,
1991).
Perhaps
more
than
any
other
species,
cold
waters
are
critical
to
maintaining
healthy
populations
of
these
native
char.
These
two
closely
related
species
are
difficult
to
taxonomically
identify
from
one
another
in
the
field
(
Hass
and
McPhail,
1991).
Cavender
(
1978)
may
have
first
recognized
that
what
had
previously
been
considered
an
interior
form
of
Dolly
Varden
was
in
fact
a
distinct
species,
now
referred
to
as
the
bull
trout.
Goetz
(
1989)
suggests
that
bull
trout
may
be
more
directly
related
to
the
Arctic
char
(
Salvelinus
alpinus),
and
in
fact
a
sister
to
the
Arctic
char­
Dolly
Varden
group.
Spalding
(
1997)
found
that
for
Washington's
Olympic
Peninsula
many
of
the
anadromous
char
previously
assumed
to
be
Dolly
Varden
keyed
out
to
be
bull
trout.
While
important
to
advancing
scientific
understanding,
the
historic
problems
with
discriminating
between
these
char
appear
to
pose
little
practical
problems
in
terms
of
setting
water
quality
criteria.
This
is
because
Dolly
Varden
and
bull
trout
are
generally
considered
to
have
very
similar
biological
requirements,
and
the
management
measures
needed
to
protect
both
Dolly
Varden
and
bull
trout
may
be
identical
(
WSDFW,
1994).

The
Washington
State
Department
of
Fish
and
Wildlife
considers
the
majority
of
native
char
stocks
in
the
state
as
being
"
Vulnerable
Populations"
requiring
special
protection.
Bull
trout
in
particular
have
received
considerable
publicity
in
recent
years
because
of
their
status
as
a
threatened
species
under
the
federal
Endangered
Species
Act.
The
results
of
a
12­
month
study
evaluating
stock
status
by
U.
S.
Fish
and
Wildlife
Service
noted
serious
declines
of
bull
trout
populations
statewide
(
USFWS,
1997).
Rieman
et
al.
(
1997)
examined
the
distribution
and
status
of
bull
trout
across
4,462
sub­
watersheds
of
the
interior
Columbia
River
basin
and
the
Klamath
River
basin
and
found
that
bull
trout
are
more
likely
to
occur
and
the
populations
are
more
likely
to
be
strong
in
colder,
higher­
elevation,
low­
to
mid­
order
watersheds
with
lower
road
densities.
They
noted
that
while
bull
trout
were
widely
Page
18
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
distributed
across
their
potential
range,
strong
populations
may
exist
in
only
6%
of
this
potential
range.

Upon
hatch,
char
fry
will
either
remain
in
the
localized
area
or
move
downstream
to
larger
streams
or
lakes
to
rear
(
Goetz,
1989;
Williams
and
Mullen,
1992;
Reiser
et
al.,
1997).
Movements
to
more
suitable
upstream
waters
has
also
been
observed
(
Fraley
and
Shepard,
1989;
Armstrong
and
Morrow,
1980,
as
cited
in
Goetz,
1989).
Unless
information
clearly
demonstrates
that
a
stretch
of
water
would
not
be
used
for
summer
rearing
even
under
natural
conditions,
temperature
criteria
assigned
to
these
migration
paths
should
also
be
set
to
protect
the
rearing
of
char
fry.

Some
research
suggests
that
age
0­
1
juvenile
char
have
cooler
temperature
preferences
than
age
1+
juveniles;
however,
there
is
little
consistency
in
the
values
identified.
The
preference
values
for
age
0­
1
bull
trout
range
from
an
average
of
4.5
°
C
(
Ratliff,
1992),
to
maximum
stream
temperatures
of
10
°
C
and
13
°
C
(
Ratliff,
1987;
and
Martin
et
al.,
1991).
Resident
forms
of
bull
trout
remain
in
or
near
their
natal
streams
for
their
entire
life.
Fluvial
and
adfluvial
forms
may
remain
in
the
area
of
their
natal
stream
for
1
to
3
years
and
then
migrate
significant
distances
to
more
productive
waters
for
greater
juvenile
growth
opportunities
(
Pratt,
1992;
Ratliff,
1992;
Riehle
et
al.,
1997;
Fraley
and
Shepard,
1989;
Goetz,
1989);
although,
some
stocks
have
also
been
observed
to
migrate
to
lakes
or
reservoirs
immediately
after
hatching
(
Reiser
et
al.,
1997).
Sea­
run
(
anadromous)
forms
will
migrate
hundreds
of
miles
to
take
advantage
of
productive
near­
shore
marine
habitat
(
Goetz,
1989).
Temperature
standards
would
ideally
be
set
in
consideration
of
these
various
life­
strategies.

The
needs
of
resident
forms
may
be
slightly
different
from
the
various
migratory
forms.
Juvenile
bull
trout
and
Dolly
Varden
have
difficulty
competing
with
several
common
salmonid
species
in
warmer
waters
(
Haas,
2001;
McMahon
et
al.,
1999).
This
may
partially
explain
why
researchers
have
observed
young
juvenile
fish
remaining
in
their
natal
stream
for
the
first
several
years
before
moving
downstream
to
warmer
and
more
productive
waters.
It
is
plausible
that
the
lack
of
significant
competitors
in
their
cold
natal
streams
may
compensate
for
the
reduced
productivity
of
these
pristine
environments.
Since
nonmigratory
resident
bull
trout
must
remain
in
and
defend
their
natal
habitat
for
their
entire
life,
temperatures
here
should
clearly
favor
bull
trout
over
competing
species
such
as
chinook
salmon
and
rainbow
trout
(
Martin
et
al.,
1991;
Mullan
et
al.,
1992;
Ziller,
1992;
Adams
and
Bjornn,
1997;
WSDFW,
1994;
Haas,
2001).
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
19
Washington's
Surface
Water
quality
Standards
ii)
Spawning
Requirements
Field
Observations
of
Spawning
Initiation
Maximum
temperatures
should
generally
be
below
12
°
C
and
on
a
fall
season
cooling
trend
at
the
time
char
enter
their
spawning
streams
(
Fraley
and
Shepard,
1989).
In
a
study
on
the
Rapid
River
in
Idaho,
pairing
behavior
was
noted
to
begin
after
average
water
temperatures
dropped
from
10
to
6.5
°
C
(
generally
equivalent
to
a
change
in
single
daily
maximum
temperatures
from
11­
7.5
°
C)
(
Schill,
Thurow,
and
Kline,
1994).
In
the
same
river,
Elle
and
Thurow
(
1994)
found
that
daily
maximum
water
temperatures
below
10
°
C
influenced
the
movements
of
spawners
both
in
and
out
of
the
Rapid
River.
While
daily
maximum
temperatures
may
need
to
fall
below
9
to
11
°
C
(
WSDFW,
1994)
for
redd
construction
to
begin,
no
authors
have
been
found
to
suggest
spawning
will
begin
at
daily
maximum
temperatures
above
10
°
C.
Most
place
the
temperature
that
triggers
spawning
below
9
°
C
(
Goetz,
1989;
Pratt,
1992;
Kramer,
1991;
Fraley
and
Shepard,
1989),
with
the
peak
of
spawning
activity
not
occurring
until
stream
temperatures
falls
below
7
°
C
(
James
and
Sexauer,
1997;
Wydoski
and
Whitney,
1979).
Reiser
et
al.
(
1997)
suggested
that
a
daily
average
temperature
between
about
6.8
to
8.1
°
C
(
generally
equivalent
to
a
single
daily
maximum
range
of
7.8­
9.1
°
C)
was
necessary
to
initiate
spawning
activity.
Temperatures
above
8
°
C
were
noted
by
Kramer
(
1994)
as
appearing
to
cause
spawning
activity
to
temporarily
cease
in
char
in
northwest
Washington
streams.
The
Washington
Water
Power
Company
(
1995)
studied
the
distribution
of
fish
in
the
lower
Clark
Fork
River
in
Idaho
and
found
that
bull
trout
spawning
was
confined
to
an
artificial
spawning
channel
created
to
mitigate
the
effects
of
the
Cabinet
Gorge
Dam.
Water
temperatures
in
theses
spawning
areas
were
consistently
cooler
(
5­
7
°
C)
than
other
areas
of
the
channel,
and
during
the
period
of
redd
construction
bull
trout
used
the
area
of
the
channel
where
the
temperature
was
11
°
C
due
to
ground
water
seeps.

Bull
trout
are
noted
to
begin
spawning
as
soon
as
conditions
are
suitable
and
redds
are
constructed.
Temperature
may
be
the
primary
cue
used
by
the
fish
to
determine
when
to
begin
migratory
movements
(
Elle
and
Thurow,
1994;
Swanberg,
1997)
as
well
as
to
initiate
spawning
(
Kramer,
1994).
It
is
important
that
temperatures
at
the
initiation
of
spawning
be
within
a
range
that
would
not
hinder
ovulation
and
would
not
cause
obvious
harm
to
offspring
of
any
early
spawning
individuals.

The
field
observations
and
citations
noted
above
are
in
strong
concurrence
that
spawning
behavior
(
pairing
and
redd
construction)
will
not
begin
7DADMax
temperatures
fall
below
8.45­
9.45
°
C,
and
that
spawning
itself
will
only
be
initiated
once
the
daily
maximum
temperature
falls
below
7.45­
8.45
°
C.
Page
20
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Laboratory
Studies
of
Spawning
Initiation
Only
a
single
study
was
found
that
addressed
the
question
of
what
temperatures
are
required
to
initiate
spawning.
In
a
study
by
Gillet
(
1991)
ovulation
in
char
was
found
to
be
completely
inhibited
at
constant
temperatures
above
11
°
C
and
slowed
down
above
8
°
C
as
compared
to
fish
held
at
5
°
C.
This
would
be
correlated
with
7DADMax
values
of
11.62­
11.93
°
C
and
8.62­
8.93
°
C,
depending
upon
the
constant
test
exposures
being
treated
as
if
they
represent
on­
day
average
and
seven­
day
daily
average
stream
exposures.
Transfers
from
8
°
C
to
5
°
C
were
found
to
stimulate
ovulations.
Gillet
also
found
that
exposure
to
temperatures
of
8
°
C
prior
to
ovulation
were
favorable
to
fecundity
rates,
and
assumed
based
on
their
work
that
very
cold
water
was
only
necessary
during
the
last
weeks
before
spawning.
The
work
of
Gillet
suggests
that
for
the
closely
related
Arctic
char,
temperatures
should
be
falling
below
8
°
C
to
stimulate
healthy
spawning,
and
falling
towards
5
°
C
to
ensure
full
survival
of
fertilized
eggs
(
only
11,
8,
an
5
°
C
were
tested,
so
specific
incubation
thresholds
were
not
determined).
Based
on
this
one
laboratory
study
it
would
appear
that
a
7DADMax
of
less
than
8.62­
8.93
°
C
is
necessary
to
allow
healthy
ovulation
to
occur
in
bull
trout.

Summary
on
Spawning
Initiation
Requirements
There
is
strong
agreement
among
both
field
and
laboratory
research
and
observations
on
the
spawning
requirements
of
bull
trout.
The
weight
of
the
evidence
supports
the
position
that
initial
spawning
behavior
(
pairing
and
redd
construction)
is
hindered
by
daily
maximum
stream
temperatures
above
9­
11
°
C
and
that
the
act
of
spawning
may
be
impaired
by
daily
maximum
water
temperatures
greater
than
8­
9
°
C.

Laboratory
Studies
on
Incubation
Success
Only
a
few
studies
were
found
that
tested
the
incubation
requirements
of
bull
trout
in
the
laboratory.
Fredenberg
(
1992)
reported
that
eggs
from
wild
bull
trout
incubated
at
an
average
temperature
of
3.1
°
C
had
egg
survival
averaging
97.4%
from
unfertilized
egg
to
eyeup
and
97.1%
from
unfertilized
egg
to
hatch.
Eggs
were
subsequently
collected
from
the
hatchery
raised
broodstock
and
incubated
at
an
average
temperature
of
5.8
°
C
(
5.0
°
C
minimum
and
6.6
°
C
maximum),
and
while
no
growth
or
survival
data
was
collected
the
author
suggested
that
it
appeared
normal.
Fredenberg,
Dwyer,
and
Barrows
(
1995)
collected
gametes
from
wild
spawning
bull
trout
from
the
Swan
River
drainage
of
northwest
Montana
during
September
of
1993
and
1994.
Fertilized
eggs
were
produced
by
paired
matings
and
incubated
in
1993
at
approximately
3.1
°
C
with
a
resultant
97.1%
survival
from
green
eggs
to
hatching.
Eggs
in
1994
were
incubated
at
approximately
6.5
°
C
with
a
resultant
95.5%
survival.

McPhail
and
Murry
(
1979)
tested
incubation
of
bull
trout
at
constant
laboratory
temperatures
of
2,
4,
6,
8,
and
10
°
C
in
a
series
of
three
replicate
tests.
In
the
one
test
lot
that
escaped
high
transit­
related
mortality
during
movement
to
the
laboratory
after
fertilization,
McPhail
and
Evaluating
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Page
21
Washington's
Surface
Water
quality
Standards
Murry
(
1979)
found
that
survival
was
excellent
(
90­
95%)
at
2,
4,
and
6
°
C,
good
(
85%)
at
8
°
C,
and
poor
(
20%)
at
10
°
C.
In
the
two
other
test
lots,
which
experienced
significant
transit­
related
stress
(
40%
egg
mortality
during
transit),
survival
rates
were
noted
to
steadily
decline
from
2
to
4,
to
6
°
C
and
drop
to
zero
at
8
and
10
°
C.
Similarly,
constant
test
temperatures
in
the
range
of
7­
11
°
C
were
reported
to
result
in
"
poor"
survival
in
hatchery
culture
by
Brown
(
1985).
McPhail
and
Murry
(
1979)
noted
that
mortalities
at
low
temperatures
(<
6.0C)
typically
occurred
at
blastopore
closure,
while
at
high
temperatures
(>
8.0)
mortality
is
associated
with
hatching.
In
studies
on
the
related
species
of
Arctic
char,
Humpesch
(
1985)
reported
optimal
incubation
to
occur
at
5
°
C.

The
above
studies
generally
found
that
constant
temperatures
in
the
range
of
3.1­
6.5
°
C
capable
of
producing
excellent
(
90­
97.4%)
survival
of
char
eggs,
with
survivals
equally
high
throughout
the
range.
At
temperatures
8
to
10
°
C
survival
rates
may
drop
precipitously
from
85%
to
0%.
The
research
suggests
that
a
constant
temperature
of
6.5
°
C
may
most
confidently
define
the
upper
limit
for
fully
protecting
incubation.
This
would
be
equivalent
to
a
7DADMax
temperature
of
7.43
°
C
when
correlated
as
if
it
were
a
one­
week
average
daily
exposure,
and
equivalent
to
a
7DADMax
temperature
of
8.5
°
C
when
correlated
as
if
it
were
an
incubation
season­
long
average
exposure
(
assuming
a
stable
stream
temperature
with
0­
2
°
C
diel
variation).
Based
on
the
above,
the
laboratory
studies
suggest
full
incubation
protection
will
occur
if
the
highest
7DADMax
temperature
during
the
incubation
period
does
not
exceed
7.43­
8.5
°
C.
It
is
important
to
note
that
this
process
is
not
trying
to
determine
an
acute
threshold
for
eggs
or
embryos.
It
only
identifies
temperatures
that
can
occur
at
the
initiation
of
incubation
that
will
fully
protect
native
char
incubation
under
a
normal
temperature
regime
(
i.
e.,
fall
cooling
trends).

It
is
important
to
note
that
even
though
2
°
C
has
been
shown
to
be
suboptimal
at
a
constant
incubation
temperature,
natural
seasonal
declines
in
temperature
down
to
2
°
C
in
the
incubation
period
are
unlikely
to
reduce
survival
rates.
Salmonids
have
been
shown
to
undergo
conditioning
in
the
early
stage
of
incubation
that
allows
excellent
survival
at
very
low
temperatures
occurring
later.
Where
the
conditioning
does
not
occur,
and
the
eggs
are
incubated
at
an
early
stage
at
very
low
temperatures,
significant
reductions
in
survival
have
been
noted
(
Murry
and
Beacham,
1986;
Seymour,
1956).
This
assertion
is
also
supported
by
work
showing
that
newly
hatched
bull
trout
alevins
are
tolerant
of
temperatures
near
0
°
C
(
Baroudy
and
Elliott,
1985)
and
that
the
lower
limit
for
hatching
in
the
related
Arctic
char
is
less
than
1
°
C.
Such
conditioning,
however,
may
not
only
protect
embryos
from
later
exposure
to
colder
waters
but
may
help
increase
tolerance
for
warm
water
fluctuations
as
well.
Bebak,
Hankins,
and
Summerfelt
(
2000)
examined
the
hatching
success
and
posthatch
survival
of
three
stocks
of
Arctic
char
and
found
that
if
incubation
had
been
well
initiated
at
a
favorable
(
6
°
C)
temperature
then
later
transfers
to
waters
as
warm
as
10­
12
°
C
still
allowed
for
excellent
hatch
rates
(
90­
98%).
It
is
noteworthy,
however,
that
they
also
found
that
survival
rates
post­
hatch
declined
over
time
at
both
10
°
C
and
12
°
C.

Field
Evidence
on
Incubation
Requirements
Page
22
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Little
evidence
was
found
from
field
studies
on
the
incubation
requirements
of
bull
trout
or
Dolly
Varden.
In
one
study
that
was
reviewed,
however,
it
was
noted
that
bull
trout
redds
in
the
upper
Flathead
River
basin
in
Montana
had
mean
temperatures
that
ranged
from
2.1­
5.4
°
C
(
Weaver
and
White,
1985).
This
season­
long
estimate
of
average
field
temperatures
would
correlate
with
a
7DADMax
temperature
range
of
4.1­
7.4
°
C.
It
is
important
to
point
out
that
just
noting
the
average
temperature
of
redds
does
not
indicate
whether
it
was
healthy,
and
is
not
very
useful
for
describing
of
the
upper
boundary
for
successful
incubation.
For
this
reason,
only
the
upper
half
of
the
range
will
be
used
to
represent
this
line
of
evidence,
the
next
preferred
option
would
be
to
not
use
this
range
at
all
to
estimate
incubation
requirements.
Based
on
the
above
discussion,
field
temperatures
of
bull
trout
redds
should
be
considered
to
have
an
upper
range
of
5.75­
7.4
°
C.

Conclusion
on
Spawning
and
Incubation
While
spawning
and
incubation
were
discussed
separately
above,
it
is
important
to
recognize
they
actually
occur
concurrently
in
the
natural
stream
environment.
Once
spawning
has
occurred
the
eggs
begin
to
incubate.
The
multiple
lines
of
evidence
created
from
the
wide
variety
of
field
and
laboratory
studies
can
be
brought
together
in
support
of
selecting
a
temperature
standard
for
application
at
the
beginning
of
the
spawning
and
incubation
period.
The
focus
on
the
temperature
at
the
initiation
of
incubation
is
viewed
as
appropriate
since
temperatures
will
be
at
their
highest
at
this
time.
They
should
also
be
on
a
trend
of
fall
cooling
that
will
soon
reach
temperatures
that
are
naturally
determined
by
the
elevation
and
geographic
location
of
the
site.
The
interim
conclusions
from
each
line
of
evidence
are
summarized
below:

Table
4.1.
Spawning
and
Incubation
requirements
of
native
char.

Line
of
Evidence
7DADM
­
Range
(
°
C)
Median
(
°
C)
Field
observations
on
spawning
initiation
7.45­
8.45
7.95
Laboratory
studies
on
spawning
initiation
8.62­
8.93
8.77
Laboratory
studies
on
incubation
success
7.43­
8.5
7.97
Field
study
on
typical
average
redd
temperatures
in
Montana
5.75­
7.4
6.57
Best
Estimate
of
Threshold
7.31­
8.32
mid.
pt.
7.82
The
range
of
these
independent
lines
of
evidence
is
5.75­
8.93
°
C
with
a
mean
range
of
7.31­
8.32
°
C
and
an
overall
midpoint
of
7.82
°
C.
This
strongly
suggests
that
a
7­
day
average
daily
maximum
temperature
of
7.82
°
C
will
fully
protect
the
spawning
and
incubation
of
char.
In
recognition
that
temperatures
of
8.0
°
C
have
been
noted
as
hindering
spawning
in
char,
it
may
not
be
advisable
in
this
situation
to
select
a
spawning
threshold
value
from
the
upper
end
of
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
23
Washington's
Surface
Water
quality
Standards
the
range
presented
above.
It
is
concluded
that
a
7­
day
average
of
the
daily
maximum
temperatures
of
7.5­
8.0
°
C
best
represents
the
highest
temperature
that
can
occur
at
the
initiation
of
spawning
without
causing
detrimental
effects
to
either
spawning
or
subsequent
incubation.

iii)
Juvenile
Rearing
Field
Distribution
Work
Examining
Fish
Presence
Dunham
et
al.
(
2001)
surveyed
the
distribution
of
juvenile/
small
bull
trout
(
Note:
small
resident
stock
fish
cannot
be
distinguished
apart
from
juveniles
of
migratory
stocks)
in
6
basins
(
109
sites)
in
Washington
state
and
found
that
summer
maximum
temperatures
best
defined
their
presence
in
streams.
In
their
study,
95%
of
the
bull
trout
were
found
in
streams
with
temperatures
above
12
°
C
with
an
interquartile
range
of
13.3­
14.7
°
C
(
with
the
median
and
mode
both
at
14
°
C).
This
is
similar
to
a
study
by
Rieman
and
Chandler
(
1999)
who
examined
581
sites
throughout
the
Western
United
States
and
found
that
the
majority
of
sites
with
juvenile/
small
bull
trout
had
summer
maximum
temperatures
of
11­
14
°
C
(
95%
were
from
waters
with
summer
maximums
less
than
18
°
C).
Goetz
(
1997b)
surveyed
13
drainages
in
Washington
and
Oregon
and
was
unable
to
find
juvenile
bull
trout
in
streams
with
temperatures
above
14
°
C.
In
a
study
in
British
Columbia,
Hass
(
2001)
found
that
the
warmest
study
streams
containing
bull
trout
(
resident
juveniles
or
adults)
had
summer
maximum
temperatures
of
16
°
C,
which
is
similar
to
the
findings
of
numerous
other
researchers
that
15­
16
°
C
formed
the
upper
temperature
limit
to
bull
trout
(
juveniles
or
adults)
summer
distributions
(
Fraley
and
Shepard
1989,
Shepard
1985,
Goetz,
1989,
Pratt,
1992,
Martin
et
al.
1991)
and
that
a
temperature
of
15
°
C
can
trigger
the
out­
migration
of
char
from
otherwise
suitable
habitat
(
Goetz,
1997).
The
general
findings
for
bull
trout
discussed
above
are
also
supported
by
the
work
of
Jensen
(
1981)
who
found
that
14
°
C
(
as
a
10­
day
mean)
appeared
to
form
a
barrier
to
the
distribution
of
the
closely
related
Arctic
char.

The
above
studies
demonstrate
that
daily
maximum
temperatures
in
the
range
of
14­
15
°
C
(
7DADMax
13.45­
14.45
°
C)
set
the
upper
boundary
for
commonly
finding
juvenile/
small
bull
trout.
It
is
important
to
note
that
some
researchers
have
both
wider
and
narrow
margins
of
distribution.
Some
authors
have
found
that
colder
summer
maximum
temperatures
(
10­
11.5
°
C)
formed
the
general
limits
to
bull
trout
distributions
in
specific
watersheds
(
Ratliff,
1987).
While
other
authors
have
reported
finding
juvenile
bull
trout
at
much
warmer
(
17­
20.5
°
C)
temperatures
(
Brown,
1992;
Goetz,
1989,
Adams
and
Bjornn,
1997;
Reiman
and
Chandler,
1999).
It
was
noted
by
Adams
(
1999)
that
bull
trout
found
at
20.5
°
C
held
in
the
coolest
water
available
in
the
area
(<
17.2
°
C)
and
looked
physically
unhealthy.
This
point
is
made
to
remind
the
reader
that
the
mere
presence
of
a
fish
does
not
indicate
that
it
is
healthy
or
that
the
stream
is
capable
of
supporting
healthy
populations.
Page
24
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Field
Work
that
Considered
Dominance,
Density,
and
Competitive
Advantage
While
studies
that
describe
the
stream
temperatures
associated
with
the
presence
of
a
species
are
a
useful
line
of
evidence,
they
cannot
be
used
to
say
whether
or
not
the
population
is
in
good
health
at
the
observed
temperature
regimes.
Researchers
have
tried
to
answer
the
question
of
what
is
healthy
for
bull
tout
in
field
studies
by
identifying
sites
with
high
densities
of
bull
trout,
and
by
identifying
temperatures
beyond
which
bull
trout
begin
to
loose
dominance
over
other
competing
species.
Health
has
also
been
assessed
more
directly
by
evaluating
the
condition
of
the
fish
using
standard
bio­
metrics.

Haas
(
2001)
found
that
a
7­
day
average
of
the
daily
maximums
of
11.6
°
C
(
correlating
with
a
single
daily
maximum
of
12.15
°
C)
consistently
determined
the
dominant
presence
and
better
condition
of
bull
trout
in
26
sites
in
the
Columbia
River
drainage
in
British
Columbia.
Haas
reasoned
that
bull
trout
populations
would
be
supported
by
maintaining
summer
maximum
temperatures
below
13
°
C,
and
noted
that
rainbow
trout
were
found
to
be
dominant
over
bull
trout
and
in
better
condition
as
summer
maximum
temperatures
approached
14­
15
°
C.
This
corresponds
well
to
the
Dunham
et
al.
(
2001)
finding
that
90%
of
the
sites
that
did
not
have
bull
trout
had
summer
maximum
temperatures
greater
than
13.5
°
C
(
non­
char
sites
had
an
interquartile
range
of
14.6­
19.8
°
C
and
a
median
of
17
°
C).
It
also
corresponds
well
with
Williams
and
Mullen
(
1992)
finding
that
rainbow
trout
excluded
the
first
two
age
classes
of
bull
trout
at
weekly
average
temperatures
above
approximately
11­
12
°
C
(
correlating
with
single
day
maximums
of
12.49­
13.49
°
C).
Williams
and
Mullan
(
1992)
found
that
bull
trout
growth
in
the
Early
Winters
Creek
basin
of
northern
Washington
steadily
increased
with
increasing
temperatures
at
three
synchronous
test
sites
having
annual
maximum
weekly
average
temperatures
of
8.7,
10.3,
and
11.7
°
C,
respectively.
The
maximum
weekly
average
of
11.7
°
C
correlates
to
a
7DADMax
of
12.63
°
C.
Growth
was
not
tested
at
any
warmer
sites,
thus
the
7DADMax
temperature
that
would
allow
for
optimal
growth
in
this
stream
system
was
not
identified,
but
would
likely
be
greater
than
12.6
°
C.
Similarly,
Martin
et
al.
(
1991)
and
Goetz
(
1989,
1997)
concluded
that
bull
trout
are
dominant
in
streams
with
summer
maximum
temperatures
less
than
13
°
C.
Saffel
and
Scarnecchia
(
1999)
examined
18
reaches
of
six
streams
and
found
that
the
density
of
bull
trout
increased
with
increasing
maximum
temperature
below
14
°
C
(
range
7.8­
13.9
°
C)
and
decreased
with
increasing
temperature
above
18
°
C
(
range
18.3­
23.3
°
C).
The
highest
densities
were
found
in
reaches
with
maximum
summer
temperatures
between
10­
13.9
°
C.
In
this
work
there
were
no
study
streams
having
summer
maximum
temperatures
within
the
range
of
14­
18
°
C,
so
it
cannot
be
said
whether
or
not
densities
would
have
begun
to
decline
just
above
14
°
C.
Kitano
et
al.
(
1994)
found
that
brook
trout
and
cutthroat
trout
coexisted
with
bull
trout
in
the
Flathead
River
basin
in
Montana
in
waters
with
a
temperature
range
of
5.3­
8.9
°
C
in
early
September,
but
they
provided
no
information
on
the
relative
health
of
these
populations
or
on
when
competition
would
be
impaired.

Based
on
the
above
works,
it
is
reasonable
to
expect
that
streams
with
summer
maximum
temperatures
of
12­
14
°
C
(
7DADMax
11.45­
13.45
°
C)
will
be
capable
of
producing
and
maintaining
strong
populations
of
bull
trout.
Not
all
of
the
research
examined,
however,
fully
supports
this
assertion.
Sexauer
and
James
(
1997)
studied
four
bull
trout
streams
in
the
Yakima
and
Wenatchee
River
watershed
in
central
Washington
that
were
selected
because
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
25
Washington's
Surface
Water
quality
Standards
they
were
considered
"
healthy"
and
found
summer
maximum
temperatures
in
the
year
the
study
was
conducted
ranged
from
9­
12.5
°
C.
Further,
Ratliff
(
1992)
and
Ziller
(
1992),
reported
that
bull
trout
began
to
lose
dominance
as
summer
average
temperatures
rose
above
7.9
°
C
(
correlating
with
single
day
maximums
of
10.45
°
C).

The
wealth
of
studies
across
the
Northwest
when
considered
in
combination
demonstrate
some
strong
patterns
of
occurrence.
Any
study
of
a
single
basin
or
stream
needs
to
be
carefully
considered
prior
to
accepting
its
conclusions.
For
example,
in
the
cold
groundwater
dominated
Metolius
River
system
in
Oregon,
Ratliff
(
1987)
reported
that
bull
trout
were
rarely
found
at
temperatures
above
10
°
C.
While
true,
it
is
important
to
recognize
that
warmer
waters
are
largely
unavailable
in
this
tributary
system.
Thus
care
should
be
exercised
before
concluding
that
10
°
C
has
been
shown
to
create
a
barrier
to
bull
trout
populations.
Similarly,
many
of
the
studies
were
conducted
over
a
single
year,
and
thus
the
long­
term
relationship
between
the
presence
healthy
populations
and
maximum
ambient
temperatures
at
the
site
is
often
not
documented.
So
while
Sexauer
and
James
(
1997)
studied
streams
with
healthy
populations,
they
only
reported
temperatures
from
a
single
year.
The
actual
temperature
regime
for
these
sites,
which
hold
healthy
char
populations,
may
include
years
with
warmer
temperatures
than
those
observed
during
the
study
year.

Summary
of
Evidence
from
Field
Studies
on
Juvenile
Rearing
A
preponderance
of
juvenile/
small
bull
trout
in
Washington
may
be
found
in
streams
with
summer
maximum
temperatures
between
13.3­
14.7
°
C
(
7DADMax
12.75­
14.15
°
C).
This
is
reasonably
consistent
with
surveys
extending
throughout
the
Western
states
that
found
most
juvenile
use
to
occur
in
stream
segments
with
maximum
temperatures
not
exceeding
14
°
C.
It
is
also
well
supported
by
studies
of
in­
stream
growth,
density,
and
competitive
dominance
which
suggest
healthy
bull
trout
streams
have
summer
temperatures
that
do
not
exceed
12­
14
°
C.
While
field
studies
are
a
very
useful
line
of
evidence
for
developing
temperature
criteria,
they
can
be
strengthened
by
cross­
checking
the
conclusions
with
laboratory
findings.

Laboratory
Studies
Supporting
Juvenile
Rearing
In
a
controlled
hatchery
environment,
Fredenberg
et
al.
(
1995)
found
that
constant
exposure
to
8.3
°
C
produced
greater
growth
of
bull
trout
at
maximum
rations
than
at
lower
test
temperatures.
This
would
be
equivalent
to
a
single­
day
maximum
of
9.79
°
C
(
7DADMax
9.24
°
C)
if
the
constant
hatchery
temperature
is
treated
as
it
represented
the
warmest
oneweek
average
stream
temperature;
and
a
single­
day
maximum
of
10.85
°
C
(
7DADMax
10.3
°
C)
if
it
were
equivalent
to
a
summer
season­
long
average
stream
temperature.

In
a
laboratory
study,
McMahon
et
al.
(
1998)
found
that
growth
was
highest
at
a
constant
12
°
C,
but
not
significantly
less
at
14
and
16
°
C
at
maximum
rations.
Growth
declined
sharply
at
temperatures
greater
than
18
°
C
and
less
than
10
°
C.
In
follow­
up
tests,
the
authors
(
McMahon
et
al.,
1999)
found
that
growth
at
maximum
and
at
66%
of
maximum
rations
Page
26
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
were
highest
at
16
°
C,
but
that
at
33%
of
maximum
ration
growth
was
maximized
at
12
°
C.
In
modeling
the
available
calories
in
low
productivity
streams
(
the
ration
used
and
the
basis
for
its
selection
was
not
provided)
against
the
growth
observed
in
their
tests,
the
authors
suggested
the
optimum
growth
range
of
12­
16
°
C
would
shift
to
8­
12
°
C.
McMahon
et
al.
(
2000)
examined
the
growth
of
bull
trout
over
a
60­
day
period
in
both
constant
and
fluctuating
temperature
regimes.
Peak
growth
at
a
moderately
restricted
ration
(
0.66)
occurred
at
12.4
°
C
at
a
constant
temperature
and
at
a
mean
of
12.2
°
C
in
fluctuating
treatments
(+/­
3
°
C
around
the
mean
­
giving
a
60­
day
average
daily
maximum
of
15.2
°
C).
Peak
growth
at
an
unrestricted
ration
occurred
at
13.2
°
C
and
at
a
severely
restricted
ration
(
0.11)
occurred
at
12.3
°
C.
In
this
work
the
authors
found
that
growth
was
higher
in
constant
exposure
tests
overall,
and
opined
this
was
due
to
the
fluctuating
temperatures
increasing
metabolism
without
increasing
food
intake.
Similar
findings
were
produced
using
the
related
Arctic
char
(
Swift,
1975;
as
cited
in
Jensen,
1995;
and
Jobling,
1983)
where
maximum
growth
occurred
at
about
12­
14
°
C
(
at
satiation
rations).
These
growth
studies
may
help
explain
why
Shepard
et
al.
(
1984)
found
that
bull
trout
growth
was
slower
in
the
middle
fork
of
the
Flathead
River,
Montana,
even
though
it
was
warmer
and
more
productive.
The
work
of
McMahon
et
al.
(
2000)
suggest
that
an
average
temperature
of
12­
13
°
C
can
produce
maximal
growth
under
both
severely
restricted
and
satiation
diets,
respectively.
A
constant
laboratory
test
temperature
exposure
of
12­
13
°
C
can
be
correlated
with
7DADMax
summer
stream
temperature
maximums
of
12.93­
13.93
°
C
based
on
correlations
with
the
warmest
one­
week
average
stream
temperature;
and
7DADMax
summer
maximums
of
14­
15
°
C
based
on
correlations
with
the
summer
season­
long
average
stream
temperature.
There
is
reason
to
also
believe
that
in
some
cases
growth
could
be
maximized,
or
not
statistically
different,
at
even
higher
constant
temperatures.
Their
work
also
suggests
that
daily
maximum
temperatures
of
15­
16
°
C
(
7DADMax
14.45­
15.45
°
C)
may
be
producing
thermal
stress
or
excess
metabolic
demands
that
may
hinder
growth
in
juvenile
char
that
are
at
otherwise
healthy
daily
mean
temperatures.
This
is
supported
by
the
findings
of
Bonneau
and
Scarnecchia
(
1996)
who
noted
that
temperatures
above
15­
16
°
C
are
associated
with
increased
metabolic
stress
and
swimming
impairment
in
bull
trout.
The
line
of
evidence
produced
using
the
above
laboratory
studies
suggests
that
streams
with
7DADMax
temperatures
not
exceeding
12.93­
15
°
C
are
likely
to
fully
support
the
growth
of
juvenile
char.

Laboratory
Studies
Examining
Competition
for
Food
Brook
trout
are
a
species
of
char
that
have
been
widely
introduced
in
the
northwest
that
have
often
been
cited
as
eliminating
bull
trout
from
their
native
habitat.
McMahon
et
al.
(
1999),
found
that
the
presence
of
brook
trout
in
sympatry
(
together)
with
bull
trout
resulted
in
significantly
greater
growth
of
brook
trout
and
significantly
lower
growth
for
bull
trout
than
occurred
with
either
species
in
allopatry
(
alone),
especially
at
constant
water
temperatures
equal
to
or
greater
than
12
°
C.
McMahon
et
al.
(
2000)
examined
bull
trout
and
brook
trout
competition
under
two
constant
(
11
and
17
°
C)
temperatures.
At
a
constant
11
°
C
there
were
not
significant
differences
in
the
growth
of
bull
trout
and
brook
trout
in
allopatry
and
sympatry.
However,
at
the
higher
temperature
(
17
°
C)
brook
trout
grew
significantly
more
(
a
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
27
Washington's
Surface
Water
quality
Standards
2.5­
times
greater
growth
rate)
than
bull
trout
in
both
allopatry
and
sympatry.
observations
were
also
made
at
temperatures
of
8
and
16
°
C.
When
in
the
presence
of
brook
trout,
bull
trout
feeding
rates
declined
50%
at
8
°
C
and
64%
at
16
°
C,
whereas
feeding
by
brook
trout
showed
no
change.
The
work
of
McMahon
et
al.
(
1999,
2000)
suggests
that
as
average
water
temperatures
rise
above
12
°
C
bull
trout
may
begin
to
loose
their
competitive
ability
against
brook
trout.
It
also
points
out
the
difficulty
of
trying
to
protect
bull
trout
from
displacement
by
brook
trout
since
the
brook
trout
are
capable
of
significantly
out­
competing
the
bull
trout
even
at
very
low
temperatures
(
8
°
C).

In
trying
to
correlate
the
laboratory
studies
on
competition
to
stream
temperatures,
it
is
important
to
recognize
that
competition
may
be
related
as
much
to
the
absolute
temperature
at
any
point
in
time
as
much
as
a
weekly
or
seasonal
average
temperature
regime.
However,
since
the
effect
of
competition
on
the
health
of
a
species
will
not
be
determined
by
a
portion
of
a
single
day
allowing
for
detrimental
competition,
it
is
still
considered
reasonable
to
correlate
average
stream
temperatures
with
the
constant
laboratory
test
temperature
at
which
competition
was
not
favored
by
water
temperature
(
12
°
C).
A
single
daily
average
and
a
weekly
average
are
used
below
to
represent
the
effects
found
with
constant
laboratory
exposures
and
make
conversions
to
a
7DADMax
temperature
metric.
The
above
cited
work
suggests
that
a
7DADMax
summer
temperature
of
12.62­
12.93
°
C
may
best
describe
a
stream
which
does
not
provide
any
thermal
advantage
in
competition
between
bull
trout
and
brook
trout.
It
is
important
to
remind
the
reader
that
the
laboratory
tests
did
not
include
constant
temperatures
within
the
range
of
12­
16
°
C
making
it
difficult
to
determine
if
a
threshold
exists
within
this
range
above
which
most
of
the
change
in
competitive
ability
occurs.

Summary
of
Laboratory
Studies
The
work
of
McMahon
et
al.
(
1999,
2000)
suggest
that
a
constant
or
average
temperature
of
12
°
C
can
produce
maximal
growth
under
even
severely
restricted
diets.
Their
work
also
suggests
that
as
water
temperatures
rise
above
12
°
C
bull
trout
begin
to
loose
their
ability
to
compete
with
brook
trout.
As
noted
above,
the
research
on
competition
seems
best
viewed
by
comparison
with
a
single
week's
average
temperature,
rather
than
as
a
single
day
of
detrimental
competition.
So,
while
the
range
produced
for
the
laboratory
growth
tests
is
compared
as
either
weekly
average
temperatures
or
season­
long
average
temperatures,
the
laboratory
study
on
brook
trout
competition
is
only
compared
here
as
a
weekly
average
temperature.
After
appropriately
converting
the
constant
laboratory
test
results
to
7DADMax
temperature
metrics,
the
studies
on
growth
and
competition
overlap
each
other.
Growth
should
be
maximized
at
a
7DADmax
of
13.5­
15.5
°
C
and
competition
with
brook
trout
should
not
be
materially
worsened
at
temperatures
below
13.5
°
C.
Page
28
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Conclusion
on
Rearing
The
multiple
lines
of
evidence
created
from
the
wide
variety
of
field
and
laboratory
studies
can
be
brought
together
in
support
of
selecting
a
summer
rearing
temperature
standard.
The
interim
conclusions
from
each
line
of
evidence
is
summarized
below:

Table
4.2.
Juvenile
rearing
of
native
char:

Line
of
Evidence
7DADM
­
Range
(
°
C)
Median
(
°
C)
Field
studies
on
limit
to
where
most
commonly
found
13.45­
14.45
13.95
Filed
studies
on
density,
and
dominance
11.45­
13.45
12.45
Laboratory
Growth
Studies
12.93­
15
13.96
Laboratory
Competition
Studies
12.62­
12.93
12.75
Best
Estimate
of
Threshold
(
averages)
12.61­
13.96
mid.
pt.
13.29
The
range
of
these
independent
lines
of
evidence
is
11.45­
15
°
C
with
a
mean
range
of
12.61­
13.96
°
C
and
an
overall
midpoint
of
13.29
°
C.
This
strongly
suggests
that
streams
having
7­
day
average
daily
maximum
summer
temperatures
not
exceeding
13.29
°
C
are
fully
protective
of
juvenile/
small
bull
trout
rearing.
After
cross
checking
this
conclusion
against
each
independent
line
of
evidence,
no
overriding
factors
of
disagreement
appear
to
exist.
The
slight
conflict
with
the
field
studies
on
density
and
dominance,
and
the
laboratory
competition
studies
do,
however,
suggest
that
rounding
the
estimate
down
to
the
nearest
0.5
°
C
increment,
rather
than
up,
may
be
more
appropriate.
It
is
therefore
concluded
that
13
°
C
as
a
7­
day
average
of
the
daily
maximum
temperatures
will
fully
support
the
life­
stage
of
juvenile/
small
char
rearing.

iv)
Migratory
Adult
and
Sub­
Adult
Char
Populations
As
noted
previously,
fluvial
and
adfluvial
forms
of
char
may
remain
in
the
area
of
their
natal
stream
for
1
to
3
years
and
then
migrate
significant
distances
to
more
productive
waters
for
greater
juvenile
growth
opportunities
(
Pratt,
1992;
Ratliff,
1992;
Riehle
et
al.,
1997;
Fraley
and
Shepard,
1989;
Goetz,
1989).
The
larger
size
of
these
migrants
is
generally
believed
to
allow
them
to
better
compete
for
resources,
and
to
make
use
of
a
larger
prey
base
that
includes
the
juvenile
fish
of
other
species.
This
is
similar
to
the
way
the
ocean
is
used
by
Pacific
salmon
to
enable
them
to
grow
to
significantly
greater
sizes
than
would
be
possible
if
they
were
to
remain
in
freshwater
tributaries.
This
may
be
a
very
important
survival
trait
of
these
migratory
populations,
and
serve
to
free
up
food
resources
in
the
tributary
system
for
juvenile
char.
These
adult
and
non­
spawning
sub­
adult
fish,
may
also
move
out
of
tributary
systems
to
hold
in
lower
main
stem
areas
during
the
winter
to
avoid
unsuitable
winter
conditions
of
ice
and
storm
flows.
In
Washington,
bull
trout
may
migrate
all
the
way
from
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
29
Washington's
Surface
Water
quality
Standards
headwater
streams
to
the
Puget
Sound
to
feed
and
rear.
Relatively
little
is
known
about
the
temperature
preferences
and
requirements
of
these
migratory
fish
which
makes
setting
temperature
criteria
for
them
problematic.

Heimer
(
1965)
examined
the
use
by
Dolly
Varden
of
an
artificial
spawning
channel
on
the
lower
Clark
Fork
River
in
Idaho
below
the
Cabinet
Gorge
Dam.
The
channel
was
constructed
in
an
area
where
cool
spring
water
would
make
the
artificial
channel
cooler
than
the
main
stem
river
during
part
of
the
year.
Temperatures
in
the
majority
of
the
spawning
channel
were
between
8­
11
°
C.
The
author
noted
that
fish
observed
in
the
spawning
area
were
consistently
in
the
areas
of
cooler
water.
As
river
temperatures
declined
in
the
fall
a
portion
of
the
fish
in
the
spawning
area
left
without
spawning.
These
fish
were
presumably
present
because
of
the
cooler
waters,
as
the
main
stem
temperatures
in
the
Clark
Fork
River
did
not
decline
to
below
13.9
°
C
until
after
September
26th.
The
Washington
Water
Power
Company
(
1995)
also
studied
the
distribution
of
fish
in
the
lower
Clark
Fork
River
in
Idaho
and
found
that
while
main
stem
river
temperatures
were
18
°
C
on
September
28th
temperatures
in
the
adjacent
channel
used
by
bull
trout
for
spawning
was
11
°
C.
These
two
studies
when
considered
together
suggest
that
bull
trout
will
actively
avoid
rivers
having
fall
water
temperatures
above
18
°
C.
The
strong
contrasting
temperatures
between
the
river
and
the
channel,
the
combination
of
spawning
and
non­
spawning
fish,
competition
with
other
salmonids
also
using
the
channel
for
refuge,
and
the
lack
of
specific
temperature
metrics
in
association
with
char
movements
all
combine
to
prevent
using
these
studies
to
estimate
a
threshold
response.

Swanberg
(
1997)
found
that
bull
trout
residing
in
the
lower
Blackfoot
River
in
Montana
migrated
out
when
daily
maximum
temperatures
reached
18­
20
°
C,
and
non­
spawning
subadult
fish
began
returning
once
maximum
temperatures
declined
to
12
°
C.
The
few
fish
that
did
not
migrate
were
found
in
association
with
the
confluence
of
a
small
cold
tributary
with
a
daily
maximum
temperature
of
12
°
C.
Elle
and
Thurow
(
1994)
found
that
adfluvial
bull
trout
in
the
Rapid
River
in
Idaho
began
leaving
the
lower
system
in
peak
numbers
seven
out
of
nine
years
as
the
daily
maximum
temperatures
began
to
exceed
10
°
C
and
returned
as
the
temperature
of
the
lower
river
declined
to
10
°
C.
These
fish
moved
to
lower
main
stem
rivers
to
hold
for
the
winter.
Movements
to
avoid
unhealthy
winter
conditions
are
common
in
salmonids,
and
Jakober
et
al.
(
1998)
found
that
bull
trout
and
westslope
cutthroat
trout
in
two
drainages
in
Montana
made
extensive
downstream
movements
as
temperatures
dropped
precipitously
in
the
Fall.
These
movements
were
most
extensive
in
mid­
elevation
streams
where
frequent
freezing
and
thawing
led
to
anchor
ice
formation
and
super
cooling
(<
0
°
C)
of
the
water.

Several
very
import
issues
and
questions
need
to
be
addressed
prior
to
setting
criteria
to
protect
these
adult
and
sub­
adult
fish.
These
include:

 
Is
it
ecologically
appropriate
to
base
temperatures
in
salmon
and
steelhead
strongholds
on
the
temperature
requirements
of
char?
 
How,
and
should,
populations
that
are
in
main
stems
to
rear
be
separated
from
those
that
may
be
leaving
inhospitable
winter
conditions
in
the
tributaries?
Page
30
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
 
Is
rearing
protected
only
by
maintaining
favorable
temperatures
in
these
lower
rivers
year­
round,
or
can
migration
out
of
the
lower
reaches
during
maximum
summer
temperatures
be
considered
a
normal
and
thus
acceptable
natural
pattern?
 
Should
temperatures
considered
protective
of
these
adult
and
sub­
adult
fish
be
based
on
assumptions
of
relative
food
abundance
and
lack
of
competition?

Until
these
questions
are
reasonably
answered,
there
does
not
seem
to
be
sufficient
foundation
in
the
research
to
confidently
support
recommending
a
temperature
threshold
value
in
lower
main
stem
rivers
that
is
colder
than
that
appropriate
for
the
protection
of
salmon
and
steelhead.

v)
Lethality
to
Adults
and
Juveniles
As
shown
above
in
the
discussion
of
juvenile
rearing,
waters
that
fully
protect
the
health
of
native
char
will
have
temperatures
well
below
those
posing
a
threat
of
acute
lethality.
Thus,
the
conclusions
that
follow
are
intended
for
application
in
the
evaluation
of
special
projects.
They
can
also
be
used
as
an
aid
in
assessing
the
relative
safety
for
char
moving
through
main
stem
rivers
predominantly
protected
for
salmon
and
steelhead.

The
only
research
found
directly
testing
one
of
Washington's
native
char
species
(
bull
trout)
was
the
works
of
McMahon
et
al.
(
1998,
1999)
as
published
in
Selong
et
al.
(
2001).
McMahon
et
al.
(
1998,
1999)
conducted
60­
day
lethality
studies
of
juvenile
bull
trout.
In
their
1998
study,
98%
survival
occurred
at
temperatures
between
7.5­
18
°
C.
Mortality
was
21%
at
20
°
C
over
the
60­
day
test
period,
but
was
100%
within
24
hours
at
26
°
C,
within10
days
at
24
°
C,
and
within
38
days
at
22
°
C.
In
the
1999
study,
survival
was
46%
at
21
°
C
and
53%
at
20
°
C,
with
the
time
to
50%
mortality
varying
from
10­
days
at
23
°
C
to
24
days
at
22
°
C.
Using
the
data
from
both
the
1998
and
1999
studies,
the
authors
determined
a
60­
day
UUILT
(
Ultimate
Upper
Incipient
Lethal
Temperature)
of
20.8
°
C
for
juvenile
bull
trout
and
estimated
that
a
7­
day
UILT
(
Upper
Incipient
Lethal
Temperature)
would
be
23.5
°
C
(
Selong
et
al.,
2001).

The
work
of
McMahon
et
al.
(
1998,
1999)
and
Selong
et
al.
(
2001)
strongly
suggest
that
lethality
will
be
prevented
at
constant
temperature
exposures
of
18­
19
°
C
or
less.
This
assertion
comes
from
two
sources
of
information.
The
first
is
that
in
tests
at
a
constant
18
°
C
there
was
at
least
98%
survival
over
a
60­
day
exposure.
The
second
is
derived
by
using
the
60­
day
UUILT
of
20.9
°
C.
After
applying
the
adjustment
factor
recommended
by
the
USEPA
(
Brungs
and
Jones,
1977)
to
change
from
a
temperature
that
kills
50%
of
the
exposed
fish
to
a
temperature
that
is
unlikely
to
kill
any
fish.
The
non­
lethal
temperature
estimate
would
change
to
a
constant
18.9
°
C
(
20.9
°
C
minus
2
°
C).
Constant
temperatures
of
18­
19
°
C
would
be
correlated
to
7DADMax
temperatures
of
21.18­
22.18
°
C
and
1­
day
maximums
(
1DMax)
of
22.13­
23.13
°
C
when
treated
as
if
they
were
based
on
weekly
temperature
exposure.
When
treated
as
if
they
were
based
on
a
season­
long
exposure
(
which
may
be
justified
in
this
case
since
the
laboratory
tests
were
60
days
long)
these
values
change
to
22.64­
23.64
°
C
(
7DADMax)
and
23.60­
24.60
°
C
(
1DMax).
This
line
of
evidence
suggests
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
31
Washington's
Surface
Water
quality
Standards
that
acute
mortality
will
be
prevented
in
acclimated
fish
by
maintaining
7DADMax
temperatures
below
21.18­
23.64
°
C
or
1­
day
maximum
temperatures
below
22.13­
24.6
°
C.

While
a
constant
temperature
exposure
would
generally
be
similar
to
an
average
temperature
in
a
natural
stream,
daily
maximum
temperatures
above
the
lowest
determine
lethal
concentration
(
20.9
°
C
with
a
60­
day
exposure)
will
begin
to
accumulate
lethal
stress.
To
avoid
exposure
to
temperatures
that
create
lethal
stress
altogether
(
would
eventually
cause
lethality
if
exposure
period
is
sufficient)
daily
maximum
temperatures
should
not
exceed
20.9
°
C
(
7DADMax
19.95
°
C).
Since
this
is
not
an
estimate
of
a
threshold
beyond
which
acute
mortality
would
be
expected,
it
should
only
be
viewed
as
a
line
of
evidence
for
establishing
a
lower­
bound
estimate
of
temperatures
that
would
avoid
lethality
and
severe
stress.

Using
the
data
of
Selong
et
al.
(
2001)
we
can
also
estimate
the
general
risk
of
mortality
from
multiple
days
of
exposure
in
the
potentially
lethal
range.
By
assuming
that
temperature
exposure
in
the
lethal
range
is
additive
(
DeHart,
1974,
1975;
Golden,
1978),
and
by
examining
the
potential
lethal
dose
that
occurs
with
each
hour
spent
over
the
lowest
lethal
level
(
20.9
°
C),
the
risk
of
mortality
can
be
reasonably
described.
The
equation
in
Selong
et
al.
(
2001)
giving
the
relationship
between
exposure
temperature
and
time
to
mortality
(
LC50)
can
be
used
to
estimate
the
number
of
hours
that
would
need
to
be
spent
at
each
daily
temperature
increment
(
one­
hour
intervals
used
herein)
above
the
an
ultimately
lethal
temperature
(
20.9
°
C)
to
cause
mortality
(
LC50).
Using
this
approach,
the
number
of
daily
cycles
of
temperature
that
would
occur
before
causing
mortality
can
be
predicted.
Based
on
this
technique,
bull
trout
in
a
stream
with
a
daily
average
temperature
of
20.9
°
C
(
21
°
C)
and
a
diel
range
above
the
daily
mean
of
4.13
°
C
would
be
expected
to
experience
high
mortality
(
LC50)
in
approximately
10
days
or
less.
In
streams
with
mean
temperatures
19
and
18
°
C,
high
mortality
(
LC50)
would
occur
after
54
and
163
days
of
repeat
exposure,
respectively.
This
approach
to
evaluating
the
risk
of
mortality
is
only
useful
in
assessing
relative
risk.
Nevertheless,
it
provides
a
good
support
for
the
position
that
weekly
average
temperatures
of
18­
19
°
C
and
summer
peak
daily
maximum
temperatures
of
22­
23
°
C
(
7DADMax
21.05­
22.05
°
C)
pose
little
risk
of
creating
acutely
lethal
conditions
in
char
populations.

While
having
relatively
sensitive
optimal
temperature
limits,
adult
and
juvenile
char
do
not
appear
unusually
sensitive
to
acute
temperature
limits.
With
acclimation,
juvenile
and
adult
char
are
very
tolerant
of
temperature
extremes
in
a
laboratory
environment
 
capable
of
withstanding
temperatures
of
 
1.2
°
C
(
below
zero)
for
up
to
5
continuous
days
and
having
upper
lethal
temperature
limits
similar
to,
but
towards
the
lower
end,
of
that
for
juvenile
Pacific
salmon
(
Selong
et
al.
2001).
Page
32
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Table
4.3.
Temperature
induced
lethality
of
in
native
char.

Line
of
Evidence
7DADMax
(
°
C)
Midpoint
(
°
C)
Direct
laboratory
observation
of
no
mortality,
and
conversion
of
LC50
to
no
effects
temperature
using
Brungs
and
Jones
(
1977).
21.18­
23.64
22.41
Laboratory
estimate
of
temperature
exerting
lethal
stress
19.95
19.95
Modeling
risk
associated
multiple
day
exposures
to
fluctuating
temperatures
21.05­
22.05
21.55
Best
Estimate
of
Threshold
20.73­
21.88
mid.
pt.
21.31
The
range
of
the
independent
lines
of
evidence
discussed
above
is
19.95­
23.64
°
C
with
a
mean
range
of
20.73­
21.88
°
C
and
an
overall
midpoint
of
21.31
°
C.
This
strongly
suggests
that
not
allowing
single
daily
maximum
temperatures
to
exceed
21.31
°
C
(
7DADMax
of
21.35
°
C)
will
prevent
acute
lethality
in
char
populations.
Based
on
the
above
it
is
concluded
that
an
annual
highest
single­
day
maximum
temperature
not
exceeding
21
°
C
will
prevent
direct
lethality
to
Washington's
juvenile
native
char.

It
is
important
to
point
out,
however,
that
only
juveniles
of
one
stock
of
bull
trout
were
tested.
Juvenile
Dolly
Varden,
other
stocks
of
juvenile
bull
trout
(
Beacham
and
Withler,
1991),
and
adults
of
either
species
could
be
somewhat
more
(
or
less)
sensitive
(
Coutant,
1970;
Becker,
1973;
Bouck
and
Chapman,
1975).
It
would
be
unwise
to
assume
that
temperatures
at
the
upper
end
of
the
range
identified
above
are
of
equal
merit
for
consideration
as
an
acute
criteria
until
more
stocks
have
been
tested.
The
data
cited
below
by
Ugedal
et
al.
(
1994)
lend
further
support
for
being
cautious
about
assuming
the
upper
end
of
the
predicted
range
would
be
fully
protective.

Support
for
the
above
findings
can
be
found
in
research
on
related
species
of
char
(
Arctic
char
and
Arctic
grayling).
The
estimated
LT50
(
lethal
to
50%
of
test
population
within
7
days)
values
for
a
variety
of
char
species
at
acclimations
of
5­
20
°
C
generally
group
between
21.5­
24
°
C.
Lohr
et
al
(
1996)
determine
a
7­
day
LC50
of
23­
25
°
C
for
juvenile
Arctic
Grayling;
Lyytikainen
et
al.
(
1997)
determined
a
14­
day
LC50
of
23­
24
°
C
for
juvenile
Arctic
Char;
and
Baroudy
and
Elliott
(
1994)
determine
a
7­
day
LC50
of
21.5­
21.6
°
C
for
Arctic
char
fry
and
parr.
These
ranges
were
determine
primarily
by
varying
the
acclimation
temperatures.
In
a
modified
CTM
test,
the
authors
also
found
that
fry
and
parr
experienced
10­
minute
LT50
values
of
24.79­
26.57
°
C
at
acclimation
temperatures
from
10
to
20
°
C.
Ugedal
et
al.
(
1994),
in
conducting
a
104­
day
growth
test
using
Arctic
char,
found
that
when
water
temperature
rose
from
an
initial
12­
13
°
C
(
daily
mean)
to
a
brief
20
°
C
(
average
temperature
for
approximately
10
days)
before
falling
again
a
slight
increase
in
mortality
(
approx
5%)
occurred
during
the
period
that
water
temperatures
reached
20
°
C.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
33
Washington's
Surface
Water
quality
Standards
Some
additional
information
from
studies
with
these
related
species
are
worth
noting
here.
At
full
acclimation
to
temperatures
of
15­
18
°
C,
sudden
exposure
to
29
°
C
water
produce
LC50
values
within
2­
4
minutes
in
Arctic
char
(
Lyytikainen
et
al.,
1997).
At
an
acclimation
of
15
°
C
a
test
temperature
of
26
°
C
produced
LC50
results
in
just
44
minutes.
Baroudy
and
Elliott
(
1994)
found
that
the
Arctic
char
alevins
had
7­
day
UILTs
of
18.67,
19.67,
20.83,
and
20.79
°
C
at
acclimation
temperatures
of
5,
10,
15,
and
20
°
C.
Alevins
experience
50%
mortality
within
ten
minutes
at
23.33,
25.09
°
C,
and
25.39
°
C
at
acclimation
temperatures
of
5,
10,
and
15
°
C.

Research
useful
for
estimating
daily
maximum
temperatures
that
will
protect
developing
embryos
and
alevins
was
only
found
for
Arctic
char
(
Baroudy
and
Elliott,
1994).
Since
the
work
of
Baroudy
and
Elliott
was
based
on
a
non­
indigenous
species
of
char,
and
tested
only
the
short­
term
effects
to
the
alevin
life­
stage
it
is
insufficient
for
setting
an
acute
lethality
threshold
for
char
incubation.
It
can
be
used,
however,
to
demonstrate
that
incubation
of
char
is
more
sensitive
then
juvenile
rearing,
and
to
suggest
that
daily
maximum
temperatures
would
likely
need
to
remain
below
16.7
°
C
to
prevent
acute
lethality
to
char
alevins.
This
estimate
was
made
by
subtracting
2
°
C
from
the
LC50
value
determined
at
an
acclimation
temperature
appropriate
for
incubation
(
5
°
C)
to
convert
to
a
temperature
that
should
not
produce
any
short­
term
mortality
(
USEPA,
1977).

b)
Salmon
and
Trout
i)
General
Life
History
Information
Chinook
Salmon
Chinook
salmon
(
Oncorhynchus
tshawytscha)
are
found
in
most
of
the
larger
streams
of
the
Columbia
River
drainage,
and
the
coastal
and
Puget
Sound
drainages.
Juvenile
chinook
salmon
spend
about
a
year
in
fresh
water
before
smolting
and
migrating
to
the
Pacific,
where
they
generally
remain
from
three­
four
years
before
returning
to
their
natal
streams
to
spawn.
Adults
begin
the
ascent
of
coastal
streams
in
late
May
and
early
June
and
principally
spawn
through
September.
Chinook
salmon
are
primarily
divided
into
two
stock
types,
based
on
the
season
they
initially
return
to
fresh
waters.
These
are
spring­
run
and
fall­
run
stocks,
but
in
some
areas
a
third,
summer­
run,
stock
may
be
identified.
Spring
run
fish
begin
entering
freshwaters
from
May
through
June,
and
typically
hold
in
deep
pools
and
at
the
mouths
of
cool
tributaries
until
the
fall
rains
begin
and
stream
levels
go
up
sufficiently
to
allow
them
to
reach
their
upriver
spawning
sites.
These
spring
run
fish
tend
to
spawn
higher
in
the
watersheds
than
the
fall
run
stocks.
Fall
chinook
migrate
up
the
streams
in
August
and
September
and
spawn
as
soon
as
the
spawning
grounds
are
reached
when
water
temperatures
are
between
5.6­
14.4
°
C.
Eggs
hatch
in
about
two
months
and
the
young
remain
in
the
gravel
for
two­
three
weeks
prior
to
emerging.
Juveniles
remain
in
freshwater
from
a
few
days
to
three
years.
Usually,
juvenile
fall
chinook
feed
for
a
short
time
and
then
migrate
to
the
ocean,
whereas
most
juvenile
spring
chinook
remain
in
the
stream
for
one
year
before
migrating
(
Wydoski
and
Whitney,
1979).
In
Washington,
96
of
the
98
chinook
stocks
Page
34
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
assessed
began
spawning
after
August
1,
and
96
of
98
had
midpoints
in
their
spawning
periods
that
were
after
August
23.
After
emerging
from
the
gravel,
chinook
fry
may
remain
in
freshwater
from
just
a
few
days
to
up
to
3
years.
Usually,
fall
chinook
feed
for
a
short
time
and
then
migrate
to
the
ocean,
whereas
most
juvenile
spring
chinook
remain
in
the
stream
for
one
year
before
migrating
to
the
ocean
(
Wydoski
and
Whitney,
1979).

Coho
Salmon
Coho
salmon
occur
all
along
the
Pacific
coast.
While
fourth
in
abundance
of
the
Pacific
salmon,
coho
provide
the
dominant
harvest
of
sport
fisherman
(
Wydoski
and
Whitney,
1979).
In
Washington,
spawning
adults
are
found
in
most
streams
of
the
upper
and
lower
Columbia
River
drainage,
and
in
the
coastal
and
Puget
Sound
drainages.
Coho
spend
their
first
(
one
to
two)
years
in
freshwater
before
becoming
smolts
and
migrating
to
the
ocean.
Spawning
generally
occurs
from
September
through
December;
although,
a
late
run
of
large
fish
is
known
to
spawn
in
January
in
the
Satsop
River.
Like
all
Pacific
salmon,
adult
coho
die
after
spawning.
Young
hatch
in
6­
8
weeks
depending
on
water
temperature
and
emerge
in
about
two­
three
weeks.
Fry
congregate
in
schools
in
the
pools
of
the
stream.
Ruggels
(
1966;
as
cited
by
Chapman
and
Bjornn,
1969)
found
that
coho
did
not
use
substrate
as
cover
in
the
winter,
and
Hartman
(
1965)
found
that
coho
tended
to
lie
near
or
on
pool
bottoms
in
aggregations.

Chum
Salmon
Chum
salmon
are
found
in
streams
of
the
Puget
Sound
and
coastal
drainages,
and
up
the
Columbia
to
the
Wind
River
(
upstream
from
the
Bonneville
Dam).
Spawning
often
occurs
just
at
the
head
of
tidewater.
Upon
emergence
from
the
stream
gravel,
chum
salmon
fry
begin
to
migrate
to
the
ocean.
Juveniles
migrate
to
the
ocean
between
March
and
June
where
they
spend
six
months
to
four
years.
In
Washington,
chum
salmon
spawn
primarily
from
October
to
December.
Redds
occur
in
medium
or
fine
gravel
in
the
stream
riffles.
Most
of
the
high
mortality
from
the
fertilized
egg
to
early
fry
state
(
70
to
over
90%)
is
reported
to
occur
during
the
embryonic
stage
due
to
suffocation
from
silt.
Eggs
hatch
in
two
weeks
to
four
and
one­
half
months
depending
upon
the
temperature.
Newly
hatched
fry
(
alevins)
absorb
their
yolk
sac
in
30­
50
days,
again
depending
on
temperature.
Fry
are
usually
in
fresh
water
for
only
a
few
days
after
emerging
from
the
gravel.
They
migrate
downstream
at
night
from
April
through
June
(
Wydoski
and
Whitney,
1979).
Little
information
was
found
on
which
to
base
a
recommendation
for
juvenile
rearing
temperature
limits
for
chum
salmon;
however,
this
is
not
a
major
concern.
Considering
that
juvenile
chum
may
only
spend
a
few
days
in
freshwaters,
the
temperature
threshold
established
for
the
pre­
emergent
stages
of
development
should
result
in
cool
waters
remaining
during
the
out­
migration
period.
Thus
even
without
a
specific
juvenile
rearing
criterion
the
freshwater
portion
of
the
juvenile
lifestage
will
be
fully
protected.
If
unique
situations
appear
to
demand
that
a
separate
rearing
standard
be
applied,
the
temperature
threshold
established
to
protect
other
Pacific
salmon
smolts
during
sea­
water
adaptation
should
be
applied.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
35
Washington's
Surface
Water
quality
Standards
Pink
Salmon
Pink
salmon
are
the
most
abundant
of
the
Pacific
Salmon.
Pink
salmon
ascend
rivers
along
the
Pacific
coast;
and
in
Washington,
the
Stillaguamish,
Skagit,
Snohomish,
Puyallup,
and
Nooksack
are
the
top
producing
watersheds.
Pink
salmon
are
similar
to
chum
salmon
in
spending
only
a
brief
time
in
fresh
water
as
juveniles
and
adults.
Spawning
in
Washington
occurs
chiefly
during
August
and
September,
usually
near
the
mouths
of
streams,
but
sometimes
spawning
occurs
far
up
into
large
rivers,
such
as
the
Skagit.
Females
usually
dig
redds
in
a
riffle
area
with
small
to
medium
size
gravel,
but
they
occasionally
spawn
in
the
tail
section
of
pools.
In
Alaska,
a
large
percentage
of
pink
salmon
spawn
in
the
intertidal
areas.
Eggs
usually
hatch
in
thee­
five
months,
depending
upon
water
temperature.
The
alevins
and
fry
remain
in
the
gravel
for
as
long
as
several
months
at
low
temperatures
in
northern
waters
before
emerging
(
Wydoski
and
Whitney,
1979).
Washington
state
appears
to
be
at
the
southern
end
of
the
range
for
streams
that
support
consistently
exploitable
spawning
runs
of
pink
salmon.
Pink
salmon
have
a
two­
year
life
cycle
which
is
so
invariable
that
fish
running
in
odd­
numbered
calendar
years
are
effectively
genetically
isolated
from
those
that
run
in
even
years
(
Bonar
et
al.,
1989).

Sockeye
Salmon
In
Washington,
populations
of
sockeye
travel
up
the
Columbia
River
to
the
headwaters
of
the
Salmon
River
in
Central
Idaho.
Populations
also
use
Quinalt
Lake
on
the
Olympic
Peninsula,
Baker
Lake,
lakes
Washington
and
Sammamish
in
the
Puget
Sound
drainage,
and
Osoyoos
Lake
and
Lake
Wenatchee
east
of
the
Cascade
Mountains.
Sockeye
differ
from
the
other
species
of
salmon
because
they
require
a
lake
environment
for
part
of
their
life
cycle.
Although
spawning
typically
occurs
in
the
gravel
of
streams
some
may
spawn
along
lake
shores
in
areas
where
ground
water
percolates
through
the
gravel.
Eggs
hatch
in
six­
nine
weeks.
The
young
remain
in
the
gravel
for
another
two­
three
weeks.
Newly
emerged
fry
move
from
the
spawning
stream
into
the
lake
associated
with
the
stream.
Young
sockeye
live
in
the
pelagic
zone
of
the
lake
for
one
to
two
years
years
before
migrating
to
the
sea
where
they
will
remain
for
two
years
before
returning
to
freshwaters
to
spawn
(
Wydoski
and
Whitney,
1979).

Steelhead
Steelhead
is
the
popular
name
given
to
the
anadromous
form
of
O.
mykiss.
The
anadromous
form
is
found
in
coastal
rainbow
and
interior
redband
trout
groups
and
occurs
from
southern
California
to
Alaska
(
Behnke,
1992).
Steelhead
populations
can
be
broadly
divided
into
spring­,
summer­,
fall­
and
winter­
run
stocks,
depending
upon
the
time
the
fish
re­
enters
freshwater.
Spring
and
summer­
run
fish
enter
fresh
waters
typically
from
May
through
August
and
move
upstream
to
hold
over
until
the
following
spring
to
spawn.
Fall
runs
typically
enter
from
September
through
November
and
spawn
in
the
spring.
Winter­
run
(
December­
March)
steelhead
may
spawn
soon
after
entering
fresh
waters.
In
general,
summer­
run
steelhead
spawn
further
upstream
in
the
watershed
than
the
fall­
or
winter­
run
fish.
With
a
spring
spawning
stock,
protection
is
achieved
by
not
encouraging
spring
water
temperatures
to
warm
above
fully
protective
levels
during
the
spawning
and
incubation
Page
36
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
periods.
Juvenile
steelhead
may
remain
in
freshwaters
for
one
to
seven
years
before
emigrating
to
sea,
depending
upon
the
stream
temperature
and
the
latitude,
although,
most
stay
in
fresh
water
for
two
years
(
Wydoski
and
Whitney,
1979;
Mullan
et
al.,
1992).
Mullan
et
al.
(
1992)
note
that
summer­
run
stocks
tend
to
remain
longer
in
freshwater
than
winter­
run
steelhead,
presumably
because
the
colder
temperatures
of
the
headwaters
streams
preferred
by
summer­
run
steelhead
retards
their
growth
rate.

Rainbow
Trout
The
name
rainbow
trout
is
commonly
applied
to
represent
any
or
all
of
the
members
of
Oncorhynchus
mykiss.
However,
O.
mykiss
in
Washington
can
be
further
divided
into
resident
coastal
rainbow
trout;
resident
interior
redband
trout;
and
steelhead,
the
anadromous
form
of
the
coastal
rainbow
and
redband
trout.
Coastal
rainbow
trout
populations
extend
from
Alaska
to
Mexico,
and
non­
anadromous,
or
resident,
populations
occur
throughout
the
entire
range
(
Behnke,
1992).
Redband
trout
are
found
east
of
the
Cascades
and
in
the
Columbia
River
basin,
and
Behnke
suggests
that
that
the
native
redband
trout
of
each
basin
has
its
own
peculiarities
and
could
probably
be
separated
into
several
new
subspecies.
Behnke
also
notes
that
in
the
desert
basins
of
the
western
states,
the
redband
trout
has
evolved
adaptations
to
live
in
extremely
harsh
environments
characterized
by
great
extremes
in
water
temperature
and
flow.

Cutthroat
Trout
In
Washington,
native
cutthroat
trout
can
be
separated
into
coastal
cutthroat
(
Oncorhynchus
clarki
clarki)
and
west­
slope
cutthroat
trout
(
Oncorhynchus
clarki
lewisi).
Coastal
cutthroat
exhibit
anadromous,
potamodromous
stream
dwelling,
potamodromous
lake­
dwelling,
and
headwater
stream­
resident
life­
history
forms;
while
west­
slope
cutthroat
exclude
anadromous
populations
(
Trotter,
1998).
Behnke
(
1992)
suggests
that
cutthroat
trout
enjoy
a
selective
advantage
over
non­
native
trout
in
many
high­
altitude
headwaters;
presumably
because
they
function
better
in
colder
waters.
He
notes
that
native
cutthroat
are
quickly
eliminated
from
waters
where
non­
native
trout
become
established.
Cutthroat
are
displaced
from
preferred
habitat
in
the
presence
of
rainbow
trout
and
coho
salmon.
The
aggressive
interaction
of
these
other
species
may
be
heightened
by
warmer
water
temperatures
and
thus
the
displacement
of
cutthroat
is
lessened
by
cooler
water
temperatures
(
Pauley
et
al.,
1989;
Trotter,
1989;
and
Mullan
et
al.,
1992).

Coastal
cutthroat
trout
(
Oncorhynchus
clarki
clarki)
occur
all
along
the
Pacific
coast
from
southern
California
to
Alaska.
Though
rarely
found
more
than
16
km
inland,
it
is
considered
the
most
abundant
of
the
cutthroat
subspecies
(
Trotter,
1998).
Sea­
run
cutthroat
live
to
a
maximum
age
of
about
ten
years,
as
compared
to
resident
forms
which
may
live
for
only
4
to
5
years
(
Behnke,
1992).

In
Washington,
sea­
run
cutthroat
may
re­
enter
fresh
water
for
spawning
anytime
from
July
through
March
(
Pauley
et
al.,
1989).
Early­
entering
stocks
in
Puget
Sound
and
Hood
Canal
typically
occur
from
July
through
November
with
the
peak
in
September
and
October.
Lateentering
migration
peaks
in
December
and
January
but
continues
through
March
(
Trotter,
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
37
Washington's
Surface
Water
quality
Standards
1989;
Pauley
et
al.,
1989).
In
southeast
Alaska,
Jones
(
1977;
as
cited
in
Pauley
et
al.,
1989)
found
that
migration
began
at
10­
12
°
C
and
peaked
at
9­
10
°
C.

Pauley
et
al.
(
1989)
suggests
that
prior
to
smolting
and
entering
saltwater,
sea
run
cutthroat
juveniles
may
migrate
up
and
downstream
several
times.
They
further
suggest
that
in
Washington
and
Oregon,
downstream
movement
is
reported
to
take
place
from
March
to
June,
but
peaks
in
mid­
May.
They
note
that
in
southeast
Alaska,
juveniles
have
been
reported
to
experience
a
peak
of
out­
migration
when
water
temperatures
are
between
4­
6
°
C.
Cutthroat
parr
may
migrate
to
sea
from
ages
one
to
six,
but
age
two
to
four
may
be
most
common
(
Trotter,
1989;
Pauley
et
al.,
1989).

Stream
resident
forms
of
coastal
cutthroat
spend
most
or
all
of
their
life
in
or
very
near
their
natal
streams
(
Trotter,
1989).
In
spring
when
water
temperatures
reach
5­
6
°
C,
mature
resident
cutthroat
move
onto
the
spawning
gravel.
Trotter
(
1989)
suggests
that
potamodromous
forms
of
coastal
cutthroat
do
not
move
into
spawning
tributaries
until
very
late
winter
or
spring
rather
than
in
autumn
to
early
winter
as
for
anadromous
fish.
Similar
to
stream
resident
forms,
they
are
reported
to
begin
spawning
as
the
water
temperature
increases
to
5­
6
°
C,
which
may
occur
from
February
through
June.
Lake­
dwelling
potamadromous
coastal
cutthroat
first
spawn
at
age
3
or
4,
and
then
spawn
almost
every
year
thereafter
for
the
remainder
of
their
lives
in
the
inlet
and
outlet
streams
of
the
lake
(
Trotter,
1989).
Few
sea­
run
cutthroat
trout
sexually
mature
before
age
4,
and
not
all
returning
fish
spawn
their
first
year
back
in
freshwater
(
Trotter,
1989).
Fish
that
do
spawn,
may
return
(
39­
41%)
to
spawn
a
second
time,
and
some
(
12%)
may
return
for
a
third
spawning
(
Trotter,
1989;
Pauley
et
al.,
1989).

West­
slope
cutthroat
trout,
the
interior
form,
have
been
found
to
spawn
in
April
and
early
May
with
a
peak
around
mid­
April
in
Montana
(
Fraley
et
al.,
1981).
Trotter
(
1999)
found
an
introduced
population
of
west­
slope
cutthroat
initiating
spawning
around
June
1
at
a
temperature
of
7
°
C
in
the
Tolt
River
in
Washington,
and
a
natural
population
initiating
spawning
on
June
29
at
a
temperature
of
11
°
C
in
a
stream
in
the
upper
Yakima
River
basin
in
Washington.
Newly
emerged
fry
move
to
low
velocity
stream
margins,
backwaters,
and
side
channels
adjacent
to
the
main­
channel
pools
and
riffles
(
Pauley
et
al.,
1989;
Trotter,
1989).
Trotter
(
1989)
suggests
that
these
young
fish
may
move
downstream
to
the
main
stem
as
early
as
the
winter
of
their
first
year,
but
more
generally
later
in
the
spring.
With
the
onset
of
winter
freshets,
the
fish
may
again
move
back
into
the
tributaries.
Older
fish
may
migrate
to
sea
where
they
will
remain
for
two
to
five
months
concentrating
in
bays,
and
estuaries
along
the
coast
gaining
weight
before
returning
to
fresh
waters.
It
is
considered
unusual
for
cutthroat
to
over­
winter
in
salt
waters
(
Pauley
et
al.,
1989).
Fry
of
potamadromous
forms
may
spend
the
first
1­
3
years
in
the
tributaries
before
commencing
a
lake­
ward
migration.
These
potamadromous
fish
may
over­
winter
in
the
lower
main
stem
of
the
river
while
anadromous
fish
are
in
the
marine
waters
(
Trotter,
1989).
Pauley
et
al.
(
1989)
cite
research
showing
that
while
in
freshwaters,
cutthroat
adults
are
associated
with
the
deeper
pools
and
slower
velocity
waters,
while
the
fry
are
found
in
shallower,
faster
areas.
Page
38
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
ii)
Spawning
Requirements
Field
and
General
Observations
on
Spawning
Temperatures:

The
authors
that
have
provided
the
following
general
observations
of
spawning
temperatures
rarely
provided
the
associated
temperature
metric.
Thus
it
is
not
clear
most
of
the
time
whether
or
not
they
were
referring
to
a
range
of
daily
average
temperatures
or
a
range
of
daily
maximum
temperatures.
Other
confounding
factors
include
that
the
authors
do
not
always
make
it
clear
that
they
are
separating
pairing
and
redd
construction
activity
with
actual
spawning,
and
generally
do
not
provide
a
copy
of
their
methodology
to
use
in
determining
if
the
temperatures
were
recorded
at
the
site
of
spawning
at
the
time
spawning
was
taking
place.
For
these
reasons,
the
following
information
on
temperatures
at
which
spawning
commonly
occurs
should
be
used
cautiously.

Chinook
Salmon
The
technical
literature
reviewed
for
this
paper
notes
a
wide
range
of
temperatures
associated
with
the
spawning
of
chinook
salmon
(
5.6­
17.7
°
C)
(
Seymour,
1956).
The
majority
of
these
temperature
observations,
however,
cite
maximum
temperatures
below
14.5
°
C
[
Bell,
1986
(
literature
Summary);
Piper
et
al.,
1982;
Wydoski
and
Whitney,
1979].
In
the
Hanford
reach
and
in
the
Snake
River
in
Washington,
redd
construction
began
as
weekly
mean
temperatures
dropped
to
15.9
°
C
and
averaged
13.6
°
C
during
the
weeks
of
spawning
initiation
from
1993­
1995
(
Groves
and
Chandler,
1999).

Coho
Salmon
Coho
salmon
(
Oncorhynchus
kisutch)
tend
to
spawn
later
and
at
lower
water
temperatures
than
the
other
Pacific
salmon.
In
Washington,
79
of
the
82
stocks
of
coho
salmon
assessed
begin
spawning
after
September
22,
and
80
of
82
have
midpoints
in
their
spawning
periods
that
occur
after
November
1.
Spawning
activity
in
coho
may
typically
occur
in
the
range
of
4.4­
13.3
°
C
[
Bell,
1986
(
literature
summary);
Piper
et
al.,
1982;
Chambers,
1956,
as
cited
in
Andrew
and
Green,
1960;
Gribanov,
1948,
and
Briggs,
1953,
as
cited
in
Sandercock,
1991].

Chum
Salmon
Chum
salmon
are
reported
to
spawn
between
1­
12.8
°
C,
with
a
range
of
7­
10.5
°
C
being
most
consistently
identified
[
Beacham
and
Murray,
1986;
Bell,
1986
(
literature
summary)].
In
Washington,
65
of
the
68
stocks
of
chum
salmon
(
Oncorhynchus
keta)
assessed
begin
spawning
after
September
1,
and
65
of
the
68
stocks
have
midpoints
in
their
spawning
ranges
that
occur
after
October
8.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
39
Washington's
Surface
Water
quality
Standards
Pink
Salmon
In
Washington,
10
of
the
12
pink
salmon
(
Oncorhynchus
gorbuscha)
stocks
assessed
begin
spawning
after
September
1,
and
10
of
12
have
midpoints
in
their
spawning
periods
which
occur
after
September
22.
Spawning
typically
occurs
between
7.2­
14
°
C
(
Bonar
et
al.,
1989),
and
may
be
optimally
supported
at
temperatures
from
7­
12.8
°
C
[
Sheridan,
1962;
Bell,
1986
(
literature
summary)].
This
potential
optimum
range
coincides
well
with
an
identified
peak
in
spawning
activity
at
10
°
C
identified
by
Sheridan
(
1962).

Sockeye
Salmon
In
Washington,
8
of
the
8
stocks
of
sockeye
salmon
(
Oncorhynchus
nerka)
assessed
begin
spawning
after
September
7,
and
8
of
8
have
midpoints
in
their
spawning
ranges
that
occur
after
October
3.
Temperatures
to
support
spawning
activity
in
sockeye
salmon
have
been
identified
as
ranging
from
7.2­
12.2
°
C
(
Piper
et
al,
1982),
with
a
preferred
range
between
10.6­
12.2
°
C
(
Bell,
1986,
literature
summary).
Chambers
(
1956;
as
cited
in
Andrew
and
Geen,
1960)
reported
that
sockeye
spawn
on
the
falling
portion
of
the
cycle
at
12.8
to
8.3
°
C.
Andrew
and
Geen
(
1960)
reported
that
spawning
temperatures
of
sockeye
in
the
Fraser
River
range
from
7.2
to
12.8
°
C
with
a
peak
at
about
10
°
C.

Steelhead
Trout
In
Washington,
101
of
the
105
spring
spawning
steelhead
stocks
assessed
ended
spawning
before
July
1
(
91
of
the
105
ended
prior
to
June
15),
and
101
of
105
have
midpoints
in
their
spawning
periods
that
occur
before
May
7.
Spawning
occurs
primarily
in
the
spring
as
water
temperatures
are
rising.
Bell
(
1986,
literature
summary)
noted
that
spawning
has
been
observed
at
temperatures
ranging
from
3.9­
21.1
°
C.
Hunter
(
1973;
as
cited
in
Swift,
1976)
noted
the
preferred
temperatures
for
spawning
range
from
4.4­
12.8
°
C.

Rainbow
Trout
Both
the
coastal
rainbow
and
the
redband
trout
spawn
in
the
spring,
stimulated
by
rising
water
temperatures.
Behnke
(
1992)
suggests
that
along
the
Pacific
coast
a
water
temperature
of
about
3­
6
°
C
may
initiate
spawning
activity,
but
that
actual
spawning
does
not
occur
until
the
temperatures
reach
6­
9
°
C.
While
this
spawning
activity
would
typically
occur
from
late
December
through
April,
in
some
very
cold
headwater
streams
local
temperatures
may
delay
spawning
until
July
or
August
for
some
stocks.
Bell
(
1986,
literature
summary)
set
the
range
for
spawning
at
2.2­
18.9
°
C;
and
Piper
et
al.
(
1982)
concluded
that
rainbow
trout
spawning
should
occur
between
10­
12.8
°
C.

Cutthroat
Trout
Bell
(
1986,
literature
review)
has
suggested
that
the
spawning
range
is
6.1­
17.2
°
C.
Fraely
et
al.
(
1981)
evaluated
the
spawning
habitat
of
west­
slope
cutthroat
trout
in
Montana.
They
found
that
the
better
spawning
streams
had
maximum
temperatures
of
11­
13
°
C
and
mean
Page
40
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
monthly
maximum
temperatures
that
exceeded
10
°
C
only
during
July
and
August.
The
poorer
quality
spawning
streams
had
higher
maximum
summer
temperatures,
with
two
having
18
°
C
and
19
°
C
as
the
average
maximum
temperatures
during
July.

Cutthroat
trout
spawn
in
the
spring,
stimulated
by
a
rising
water
temperature.
While
temperatures
of
3­
6
°
C
may
initiate
spawning
activity
by
coastal
cutthroat
from
late
December
through
April,
actual
spawning
may
not
occur
until
temperatures
reach
6­
9
°
C.
Spawning
may
extend
from
December
through
May
in
Washington,
but
peaks
in
February
(
Trotter,
1989;
Wydoski
and
Whitney,
1979;
Pauley
et
al.,
1989;
Behnke,
1992).
Sea­
run
cutthroat
tend
to
spawn
close
to
deep
pools
(
Pauley
et
al.,
1989)
in
low
gradient
areas
of
small
tributaries
(
Wydoski
and
Whitney,
1979;
Trotter,
1989)
where
they
may
avoid
competition
for
rearing
area
with
steelhead
and
coho
salmon.
Pauley
et
al.
(
1989)
suggest
that
searun
cutthroat
travel
farther
upstream
to
spawn
than
either
steelhead
or
coho
salmon
where
they
rear
sympatric
with
resident
cutthroat
populations.
While
cutthroat
home
very
precisely
to
their
natal
streams
to
spawn,
immature
fish
may
migrate
from
marine
waters
to
non­
natal
areas
to
feed
(
Pauley
et
al.,
1989).

Summary
of
Reported
Spawning
Temperatures:
While
both
salmon
and
trout
have
been
occasionally
cited
has
having
a
spawning
range
that
extends
as
high
as
17.2­
17.8
°
C
(
7DADMax
16.35­
16.95
°
C),
these
are
uncommon
citations
and
are
sometimes
described
as
poorer
quality
spawning
streams.
Generally,
maximum
temperatures
below
12.8­
14.5
°
C
(
7DADMax
11.95­
13.65
°
C)
appear
to
most
consistently
define
the
thermal
conditions
under
which
spawning
will
be
initiated
in
the
fall
spawning
Pacific
salmon.
Spring
spawning
species
such
as
Steelhead
trout
and
cutthroat
trout
commonly
begin
spawning
as
temperature
warm
above
4­
6
°
C.
Most
authors
reviewed
place
the
late
spring
spawning
temperature
range,
or
define
quality
spawning
stream
summer
maximum
temperatures,
as
being
12.8­
13
°
C
(
7DADMax
11.95­
12.15
°
C).
Thus
for
all
salmonids,
temperatures
below
12.8­
14.5
°
C
(
7DADMax
11.95­
13.65
°
C)
are
generally
viewed
as
the
upper
limit
for
spawning.
For
the
purpose
of
this
analysis,
the
studies
are
being
treated
as
if
they
were
representing
the
daily
maximum
temperature,
unless
the
author
suggested
otherwise.
This
interpretation
by
itself
could
be
viewed
as
being
biased
towards
the
protective
side.
However,
it
is
reasonable
in
light
of
the
way
temperature
ranges
are
typically
used
in
the
literature
(
as
extreme
point
estimates),
and
in
light
of
the
fact
that
the
temperature
ranges
are
not
typically
related
to
an
estimate
of
reproductive
success.

Prespawning
Effects
of
Temperature:

The
effect
of
temperature
exposure
prior
to
actual
spawning
can
effect
the
development
of
viable
offspring
and
needs
to
be
considered
in
setting
a
temperature
standard.
It
has
been
found
that
temperature
can
effect
both
the
health
of
the
spawners
and
their
potential
reproductive
success
prior
to
the
act
of
spawning.
One
of
the
ways
in
which
temperature
affects
the
health
of
spawners
is
by
increasing
the
risk
of
mortality
from
warm
water
diseases
(
Schreck
et
al.,
1994;
Bumgarner
et
al.,
1997;
and
ODFW,
1992)
prior
to
spawning
(
disease
is
discussed
separately
in
this
paper).
Another
way
that
warm
water
affects
the
success
of
spawners
is
through
its
effect
on
the
health
of
the
unfertilized
eggs
as
well
as
the
maturation
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
41
Washington's
Surface
Water
quality
Standards
timing
of
adult
salmon.
This
information
is
important
to
setting
temperatures
in
both
spawning
streams
and
along
the
final
migratory
route
taken
by
adult
spawners.

Chinook
Salmon
Several
researchers
have
examined
the
effects
of
holding
mature
adult
chinook
prior
to
spawning
at
warm
temperatures.
Holding
mature
adults
at
warm
temperatures
has
been
found
to
the
reduce
the
survival
of
eggs
(
Hinze,
1959;
as
cited
in
CDWR,
1988).
Rice
(
1960)
found
that
holding
broodstock
at
temperatures
above
15.6
°
C
reduced
survival
of
eggs
to
the
eyed
stage
by
12.7%
as
compared
to
holding
broodstock
at
8.3­
15.6
°
C
(
mean
11.95
°
C).
Adult
immigrants
held
at
temperatures
greater
than
15.6
or
less
than
3.3
°
C
were
also
found
to
produce
eggs
that
are
less
viable
in
a
study
by
Hinze,
Culver,
and
Rice
(
1956;
as
cited
in
CDWR,
1988).
The
greatest
survival
(
95%)
was
from
adults
taken
at
temperatures
in
the
range
of
11.7­
12.2
°
C.
Berman
and
Quinn
(
1989)
cite
a
personal
communication
with
the
manager
of
the
Kalama
State
Fish
Hatchery
as
finding
egg
moralities
of
50%
or
more
from
adults
held
in
river
waters
fluctuating
from
14.4­
19.4
°
C
(
mean
16.9
°
C).
The
current
supervisor
of
the
Kalama
Falls
Hatchery,
Ron
Castaneda,
notes
that
they
still
attribute
some
increased
losses
to
holding
temperature
around
15.6­
17.8
°
C;
although,
they
have
not
had
conditions
of
mortality
as
high
as
50%.
Similarly,
Marine
(
1992;
as
cited
in
USEPA,
2001)
noted
that
conventional
hatchery
practice
is
to
consider
chinook
broodstock
stressed
at
temperatures
above
15
°
C,
and
noted
that
survival
declines
dramatically
in
holding
ponds
when
temperatures
exceed
17
°
C.
M.
Everson
with
the
Oregon
Department
of
Fish
and
Wildlife
is
cited
in
a
personal
communication
by
Marine
(
1992;
as
cited
in
USEPA,
2001)
as
acknowledging
that
wild
spring
chinook
in
the
lower
Rouge
River
exhibited
high
prespawning
mortality
in
1992
when
water
temperatures
ranged
from
18­
21
°
C.
The
information
cited
above
suggests
that
average
temperature
exposures
of
15.6­
17
°
C
are
associated
with
reduction
in
reproductive
success.
Average
temperatures
of
approximately
12
°
C
were
associated
with
high
survival
rates.

Sockeye
Salmon
Andrew
and
Geen
(
1960)
noted
that
average
daily
temperatures
in
excess
of
12.8
°
C
during
the
spawning
period
appear
to
increase
the
numbers
of
female
sockeye
salmon
that
die
unspawned.
In
one
year
when
the
average
daily
water
temperatures
at
the
peak
of
spawning
ranged
from
12.8­
15.6
°
C
(
mean
14.2
°
C),
spawning
success
was
only
45%.

Steelhead
and
Rainbow
Trout
Smith
et
al.
(
1983)
and
Piper
et
al.
(
1982)
cite
work
demonstrating
that
adult
brood­
fish
should
be
held
at
temperatures
below
12.2­
13.3
°
C
prior
to
spawning
to
produce
good
quality
eggs,
while
holding
temperatures
above
13
°
C
have
been
found
to
reduce
invivo
postovulatory
egg
survival
(
Flett
et
al.,
1996,
and
Billard
and
Gillet,
1981;
as
cited
in
Billard,
1985).
Temperatures
of
18
°
C
or
higher
have
been
found
to
reduce
the
volume
of
male
sperm,
and
a
temperature
of
20
°
C
has
found
to
cause
a
drop
in
egg
fertility
invivo
to
5%
after
four
and
one­
half
days
(
Billard
and
Breton,
1977).
But
at
10
°
C,
fertility
of
the
eggs
held
in
Page
42
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
the
hen
trout
remained
high.
Steroid
biosynthesis
has
been
noted
to
be
suppressed
in
rainbow
trout
testes
at
17
°
C
(
Manning
and
Kime,
1985;
as
cited
in
Macdonald
et
al.,
2000).
Saki
et
al.
(
1975;
as
cited
in
De
Gaudemar
and
Beal,
1998)
found
that
embryonic
and
post
hatching
survival
in
O.
mykiss
decreased
significantly
if
they
remained
ripe
in
the
body
cavity
for
more
than
5­
7
days
after
ovulation,
and
fertility
could
approach
zero
after
two
weeks
(
Stein
and
Hochs,
1979;
as
cited
in
De
Gaudemar
and
Beal,
1998).
To
avoid
invitro
damage
to
the
eggs
of
steelhead
and
rainbow
trout
it
appears
that
constant
temperatures
should
be
maintained
below
12.2­
13.3
°
C.

Summary
on
Prespawning
Effects:
A
consistent
finding
among
the
studies
reviewed
was
that
average
temperatures
greater
than
12­
13
°
C
(
7DADMax
of
13.48­
14.48
°
C
when
study
results
are
treated
as
daily
average,
and
13.96­
14.96
°
C
when
treated
as
a
weekly
average)
(
conversion
assumes
a
diel
variation
of
2­
4
°
C)
pose
a
real
risk
of
reducing
spawner
viability,
and
that
high
losses
can
occur
when
average
temperatures
exceed
14
°
C
(
7DADMax
15.48­
15.96
°
C)
in
the
weeks
immediately
prior
to
spawning.
An
important
consideration
is
that
the
viability
of
the
eggs
in
ripe
fish
will
decline
both
with
time
held
in
utero
and
with
increasing
temperatures
above
12
°
C,
thus
temperatures
should
be
suitable
for
spawning
within
a
few
days
of
when
ripe
adults
arrive
on
the
spawning
grounds
to
avoid
losses.
It
seems
most
appropriate
to
treat
the
laboratory
test
results
that
were
based
on
constant
test
exposures
as
daily
average
and
weekly
average
temperatures
(
both
are
used
to
create
the
likely
range
within
which
the
best
estimate
would
occur)
when
converting
to
the
standardized
7DADMax
metric
used
in
this
analysis.
This
is
because
the
period
of
time
that
exposure
would
be
effecting
ripe
eggs
and
adult
fish
in
spawning
condition
would
be
around
one
to
four
weeks.
Thus
to
avoid
prespawning
losses
of
eggs
the
highest
7DADMax
temperature
should
not
exceed
13.48­
14.96
°
C.
Given
that
the
effect
of
concern
is
a
lethal
endpoint
(
rather
then
sublethal
such
as
a
reduction
of
growth)
it
would
be
most
advisable
not
to
assume
the
upper
end
of
this
range
is
fully
protective.

Incubation
and
Pre­
emergent
Development
Chinook
Salmon
Once
spawning
has
taken
place,
the
eggs
of
chinook
salmon
hatch
in
about
2
months
and
the
young
remain
in
the
gravel
for
2­
3
weeks
prior
to
emerging.
Many
researchers
have
tested
incubation
survival
at
constant
exposure
to
various
test
temperatures.
Complete
mortality
(
100%)
has
been
noted
at
incubation
temperatures
from
13.9
to
19.4
°
C
(
Donaldson,
1955;
Garling
and
Masterson,
1985;
Seymour,
1956;
Eddy,
1972,
as
cited
in
Raleigh,
Miller,
and
Nelson,
1986).
Significant
mortality
(
over
50%)
has
been
noted
at
constant
incubation
temperatures
from
9.9
to
16.7
°
C
(
Donaldson,
1955;
Seymour,
1956;
Burrows,
1963,
and
Bailey
and
Evans,
1971;
as
cited
in
Alderdice
and
Velsen,
1978;
Hinze,
1959;
as
cited
in
Healy,
1979).
A
constant
incubation
temperature
of
8
°
C
produced
more
robust
alevin
and
fry
than
constant
exposure
to
either
4
or
12
°
C
in
a
study
by
Murry
and
Beacham
(
1986),
and
Velsen
(
1987)
compiled
data
showing
that
the
best
survival
(>
92.9%)
occurred
between
7.2­
9.6
°
C.
Heming
(
1982),
however,
found
good
survival
at
both
10
°
C
and
12
°
C.
Heming
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
43
Washington's
Surface
Water
quality
Standards
tested
the
survival
in
both
incubation
trays
and
artificial
redds.
Survival
rates
declined
as
the
temperatures
increased
from
6
to
8,
10,
and
12
°
C.
The
greatest
survival
(
91.7­
98%)
occurred
at
6
and
8
°
C,
but
it
was
still
very
good
(
90.2­
95.9%)
at
10
°
C.
Incubation
at
12
°
C
consistently
had
the
lowest
survival
(
84.6­
89.3%).
Heming
also
tested
survival
rates
from
incubation
to
hatching
against
survival
rates
from
hatching
through
complete
yolk
absorption.
His
work
suggests
higher
incubation
temperatures
may
create
a
metabolic
energy
deficit
for
pre­
emergent
salmon
that
increases
mortality.
Once
alevin
have
hatched
and
absorbed
their
yolk
sacs
they
will
need
to
make
a
transition
to
active
feeding.
Heming
and
McInery
(
1982)
found
that
temperatures
of
6,
8,
and
10
°
C
resulted
in
an
average
survival
of
98.4%
during
this
transitional
period,
while
12
°
C
was
associated
with
a
decrease
in
survival
to
89.2%.
Heming
(
1982;
as
cited
in
Beacham
and
Murray,
1986)
found
the
maximum
conversion
of
yolk
to
tissue
weight
occurred
at
6
°
C
or
below.
Seymour
(
1956)
noted
a
9­
fold
increase
in
abnormalities
in
fry
incubated
at
15.6
°
C
and
higher
when
compared
to
those
incubated
between
4.4­
12.8
°
C.
Seymour
also
noted
that
fry
incubated
at
4.4
°
C
emerged
larger
than
those
reared
at
higher
temperatures,
however,
subsequent
fry
growth
was
maximized
at
12.8
°
C.
Considered
together,
the
work
of
the
authors
cited
above
strongly
suggest
that
constant
temperatures
above
8­
9
°
C
or
below
5
°
C
may
reduce
the
survival
of
chinook
salmon
embryos
and
alevins.
A
constant
10
°
C
has
been
shown
to
produce
mixed
results
with
some
authors
finding
high
survival
rates
and
others
noting
significant
losses.
While
constant
temperatures
of
11­
12
°
C
can
still
result
in
good
success,
the
results
are
consistently
less
than
what
is
produced
at
lower
temperatures.

Some
researchers
have
tried
to
mimic
the
naturally
fluctuating
and
falling
temperatures
actually
experienced
by
incubating
eggs,
or
have
made
stepwise
reductions
in
the
incubation
temperatures
as
incubation
progressed.
Initial
incubation
temperatures
from
15.6­
16.7
°
C
have
been
associated
with
significant
to
total
losses
of
young
fish
through
the
incubation
to
early
fry
development
phase
(
Healy;
1979;
Brice,
1953;
CDWR,
1988;
Jewett,
1970;
as
cited
in
CDWR,
1988).
Rice
(
1960)
found
that
source
waters
declining
from
15.6­
8.3
°
C
resulted
in
satisfactory
egg
development,
though
did
not
provide
survival
rates
and
may
not
have
considered
survival
through
to
the
fry
stage.
Johnson
and
Brice
(
1953)
found
survival
to
often
exceed
90%
where
initial
water
temperatures
(
as
a
daily
mean)
were
below
12.2
°
C.
Healy
(
1979)
found
that
highest
survival
(
97%)
occurred
in
creek
water
where
the
daily
maximum
reached
12.8
°
C
only
a
few
times
during
the
first
two
weeks
of
development;
but
also
noted
that
survival
was
still
very
good
(
90­
94%)
where
the
initial
daily
maximum
temperatures
were
between
12.8­
14.2
°
C.
Olson
and
Nakatani
(
1969)
found
53.7­
88%
survival
in
egg
lots
started
at
12.5
°
C,
experiencing
a
brief
increase
to
14.7
°
C
(
daily
mean)
in
the
first
week,
and
then
quickly
dropping
back
to
12­
12.5
°
C
and
assuming
a
seasonal
downward
trend
in
temperature
(
test
water
paralleled
both
diel
and
seasonal
fluctuations
 
and
no
control
lots
were
provided).
Olson
and
Foster
(
1955)
found
the
greatest
survival
at
an
initial
test
temperature
(
daily
mean)
of
11.6
°
C
(
92.2%),
but
reported
no
appreciable
differences
in
survival
rates
at
initial
test
temperatures
of
13.8,
15,
and
16
°
C
(
89.9­
83.9%)
(
test
water
paralleled
seasonal
daily
average
temperatures).
Seymour
(
1956)
tested
four
geographically
distinct
stocks
of
chinook.
Taking
into
consideration
both
mortality
and
growth
rate,
the
optimum
temperature
was
estimated
as
11.1
°
C
for
eggs
and
fry.
The
mortality
rate
was
considered
low
at
all
stages
of
development
for
lots
reared
between
4.4
°
C
Page
44
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
and
12.8
°
C
(
daily
mean).
Lots
with
initial
temperatures
of
18.3
°
had
the
highest
mortality
(
11,
24,
40,
and
100%).
In
the
cyclic
and
fluctuating
temperature
tests
reviewed
here,
having
temperatures
at
the
beginning
of
incubation
that
are
below
11­
12.8
°
(
daily
mean)
are
typically
associated
with
excellent
survival
rates.
This
range
compares
well
with
the
optimal
temperature
range
of
8­
12
°
C
recommended
by
the
Independent
Scientific
Group
(
1996).

Donaldson
(
1955)
transferred
eggs
to
more
optimal
(
10­
12.8
°
C)
constant
incubation
temperatures
after
various
periods
of
exposure
to
higher
temperatures.
He
found
that
tolerance
to
temperature
exposure
varies
with
the
stage
of
development.
He
also
found
20%
mortality
could
be
induced
by
exposing
eggs
to
19.4
°
C
for
one
day,
18.3
°
C
for
thee
days
and
17.2
°
C
for
less
than
ten
days.
Donaldson's
work
lends
further
support
to
the
observations
made
by
other
authors
such
as
Jewett
(
1970;
as
cited
in
CDWR,
1988)
that
the
latent
effects
of
holding
eggs
at
higher
than
optimal
temperatures
continues
through
the
period
of
absorption
of
the
yolk
sac,
thus
using
mortality
estimates
at
the
time
of
hatching
underestimate
the
total
temperature
induced
mortality.
Donaldson
found
the
developmental
stages
associated
with
the
greatest
percentages
of
temperature
induced
mortality
were:
1)
the
time
up
until
the
closure
of
the
blastopore
(
200
T.
U.);
2)
the
period
just
previous
to
and
during
hatching;
and
3)
when
fry
are
adapting
themselves
to
feeding.
He
also
found
that
when
eggs
were
exposed
to
test
temperatures
(
17.2,
18.3,
and
19.4
°
C)
past
the
eye
pigmentation
stage
(
350
T.
U.)
the
time
necessary
for
complete
hatching
doubled,
and
the
frequency
of
common
abnormalities
increased
with
both
the
higher
temperatures
and
longer
exposures.
Donaldson's
work
also
suggests
that
a
week
or
more
exposure
at
an
average
of
17
°
C
would
likely
cause
at
least
moderate
mortality
(
20%),
even
if
the
water
were
quickly
and
substantially
cooled
after
that
initial
exposure.
Murry
and
Beacham
(
1986)
found
that
initial
incubation
at
4
°
C
reduced
survival
even
with
later
transfer
(
at
completion
of
epiboly)
to
warmer
waters
(
8
°
C
and
12
°
C).
Transfers
after
epiboly
or
completion
of
eye
pigmentation
from
4
to
12
°
C
and
from
12
to
4
°
C
also
caused
an
increase
in
alevin
mortality.
The
authors
also
found
that
a
decreasing
temperature
regime
produced
longer
and
heavier
alevins
and
fry.
Combs
(
1965)
found
that
eggs
developed
to
the
128­
cell
stage
at
5.8
°
C
could
then
tolerate
1.7
°
C
for
the
remainder
of
the
incubation
period
with
only
moderate
losses.
Mortality
of
14.5%
was
observed
with
a
transfer
time
of
72
hours,
while
only
3.3%
mortality
occurred
with
a
transfer
at
144
hours.
These
three
works
taken
together
suggest
that
the
effects
of
suboptimal
initial
incubation
temperatures
may
not
be
nullified
by
later
changes
in
the
temperature
regime
to
more
optimal
levels;
that
sudden
changes
in
temperature
at
either
early
or
later
stages
of
development,
regardless
of
the
direction
of
that
change,
can
be
harmful
to
pre­
emergent
life
stages;
and
that
initial
incubation
at
optimal
temperatures
may
condition
eggs
and
embryos
such
that
they
can
withstand
very
low
winter
temperature
regimes.
Neitzel
and
Becker
(
1985)
also
conducted
work
on
the
effects
of
short­
term
increases
in
temperature
that
can
be
used
to
support
daily
maximum
temperature
criteria.
Neitzel
and
Becker
(
1985)
used
chinook
salmon
to
try
and
determine
the
effects
of
short
term
de­
watering
of
redds
by
hydropower
facilities.
Neitzel
and
Becker
found
that
sudden
increases
in
temperatures
from
10
°
C
to
above
22
°
C
for
1­
8
hours
significantly
reduced
survival
of
cleavage
eggs
in
chinook
salmon.
Controls
held
at
10
°
C
experienced
very
low
mortalities
(
less
than
2%).
Mortality
in
treatment
groups
was
8­
10%
at
22
°
C
after
2
hours
exposure,
and
was
22%
after
a
one­
hour
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
45
Washington's
Surface
Water
quality
Standards
exposure
at
23.5
°
C.
They
further
found
that
decreasing
the
temperature
from
10
°
C
to
near
freezing
(
0.0
°
C)
for
up
to
24
hours
did
not
increase
mortality
in
eggs,
embryos,
or
alevin.

Considering
the
work
of
Donaldson
(
1955)
and
Neitzel
and
Becker
(
1985)
together
allows
a
reasonable
estimate
of
temperatures
that
would
avoid
causing
acute
effects
to
eggs
and
embryos.
Donaldson
found
that
a
constant
24­
hour
exposure
to
19.4
°
C
caused
modest
lethality.
This
is
relatively
comparable
to
Neitzel
and
Becker's
findings
that,
a
1­
8
hour
exposure
to
22
°
C
caused
moderate
lethality.
Adjusting
this
range
(
19.4­
22
°
C)
by
a
reduction
of
2
°
C
would
be
expected
to
reduce
the
likelihood
of
any
mortality
occurring
at
all,
and
produces
a
single
daily
maximum
or
one­
hour
duration
limit
of
17.4­
20
°
C
to
fully
protect
incubating
eggs
and
embryos
from
acute
lethality.

While
there
is
some
disagreement,
the
literature
is
very
consistent
overall
regarding
the
optimal
incubation
requirements
for
chinook
salmon.
To
provide
full
protection
from
fertilization
through
initial
fry
development
for
chinook
salmon,
average
daily
temperatures
should
remain
below
11­
12.8
°
C
at
the
initiation
of
incubation,
and
the
seasonal
average
should
not
exceed
8­
9
°
C
throughout
the
incubation­
through­
emergence
period.
To
protect
eggs
and
embryos
from
acutely
lethal
conditions
the
highest
single
day
maximum
should
not
exceed
17.5­
20
°
C.

Coho
Salmon
Embryo
survival
is
consistently
maximized
in
tests
conducted
at
constant
temperature
exposures
between
2.5­
6.5
°
C,
and
is
only
slightly
less
successful
between
1.3­
10.9
°
C
(
Dong,
1981;
Tang
et
al.,
1987;
Murry
et
al.,
1988;
Velsen,
1987).
Davidson
and
Hutchinson
(
1938;
as
cited
in
Sandercock,
1991)
suggested
the
optimum
temperature
for
egg
incubation
is
from
4­
11
°
C.
Mortalities
tend
to
become
moderate
(
21­
26%)
between
11­
12.5
°
C,
and
between
12.5
°
C
and
13.5
°
C
mortalities
of
50%
can
be
expected.
Above
14­
14.4
°
C,
tests
commonly
report
at
or
near
100%
mortality.
Alevin
survival
may
be
excellent
(
97%)
between
1.3­
10.9
°
C
(
Dong,
1981;
Tang
et
al.,
1987),
and
the
most
robust
fry
are
at
incubation
temperatures
between
4­
8
°
C
(
Dong,
1981;
Murry
et
al.,
1988).
Alevin
mortalities
of
51­
59%
occur
at
12.5
°
C
(
Dong,
1981),
and
100%
mortality
occurs
at
14­
14.4
°
C
(
Dong,
1981;
Murry
et
al.,
1988).

From
the
studies
discussed
above
we
can
be
relatively
confident
that
egg
survival
would
be
consistently
maximized
at
exposure
to
constant
temperatures
between
2.5­
6.5
°
C,
but
may
still
be
excellent
for
many
stocks
at
temperatures
between
1.3­
10.9
°
C.
Alevin
and
fry
survival
and
health
may
be
optimized
at
exposure
to
constant
temperatures
between
4­
8
°
C,
but
survival
may
remain
excellent
up
to
10.9
°
C.
Based
on
the
preceding,
the
incubation
of
coho
salmon
should
be
fully
support
by
maintaining
the
average
temperature
throughout
the
incubation
period
should
at
or
below
8­
10
°
C.

There
is
no
information
available
that
suggests
coho
salmon
embryos
and
alevin
would
be
more
sensitive
to
short
term
(
daily
peak)
increases
in
temperature
than
any
other
Pacific
salmon.
The
one
study
reviewed
that
looks
at
short
term
temperature
changes
for
coho
was
Page
46
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
that
by
Tang
et
al.
(
1987).
In
that
study,
incubation
temperatures
were
increased
from
10.2
to
17
°
C
and
lowered
from
10.2
to
4
°
C
for
eight
hours.
In
neither
test
did
these
modest
changes
result
in
any
statistically
significant
increase
in
mortality.
Additionally,
one
field
study
reported
coho
alevins
surviving
very
"
substantial"
daily
peak
temperatures
with
no
clear
change
in
later
juvenile
abundance
(
summer
peak
stream
temperatures
of
24
and
30
°
C
were
noted;
although,
its
is
highly
unlikely
the
eggs
were
actually
exposed
to
these
temperatures
and
were
most
likely
protected
by
placement
in
groundwater
seeps)
(
Hall
and
Lantz,
1969).
Since
no
clear
basis
has
been
found
for
setting
a
daily
peak
temperature
specific
to
coho
incubation,
it
is
suggested
that
the
daily
maximum
value
chosen
for
chinook
be
considered
appropriate
for
coho
salmon
as
well.

Pink
Salmon
The
range
for
successful
incubation
has
been
suggested
to
be
from
4.4
°
C
to
13.3
°
C
(
Bonar
et
al.,
1989).
Murray
and
Beacham
(
1986)
reported
excellent
survival
(
91­
97%)
with
initial
fertilization
occurring
at
constant
14
°
C
and
a
0.5
°
C
drop
in
temperature
every
three
days
down
to
5
°
C.
When
they
allowed
temperatures
to
drop
further
to
4
and
2
°
C
survival
was
reduced.
Murray
and
McPhail
found
survival
of
94%
from
fertilization
to
emergence
at
5
°
C,
and
Beacham
and
Murray
(
1986)
found
the
greatest
survival
for
5
stocks
and
21
families
of
pink
salmon
tested
at
8
°
C.
Velsen
(
1987)
compiled
data
that
showed
the
best,
though
highly
variable,
survival
(
generally
>
89.5%)
occurred
between
8­
13
°
C.
Survival
decreased
at
an
incubation
temperature
of
11
°
C
in
a
test
by
Murray
and
McPhail
(
1988),
and
was
50%
at
15­
15.5
°
C
(
Beacham
and
Murray,
1990).
Temperatures
of
5­
8
°
C
produced
the
largest
alevins
in
a
study
by
Murray
and
McPhail
(
1988),
and
8
°
C
produced
the
longest
(
Beacham
and
Murray,
1986)
and
fry
heaviest
(
Murray
and
McPhail,
1988).
Konecki
et
al.
(
1995)
incubated
embryos
from
ten
western
Washington
populations
and
found
significant
variation
(
1­
3
weeks)
in
incubation
rates
 
suggesting
that
some
differences
in
temperature
optimums
may
occur
between
stocks.

Survival
of
the
alevin
life­
stage
was
found
to
be
generally
excellent
(>
97%)
for
21
families
of
pink
salmon
tested
at
temperatures
ranging
from
4­
12
°
C
(
Beacham
and
Murray,
1986).
Survival
to
emergence
was
reported
as
low
at
14
°
C
(
Murray
and
McPhail,
1988).
Examining
low
incubation
temperatures,
Beacham
and
Murray
(
1986)
found
that
temperatures
of
4
°
C
consistently
resulted
in
the
lowest
survival
for
5
stocks
and
21
families
of
pink
salmon,
and
in
a
1990
study
found
50%
mortality
at
5
°
C.
Complete
mortality
has
been
found
at
an
incubation
temperature
of
2
°
C
(
Murray
and
McPhail,
1988;
Beacham
and
Murray,
1990;
Bailey
and
Evans,
1971).
Murray
and
Beacham
(
1986)
transferred
embryos
in
a
late
stage
of
development
from
8
°
C
to
1
°
C
and
found
that
while
northern
stocks
had
100%
survival,
southern
stocks
experienced
moralities
ranging
from
38­
60%.

Based
on
the
research
cited
above,
constant
temperatures
in
the
range
of
4.5­
12
°
C,
and
a
constantly
declining
temperature
regime
beginning
at
14
°
C
are
capable
of
producing
excellent
and
perhaps
optimal
survival
rates
of
incubating
pink
salmon.
However,
a
constant
temperature
of
8
°
C
appears
to
produce
the
most
consistently
optimal
results;
and
while
tests
up
to
12­
13.3
°
C
were
found
to
produce
excellent
results,
several
tests
found
temperatures
of
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
47
Washington's
Surface
Water
quality
Standards
11­
12
°
C
to
produce
lower
survival
and
smaller
fry.
Further,
in
natural
streams
the
temperatures
will
not
dependably
decline
at
a
steady
rate,
and
constant
temperatures
of
15­
16
°
C
have
resulted
in
high
mortality.
In
consideration
of
all
of
these
issues
it
should
be
assumed
that
constant
temperatures
in
the
range
of
8­
10
°
C
represent
upper
range
for
fully
supporting
the
embryonic
development
of
pink
salmon.
Therefore,
to
fully
protect
the
incubation
of
pink
salmon
the
daily
average
temperature
at
the
initiation
of
incubation
should
be
maintained
below
12­
14
°
C
and
the
average
temperature
throughout
incubation
through
emergence
should
be
not
exceed
8­
10
°
C.

Chum
Salmon
Incubation
survival
from
fertilization
to
emergence
is
variable,
but
can
be
excellent
anywhere
from
4­
12
°
C
(
Murray
and
Beacham,
1986;
Beacham
and
Murray,
1985).
In
the
initial
period
of
embryonic
development
temperatures
in
the
range
of
8­
12
°
C
produce
the
highest
survival.
However
in
later
stages
of
incubation,
temperatures
in
the
range
of
5­
8
°
C
produce
the
best
survival
as
well
as
the
largest
and
heaviest
alevin
and
fry
(
Beacham
and
Murray,
1986).
Temperatures
of
12
°
C
in
the
later
developmental
stages
can
result
in
heavy
losses
in
some
stocks
(
Beacham
and
Murray,
1985;
Beacham
and
Murray,
1986).
The
optimal
temperature
range
for
conversion
of
yolk
to
tissue
weight
was
estimated
to
be
from
6­
10
°
C
(
Beacham
and
Murray,
1986),
and
optimal
respiration
efficiency
has
been
estimated
to
range
from
11­
12.5
°
C
for
prolarvae
and
larvae
(
Zinichev
and
Zotin,
1988).
Constant
incubation
at
temperatures
of
14
°
C
and
16
°
C
as
well
as
at
2.5
°
C
have
been
associated
with
embryonic
mortalities
of
50%
(
Beacham
and
Murray,
1990).
The
alevin
stage
of
development
(
late),
however,
was
shown
to
have
very
high
survival
rates
when
exposed
to
temperatures
as
low
as
2
°
C.
Beacham
and
Murray
(
1987)
found
that
10
°
C
resulted
in
higher
growth
and
survival
(
96%)
of
emergent
fry.
Fry
survival
rates
were
62%
at
13
°
C,
and
73%
at
16
°
C.

Based
on
the
literature
reviewed,
constant
incubation
temperatures
from
4­
12
°
C
can
produce
excellent
and
incubation
results;
however,
some
researchers
have
noted
reductions
in
survival
occurring
at
the
ends
of
this
range.
Constant
incubation
temperatures
in
the
range
6­
10
°
C
would
be
most
consistently
fully
protective
for
chum
salmon.
Average
temperatures
at
the
initiation
of
incubation
should
not
exceed
11­
12
°
C
to
allow
for
the
full
protection
of
developing
eggs.
To
provide
for
the
full
protection
of
the
period
of
development
from
incubation
through
emergence
in
chum
salmon,
the
daily
average
temperature
at
the
beginning
of
the
incubation
period
should
not
exceed
11­
12
°
C
and
the
seasonal
average
temperature
should
not
8­
10
°
C.
Page
48
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Sockeye
Salmon
Murray
and
McPhail
(
1988)
and
Combs
(
1965)
report
that
sockeye
salmon
are
more
tolerant
of
low
incubation
temperatures
and
less
tolerant
of
high
incubation
temperatures
than
the
other
Pacific
Salmon.
At
constant
exposure,
Combs
(
1965)
reported
that
temperatures
within
the
range
of
4.4­
12.7
°
C
produced
similarly
high
survival
rates
(
85.8­
90.9%),
with
the
highest
occurring
at
5.8
°
C.
Combs
found
incremental
increases
in
mortality
of
53­
67%
occurred
when
the
temperature
was
lowered
from
5.8
to
4.4
°
C
or
raised
from
12.7
to
14.2
°
C.
Velsen
(
1987)
found
that
while
survival
rates
were
highly
inconsistent
between
1.1­
15
°
C,
the
best
survival
generally
occurred
between
3.1­
5.8
(
generally
>
90%),
with
fair
survival
(>
70%)
occurring
in
the
range
of
2.1­
12.7
°
C,
and
survival
rates
consistently
poor
(
17­
76%)
above
14
°
C.
Murray
and
McPhail
(
1988)
found
that
survival
was
highest
at
8
°
C
(
79%)
but
only
40%
at
both
11
and
5
°
C.
Andrew
and
Geen
(
1960)
reported
that
in
the
first
two
years
of
a
four
year
field
study,
eggs
initially
incubated
at
temperatures
of
7.2
°
C
had
lower
survival
than
those
initially
incubated
at
10,
12.8,
and
15.6
°
C.
In
a
follow­
up
experiment
the
following
two
years,
they
found
that
eggs
exposed
to
temperatures
of
15.6
to
16.7
°
C
for
"
short
periods
of
time"
suffered
severe
losses
during
the
period
of
exposure,
and
that
temperatures
of
16.7­
18.3
°
C
caused
extensive
losses
both
during
and
following
the
exposure.
In
a
study
by
Craig
et
al.
(
1996)
the
temperature
range
of
8­
10
°
C
resulted
in
the
optimum
1:
1
male
to
female
sex
ratio
in
offspring;
although,
the
study
design
really
only
allows
the
conclusion
that
temperature
in
the
early
stage
of
development
affects
sex
determination.

In
the
studies
cited
above,
constant
temperatures
in
the
range
of
4­
12.7
°
C
produce
variable
but
oftentimes
excellent
survival
rates
in
sockeye
salmon;
but
that
the
range
6­
10
°
C
appears
most
consistently
optimum.
Severe
losses
have
been
noted
at
and
above
15.6
°
C,
even
with
short
term
exposure
under
fluctuating
field
conditions,
and
poor
survival
occurred
at
constant
temperatures
of
14­
14.2
and
above.
Taken
together
this
information
suggests
that
daily
average
temperatures
in
the
early
incubation
period
should
not
exceed
11­
12.7
°
C,
and
the
average
temperature
throughout
the
period
of
incubation
to
emergence
should
not
exceed
8­
10
°
C
to
fully
protect
the
incubation
success
of
sockeye
salmon.

Steelhead
and
Rainbow
Trout
Fuss
(
1998)
considered
the
range
5.6­
11.1
°
C
as
being
optimal
for
steelhead
egg
survival
in
the
Washington
State
hatchery
program,
and
Bell
(
1986)
suggests
that
10
°
C
is
the
preferred
hatching
temperature
for
steelhead
eggs.
Rombough
(
1988)
found
less
than
4%
embryonic
mortality
at
6,
9,
and
12
°
C,
but
noted
an
increase
to
15%
mortality
at
15
°
C.
Alevin
mortality
was
less
than
5%
at
all
temperatures
tested,
but
alevins
hatching
at
15
°
C
were
considerably
smaller
and
appeared
less
well
developed
than
those
incubated
at
the
lower
test
temperatures.
Redding
and
Schreck
(
1979)
similarly
found
that
emergent
fry
were
larger
at
12
°
C
than
at
16
°
C.

Velsen
(
1987)
compiled
data
on
the
incubation
survival
of
both
anadromous
(
steelhead)
and
non­
anadromous
rainbow
trout
that
showed
survival
was
consistently
high
(>
92%)
between
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
49
Washington's
Surface
Water
quality
Standards
4­
9
°
C,
and
fair
(>
78%)
between
3­
15
°
C,
but
very
poor
(
7%)
above
16
°
C.
Kamler
and
Kato
(
1983)
tested
incubation
survival
of
rainbow
trout
at
9,
10,
12,
14,
and
16
°
C.
They
found
the
highest
survival
of
eggs
at
10
°
C
and
12
°
C,
slightly
lower
survival
at
14
°
C,
and
abrupt
drops
in
survival
at
both
9
°
C
and
16
°
C.
Survival
to
the
swim­
up
stage
in
two
strains
of
rainbow
trout
had
94­
98%
survival
at
7
°
C,
72­
95%
at
4
°
C,
and
<
12­
41%
survival
at
2
°
C
(
Stonecypher
and
Hubert,
1994).
Kwain
(
1975)
found
the
lowest
mortalities
of
rainbow
trout
occurred
at
7
and
10
°
C,
and
Kashiwagi
et
al.
(
1987;
as
cited
in
Taylor
and
Barton,
1992)
found
optimal
hatching
occurred
at
10
°
C.
Humpesh
(
1985)
found
that
optimal
hatching
(>
90%)
occurred
between
7­
11
°
C
in
rainbow
trout,
and
Alekseeva
(
1987;
as
cited
in
Taylor
and
Barton,
1992)
suggested
that
optimal
incubation
occurs
with
temperatures
rising
from
5.3­
10.5
°
C.
Constant
temperatures
10­
12
°
C
commonly
form
the
upper
end
of
the
temperature
range
that
provides
excellent
support
for
the
incubation
and
hatching
of
anadromous
and
non­
anadromous
forms
of
O.
Mykiss.
Losses
become
moderate
at
temperatures
of
14­
15
°
C
and
severe
above
16
°
C.
Maintaining
a
seasonal
average
temperature
of
7­
10
°
C
appears
to
provide
for
the
most
consistent
occurrence
of
optimal
survival
and
development
in
O.
Mykiss.
Thus,
it
can
be
reasonably
concluded
that
the
average
temperature
throughout
the
development
of
O.
Mykiss
should
not
exceed
be
7­
10
°
C,
and
the
maximum
daily
average
temperature
at
the
time
of
hatching
should
be
maintained
below
11­
12
°
C.

Statewide
spawning
dates
were
not
found
for
non­
anadromous
rainbow
trout.
In
the
absence
of
such
information,
it
is
suggested
that
dates
associated
with
the
anadromous
forms
of
O.
mykiss
be
considered
if
need
arises
to
set
dates
of
application
for
the
spawning
and
incubation
criteria.

Cutthroat
Trout
Eggs
of
sea­
run
cutthroat
incubate
for
6­
7
weeks
before
they
hatch
and
the
alevin
remain
in
the
gravel
for
about
another
2
weeks
before
they
emerge
(
Trotter,
1989;
Pauley
et
al.
1989).
Fry
may
emerge
from
March
through
June,
depending
on
the
location
and
time
of
spawning,
but
peak
emergence
occurs
in
mid
April
(
Trotter,
1989;
Wydoski
and
Whitney,
1979).

Pauley
et
al.
(
1989)
cite
studies
demonstrating
that
the
optimum
temperature
for
incubation
is
10­
11
°
C.
Bell
(
1986)
has
suggested
that
the
range
for
hatching
of
cutthroat
trout
eggs
is
from
4.4­
12.8
°
C.
Smith
et
al.
(
1983)
found
that
west­
slope
cutthroat
trout
eggs
held
in
creek
water
with
a
fluctuating
temperature
of
2­
10
°
C
had
significantly
better
survival
than
eggs
held
at
a
constant
10
°
C.
Stonecypher
and
Hubert
(
1994)
found
that
survival
to
swim­
up
stage
in
Snake
River
cutthroat
trout
was
95%
at
7
°
C,
approximately
87%
at
4
°
C,
and
less
than
16%
at
2
°
C.
Hubert
and
Gern
(
1995)
found
68.6%
survival
in
a
control
population
held
at
7
°
C
when
testing
the
effects
of
lowering
incubation
temperatures
in
the
early
stage
of
development.
Mortality
rates
were
no
different
from
controls
when
temperatures
were
lowered
to
3
°
C
at
least
13­
15
days
after
fertilization
but
were
higher
if
the
cooling
took
place
sooner.
Page
50
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
It
is
somewhat
problematic
to
set
standards
for
the
incubation
of
cutthroat
trout
that
can
be
reasonably
applied
statewide.
As
a
spring
spawning
species
that
often
spawns
high
in
the
watershed,
cutthroat
trout
have
a
very
broad
period
of
spawning
when
examined
statewide.
Stocks
that
exist
in
lower
or
warmer
watersheds
will
spawn
as
early
as
February
when
temperatures
rise
above
6
°
C,
while
stocks
that
exist
in
high
elevation
snow
fed
streams
may
need
to
wait
until
late
June
or
July
for
waters
to
be
sufficiently
warm
(
6­
11
°
C)
to
allow
successful
spawning.
Were
it
not
for
risk
of
egg
loss
due
to
late
winter
and
spring
freshets,
one
could
suggest
that
the
spring
spawning
strategy
is
relatively
unencumbered
by
changes
in
the
temperature
regime.
While
earlier
spawning
subjects
cutthroat
eggs
to
higher
risks
of
physical
damage,
the
earlier
hatch
also
places
surviving
resident
fry
in
a
good
position
to
maximize
summer
growth
and
thus
increase
their
survival
opportunities
over
the
following
winter.
It
may
well
be
that
the
superior
growth
obtained
in
the
oceanic
phase
of
anadromous
forms
may
make
minor
differences
in
weight
gains
from
earlier
emergence
of
less
value,
but
this
relationship
remains
to
be
tested.
In
general,
specific
stocks
will
have
tailored
their
spawning
and
emergence
periods
to
optimize
both
incubation
survival
and
early
fry
growth.
Significant
changes
in
the
temperature
regime,
such
as
earlier
spring
warming
will
bring
unknown
risks
to
individual
populations.
Therefore,
while
a
fully
protective
temperature
regime
is
suggested
in
this
document
for
cutthroat
trout,
it
would
be
best
tailored
to
the
historic
patterns
of
spawning
found
in
specific
stocks.
To
initiate
spawning
in
most
stocks
the
water
temperatures
must
at
least
warm
to
daily
maximums
of
6­
7
°
C,
though
some
stocks
may
not
begin
spawning
until
temperatures
reach
11
°
C.
Specific
studies
on
incubation
survival
suggest,
however,
that
incubation
may
be
optimized
with
constant
temperature
exposures
in
the
range
of
7­
11
°
C;
although,
the
data
suggests
their
could
sometimes
be
incubation
losses
at
constant
water
temperatures
of
10­
11
°
C.
Statewide,
most
cutthroat
spawn
in
mid­
February
and
hatch
by
the
end
of
March.
It
is
suggested
that
in
the
absence
of
more
watershed­
specific
information
April
1
could
be
used
to
compare
the
incubation
threshold
recommendations
statewide.
Given
that
the
studies
reviewed
create
some
contradictions,
it
is
believed
that
the
middle
portion
of
the
possible
fully
protective
range
should
be
considered
most
defensible
for
use
in
setting
any
final
recommendations.
To
fully
protect
the
period
of
incubation
through
emergence
in
cutthroat
trout,
the
available
information
suggests
that
the
average
temperature
throughout
the
entire
development
period
should
not
exceed
8­
9
°
C,
and
the
daily
average
temperatures
should
not
exceed
11­
12
°
C
until
after
emergence.

Summary
of
Laboratory
and
Field
Studies
on
Incubation
Success:
Across
all
species
of
salmonids
examined,
full
protection
would
be
achieved
by
maintaining
an
overall
average
temperature
throughout
the
period
of
incubation
through
emergence
of
8­
9
°
C,
and
by
not
exceeding
12­
12.8
°
C
as
a
daily
average
temperature
at
the
initiation
of
incubation
in
fall
spawning
stocks
and
at
the
time
of
emergence
in
spring
spawning
stocks
(
see
Table
4.4,
below).
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
51
Washington's
Surface
Water
quality
Standards
Table
4.4.
Constant
or
average
temperature
exposures
estimated
to
allow
for
full
protection
of
the
incubation
of
individual
salmon
and
trout
species.

Species
Daily
Average
Temperature
at
Initiation
Seasonal
Average
Temperature
Chinook
11­
12.8
8­
9
Coho
­­­­
8­
10
Chum
12­
14
8­
10
Pink
11­
12
8­
10
Sockeye
11­
12.7
8­
10
Steelhead/
Rainbow
11­
12
7­
10
Cutthroat
11­
12
8­
9
Summary
11.16­
12.58
(
mid.
11.87)
7.86­
9.71
(
mid.
8.79)

Converting
these
temperatures
to
a
standardized
7DADMax
metric
will
facilitate
the
comparison
with
other
lines
of
evidence
on
this
life
stage.
It
seems
appropriate
to
assume
the
overall
average
temperature
(
based
on
constant
temperature
exposure
throughout
the
incubation
period)
represents
a
seasonal
average
(
which
gives
a
7DADMax
of
11.29­
12.29,
and
the
initial
incubation
temperature
(
based
on
tests
that
allowed
temperatures
to
cool
during
incubation
and
that
recorded
temperature
as
a
daily
average)
is
represented
best
as
either
a
daily
average
(
7DADMax
13.48­
13.77
°
C)
or
a
weekly
average
(
7DADMax
13.96­
14.76
°
C)
(
conversions
are
based
on
assuming
stream
diel
temperature
ranges
of
2­
4
°
C).

Table
4.5.
Spawning
and
incubation
requirements
of
salmon
and
trout.

Line
of
Evidence
7DADM
­
Range
(
°
C)
Median
(
°
C)
General
observations
on
spawning
initiation
(
generally
field
observations)
11.95­
13.65
12.8
Prespawning
effects
(
lab
and
field
studies)
13.48­
14.96
14.22
Incubation
success
(
as
highest
1
week
period)
13.48­
14.76
14.12
Incubation
success
(
as
a
average
across
the
entire
life­
stage)
11.29­
12.29
11.79
Best
Estimate
of
Threshold
12.55­
13.92
mid.
pt.
13.24
The
range
of
these
independent
lines
of
evidence
is
11.29­
14.96
°
C
with
a
mean
range
of
12.55­
13.92
°
C
and
an
overall
midpoint
of
13.24
°
C.
This
suggests
that
to
fully
protect
salmon
and
trout
during
the
period
from
incubation
through
emergence,
temperatures
should
not
exceed
a
7­
DADMAx
of
13.24
°
C.
After
cross
checking
the
conclusion
against
each
independent
line
of
evidence,
no
overriding
factors
of
disagreement
appear
to
exist.
The
largest
disagreement
is
with
the
season­
wide
estimate
of
incubation
success,
however,
this
seems
well
counterbalanced
by
the
estimate
of
initial
incubation
temperatures
relating
to
full
protection.
The
similarity
between
species
for
each
line
of
evidence,
and
the
similarity
Page
52
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
between
each
of
the
lines
of
evidence
lend
greater
credibility
to
the
overall
estimate
of
a
temperature
regime
that
will
fully
protect
the
salmonids
from
incubation
through
emergence.

To
protect
eggs
and
embryos
from
acutely
lethal
conditions
that
may
arise
under
unique
situations,
the
highest
single
day
maximum
should
not
exceed
17.5­
20
°
C.
(
Note:
the
conversion
estimates
would
have
been
raised
by
1.12,1.22,
and
1.35
°
C
for
the
daily
average,
weekly
average,
and
seasonal
average
estimates,
respectively,
if
it
were
assumed
the
streams
would
have
an
average
seasonal
diel
fluctuation
of
4­
6
°
C
rather
than
2­
4
°
C,
however,
a
lower
range
seems
more
appropriate
for
use
in
the
incubation
period).

iii)
Juvenile
Rearing
General
Introduction:

Attaining
good
growth
is
very
important
to
the
survival
of
young
salmon.
In
a
study
by
Burrows
(
1963),
it
was
found
that
doubling
the
weight
of
sockeye
salmon
fingerlings
released
from
a
hatchery
resulted
in
a
tripling
of
the
adult
return
rate.
However,
in
determining
temperatures
that
will
fully
protect
juvenile
salmonids,
growth
is
not
the
only
endpoint
of
concern.
It
is
important
to
note
that
disease,
predation,
and
smoltification
concerns
are
all
important
issues
that
need
to
be
considered.
Additionally,
when
considering
growth
rates,
it
is
important
to
distinguish
between
studies
conducted
under
constant
and
cyclic
temperature
regimes,
and
to
recognize
that
the
feeding
rates
used
in
testing
will
affect
the
results
of
the
test.
As
food
becomes
more
limited,
the
temperature
that
provides
for
optimal
growth
declines.
As
food
becomes
more
plentiful,
fish
can
grow
larger
in
warmer
waters.
Since
most
tests
are
conducted
at
high
to
very
high
rations,
the
results
must
be
cautiously
applied
to
natural
stream
environments
where
food
availability
can
be
more
limiting.

Laboratory
Growth
Studies
Conducted
at
Constant
Temperatures:

Chinook
Salmon
In
constant
temperature
experiments
conducted
at
high
feeding
rates,
maximum
growth
tends
to
be
associated
with
temperatures
in
the
range
of
15­
19
°
C
(
Hillman,
1991;
Cech
and
Myrick,
1999;
Brett
et
al.,
1982;
Banks
et
al.,
1971).
Banks
et
al.
(
1971)
found
that
growth
was
similar
at
15.6
°
C
and
18.3
°
C,
with
15.6
°
C
being
slightly
higher
in
2
of
4
test
lots.
In
the
two
lots
where
18.3
°
C
produced
better
growth;
however,
the
fish
in
this
warmer
water
showed
higher
rates
of
disease
incidence.
Temperatures
above
19
°
C
were
associated
with
reduced
feeding
and
growth,
as
well
as
increased
problems
with
disease.
Marine
and
Cech
(
1998,
as
cited
in
USEPA,
2001)
found
that
growth
was
substantially
reduced
at
21­
24
°
C
when
compared
to
13­
16
°
C.
Brett
et
al.
(
1982)
estimated
that
under
natural
ration
levels
the
optimum
of
19
°
C
would
be
reduced
to
14.8
°
C
and
no
growth
would
be
possible
at
21.4
°
C.
Seymour
(
1956)
studied
three
Washington
and
one
California
stock
of
chinook
salmon
and
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
53
Washington's
Surface
Water
quality
Standards
concluded
that
the
general
optimum
temperature
for
growth
and
survival
of
chinook
fingerlings
was
14.4
°
C.
Zaugg
and
Wagner
(
1973)
found
that
at
the
end
of
a
16­
week
test,
growth
was
roughly
27%
greater
at
12
°
C
than
at
8
°
C.
Based
on
the
foregoing
works,
maximum
growth
should
be
expected
with
constant
exposure
at
15.6­
19
°
C.
The
increase
in
growth
above
15.6
°
C,
however,
was
inconsistently
greater,
and
under
natural
rations
the
maximum
growth
rate
may
decline
by
as
much
as
4.2
°
C
(
14.8
°
C).

Coho
Salmon
Maximum
growth
of
coho
fed
to
satiation
in
the
laboratory
occurred
at
constant
17
°
C
in
a
study
by
Shelbourn
(
1980).

Sockeye
Salmon
Brett
et
al.
(
1969;
as
cited
in
USEPA,
2001)
found
that
optimum
growth
occurs
at
15
°
C
on
unrestricted
rations,
but
when
the
feeding
rate
was
reduced
to
1.5%
of
body
weight/
day
the
optimal
growth
rate
lowered
to
10
°
C
and
growth
was
zero
at
15
°
C.
Brett
and
Glass
(
1973)
noted
that
10
°
C
resulted
in
the
most
efficient
use
of
food
and
that
the
reduction
from
15
to
10
°
C
created
a
33%
reduction
in
the
demand
for
food.
Brett
and
Groves
(
1979)
note
that
at
temperatures
below
10
°
C
food
conversion
efficiency
rises
most
rapidly
from
the
base
level
(
maintenance
ration)
and
reaches
peak
efficiency
at
an
intermediate
ration.
This
relation
changes
progressively
with
rising
temperature
such
that
at
17
°
C
and
above
the
highest
efficiency
occurs
on
a
maximum
ration
diet.
They
note
that
for
sockeye
salmon,
over
the
complete
range
of
tolerable
temperatures
maximum
food
conversion
efficiency
occurs
at
11
°
C.
Brett
(
1971;
as
cited
in
USEPA,
2001)
found
swimming
capacity,
metabolic
scope,
growth
on
excess
rations,
and
ingestion
were
maximized
at
15
°
C,
and
that
15
°
C
was
also
the
final
preferendum
temperature.
Wurtsbaugh
and
Davis
(
1977)
found
that
optimum
growth
occurred
at
16.5
°
C
on
full
rations.
Donaldson
and
Foster
(
1941;
as
cited
in
Brett,
Hollands,
and
Alderdice,
1958)
discovered
that
growth
in
young
sockeye
tends
to
be
poor
to
none­
at­
all
at
temperatures
above
21
°
C
and
below
4
°
C.
Brett
(
1956)
noted
that
sockeye
juveniles
may
cease
feeding
entirely
when
temperatures
reach
21
°
C.
Temperatures
in
the
range
of
4­
7
°
C
have
been
associated
with
poor
growth
(
Brett,
1956).
The
foregoing
works
suggest
that
maximum
growth
may
occur
at
constant
temperatures
of
15­
16.5
°
C
under
unrestricted
rations,
but
that
under
severely
restricted
rations
the
point
of
maximum
growth
may
decline
as
much
as
5
°
C
(
to
10
°
C).

Steelhead
and
Rainbow
Trout
Olson
and
Tempelton
(
undated
manuscript;
as
cited
in
USEPA,
1973)
reportedly
found
that
the
most
favorable
range
for
steelhead
growth
was
between
5­
17
°
C,
with
a
physiological
optimum
in
the
vicinity
of
15
°
C.
The
amount
of
food
necessary
to
maintain
the
fishes'
weight
increased
rapidly
as
temperatures
rose
above
12
°
C
with
no
growth
occurring
at
approximately
23
°
C
despite
the
presence
of
excess
food.
It
was
suggested
that
the
most
efficient
growth,
within
the
consumption
ranges
believed
to
occur
in
nature,
is
at
the
temperature
of
5­
14
°
C
in
the
early
spring,
11­
14
°
C
in
early
summer,
14­
17
°
C
in
late
Page
54
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
summer,
11­
17
°
C
in
fall,
and
5­
8
°
C
in
winter.
Behnke
(
1992)
cites
work
showing
that
rainbow
trout
reduce
and
finally
cease
feeding
as
temperatures
rise
to
between
22­
25
°
C,
often
well
below
the
lethal
temperature.

Cech
and
Myrick
(
1999)
tested
winter­
run
steelhead
from
the
American
river
in
California
at
three
different
temperature
(
11,
15
and
19
°
C)
and
ration
levels
that
varied
from
82%
to
100%
of
satiation.
They
found
that
while
a
12%
reduction
in
feeding
rate
resulted
in
a
statistically
significant
reduction
of
growth
at
19
°
C,
the
growth
rate
was
still
higher
than
that
observed
at
any
feeding
levels
at
11
°
C
and
15
°
C.
This
suggests
that
the
maximum
growth
rate
under
more
moderate
level
of
feeding
would
likely
occur
between
15­
19
°
C.

Behnke
(
1992,
citing
Dwyer
et
al.
1981,
1983a,
1983b)
suggested
that
the
optimum
temperature
for
the
growth
and
food
assimilation
in
salmonids
occurs
between
13­
16
°
C.
Taylor
and
Barton
(
1992)
recommended
temperature
criteria
for
Alberta
in
British
Columbia
of
12­
19
°
C
as
a
7­
day
average.
In
a
review
on
rainbow
trout,
Moyle
(
1976)
opined
that
the
optimal
temperature
for
growth
and
the
completion
of
most
life
history
stages
was
13­
21.
The
optimal
temperature
range
was
cited
by
Bell
(
1986)
as
being
12.2­
18.9
°
C
with
the
most
optimal
set
at
13.9
°
C.
Piper
et
al.
(
1982)
in
a
hatchery
management
review
paper
set
the
optimal
at
10­
16.7
°
C.
Mckee
and
Wolf
(
1963;
cited
in
Wedemeyer
et
al.,
date
absent
from
copy)
are
reported
to
have
found
13
°
C
to
be
optimal.

Behnke
suggests
that
redband
trout
from
an
Oregon
desert
basin
have
been
demonstrated
to
have
an
optimum
feeding
temperature
at
some
untested
temperature
higher
than
19
°
C
(
citing
unpublished
work
from
the
U.
S.
Fish
and
Wildlife
Service,
Cultural
Development
Center,
Boozeman,
Montana).
This
was
contrasted
to
work
done
at
the
same
research
facility
on
other
forms
of
rainbow
trout
that
showed
growth
optimum
feeding
peaked
at
10­
16
°
C.
Behnke
(
1992)
suggests
that
the
work
done
to
date
demonstrates
that
the
desert
redband
have
a
functional
feeding
temperature
that
is
higher
than
rainbow
trout
that
have
evolved
in
less
harsh
environments
of
temperature
and
water
flow.
Kaya
(
1978)
compared
an
introduced
population
of
interior
rainbow
trout
from
the
Firehole
River
in
Montana
to
two
hatchery
strains.
Temperatures
in
the
Firehole
River
at
times
reach
summer
maximums
as
high
as
29.5
°
C
due
to
thermal
springs.
The
planted
stock
has
been
living
in
the
river
for
approximately
20
generations,
yet
it
was
found
that
neither
the
functional
feeding
temperature
or
the
upper
incipient
lethal
temperature
had
increased
in
comparison
to
the
hatchery
stocks.

Reimers
(
1957)
found
that
raising
water
temperatures
(
from
9.4
to
16.7
°
C)
at
the
conclusion
of
starvation
testing,
increased
metabolic
demands
sufficiently
to
cause
mortality
in
the
weakened
fish,
and
found
that
fish
that
began
the
trials
as
well
nourished
hatchery
rainbow
trout
survived
better
than
stream­
conditioned
wild
brown
trout.
This
was
to
point
out
the
importance
of
beginning
the
winter
period
with
good
energy
reserves.

Hokanson
et
al.
(
1997)
found
that
a
constant
exposure
to
17.2
°
C
produced
the
greatest
growth
rates
in
trout
fed
to
satiation
over
a
30­
day
test
period.
Maximum
growth
was
estimated
to
occur
between
15­
20
°
C
(
mean
17.5
°
C)
by
Cho
and
Kaushik
(
1990;
as
cited
in
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
55
Washington's
Surface
Water
quality
Standards
Dockray
et
al.,
1996).
Sadler
et
al.
(
1986)
found
that
growth
and
food
conversion
efficiency
were
greater
at
16
°
C
as
compared
with
10
°
C.
Myrick
and
Cech
(
2000)
found
that
the
growth
rates
of
two
strains
of
rainbow
trout
from
California
tended
to
increase
as
temperatures
increased
from
10
to
19
°
C,
with
the
significant
increase
occurring
from
10­
14
°
C,
and
to
decrease
to
near
zero
as
temperatures
increased
from
19
to
25
°
C.
They
also
found
that
while
the
growth
of
the
two
strains
was
similar
at
low
temperatures,
the
hatchery
strain
grew
much
better
then
the
wild
strain
when
exposed
to
temperatures
in
the
upper
range
(
22
and
25
°
C).
The
authors
noted
that
the
increasing
trend
in
food
consumption
and
growth
rates
for
the
two
strains
between
14­
19
°
C,
suggests
the
optimal
temperature
lies
between
these
to
temperatures.
Huggins
(
1975)
suggested
older
fish
sometimes
demonstrate
more
intermediate
temperature
preferences.
While
the
conclusions
of
the
foregoing
authors
are
highly
variable,
the
estimates
at
the
low
end
of
the
range
all
come
from
recommendation
papers
or
indirect
citations.
For
this
reason,
preference
is
being
given
the
conclusions
of
research
papers
directly
reviewed.
It
is
therefore
estimated
that
growth
may
be
maximized
at
temperatures
as
high
as
17.2­
19
°
C
under
satiation
rations.

Cutthroat
Trout
Pauley
et
al.
(
1989)
cite
research
concluding
that
the
optimal
temperature
for
juveniles
is
15
°
C,
and
that
equilibrium
and
ability
to
swim
is
lost
between
28­
30
°
C.
Hickman
and
Raleigh
(
1982;
as
cited
by
Barton,
1996)
suggest
that
the
optimal
range
for
cutthroat
trout
is
12­
15
°
C.
Data
reviewed
by
Carlander
(
1969;
as
cited
in
Gresswell,
1995)
suggest
optimum
water
temperatures
between
4.4
and
15.5
°
C
for
Yellowstone
cutthroat
trout.
A
temperature
of
15­
15.5
°
C
is
consistently
cited
as
optimal
for
the
growth
of
cutthroat
trout.

Table
4.6.
The
following
summarizes
the
conclusions
from
laboratory
tests
where
the
fish
were
exposed
to
constant
test
temperatures:

Species
Temperature
(
°
C)
Feeding
Rate
Comments
Author
Chinook
15.6
and
18.3
Satiation
Highest
growth
each
in
2
of
4
tests,
Higher
disease
at
18.3
Banks
et
al.,
1971
19
Satiation
Brett
et
al.,
1982
14.8
60%
of
satiation
A
4.2C
reduction
from
max
at
satiation
feeding
Brett
et
al.,
1982
14.4
Satiation
Three
Washington
and
one
Calif.
stocks
Seymour,
1956
Coho
17
Satiation
Shelbourn,
1980
Sockeye
15
Satiation
Brett
et
al.,
1969
10
1.5%
body
A
5C
reduction
from
Brett
et
al.,
Page
56
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
wt.
max
at
satiation
1969
16.5
Satiation
Wurtsbaug
and
Davis,
1977
17
Satiation
Highest
growth
efficiency
Groves,
1979
Steelhead
15
Satiation
Olson
and
Tempelton
19
Satiation
and
88%
Cech
and
Myrick,
1999
13.9
Satiation
Bell,
1986
13
Satiation
Mckee
and
Wolf
Redband
19
Satiation
Maximum
may
be
higher
Behnke,
1992
Rainbow
17.2
Satiation
Hokanson
et
al.,
1997
17.5
Satiation
Cho
and
Kaushik,
1990
19
Satiation
Myrick
and
Cech,
2000
15
Satiation
Based
on
peak
of
plotted
results
Grabowski,
1973
Cutthroat
15
Satiation
Pauley,
1989
17.5
Satiation
Based
on
peak
of
plotted
results
Clarke,
1978
Summary
16.41(
mean)
Satiation
Range
13­
19
Optimal
growth
rates
under
high
feeding
levels
have
been
found
at
constant
temperature
exposures
from
13­
19
°
C
with
a
mean
of
16.41
°
C.
However,
most
studies
group
the
estimate
more
narrowly,
between
14­
17.5
°
C
(
15.75
°
C).
When
feeding
is
reduced
substantially,
however,
the
temperature
that
will
allow
for
maximum
growth
has
been
noted
to
decrease
by
4­
5
°
C.
Thus
under
natural
feeding
regimes,
where
food
availability
is
more
restricted
during
the
high
temperature
months
of
summer,
maximum
growth
rates
would
be
more
likely
to
be
obtained
at
a
constant
temperatures
of
11­
13.5
°
C
(
assuming
a
4
°
C
reduction
due
to
restricted
feeding).
If
treated
as
either
a
weekly
average
or
summer
average
temperature
regime
this
constant
temperature
range
would
be
converted
to
estimated
7­
DADMax
temperature
ranges
of
14.18­
16.68
°
C
(
15.43)
and
15.64­
18.14
°
C
(
16.89),
respectively.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
57
Washington's
Surface
Water
quality
Standards
Ranges
Identified
in
the
Literature
as
Optimal
for
Growth:

The
upper
end
of
the
ranges
estimated
as
optimal
for
growth
in
the
literature
ranged
from
15­
21
°
C
with
a
midpoint
of
18
°
C
(
Table
4.7).
It
was
not
always
clear
from
the
literature
why
the
authors
chose
the
specific
ranges
that
were
recommended,
whether
or
not
they
had
tried
to
set
the
ranges
to
maximum
growth,
or
what
temperature
metric
(
i.
e.,
daily
average,
daily
maximum,
etc.)
they
were
associating
with
their
recommended
ranges.
Thus,
this
line
of
evidence
should
be
considered
with
caution.

Table
4.7.
A
summary
of
the
ranges
identified
in
the
literature
as
producing
optimal
growth
conditions
Species
Estimated
Optimal
Range
Comments
Author
Steelhead
and
Rainbow
Trout
14­
17
Estimate
for
the
late
Summer
regime
Behnke,
1992
10­
16
Growth
optimum
feeding
Behnke,
1992
5­
17
Olson
and
Templeton
13­
16
Dwyer
et
al.,
1981,
1983a,
1983b
12­
19
Recommended
7­
day
average
criteria
Taylor
and
Barton,
1992
13­
21
Moyle,
1976
12.2­
18.9
Bell,
1986
15­
20
Cho
and
Kaushik,
1990
14­
19
Optimal
temperature
lies
between
14­
19
Myrick
and
Cech,
2000
Cutthroat
Trout
12­
15
Raleigh,
1982
4.4­
15.5
Carlander,
1969
Summary
Upper
Range:
15­
21
(
18)

Comparison
Test
Regimes
with
Better
Growth:

The
results
in
the
table
4.8
below
should
only
be
used
to
generally
support
other
studies
since
they
did
not
determine
specific
temperatures
that
resulted
in
maximum
growth.
These
results
suggest,
however,
that
constant
temperatures
above
21
°
C
and
below
10
°
C
may
be
detrimental
to
growth,
and
that
constant
temperatures
in
the
range
of
12­
14.5
are
generally
favorable.
When
feeding
is
reduced
substantially,
however,
the
temperature
that
will
allow
for
similar
levels
of
growth
may
be
decreased
substantially
(
4­
5
°
C).
Thus
under
natural
feeding
regimes,
where
food
availability
is
more
restricted
during
the
high
temperature
months
of
summer,
maximum
growth
rates
would
be
more
likely
to
be
obtained
at
a
constant
temperatures
of
8­
10.5
°
C
(
assuming
a
4
°
C
reduction
due
to
restricted
feeding).
If
treated
as
either
a
weekly
average
or
summer
average
temperature
regime
this
constant
temperature
Page
58
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
range
would
be
converted
to
estimated
7­
DADMax
temperature
ranges
of
11.18­
13.68
°
C
(
12.43)
and
12.64­
15.14
°
C
(
13.89),
respectively.
This
line
of
evidence
when
viewed
on
its
own
merits
suggests
that
relatively
good
growth
will
occur
with
7DADMax
temperatures
of
11.18­
15.14
°
C
as
compared
with
temperatures
significantly
(
4
°
C
or
more)
warmer
or
colder.

Table
4.8.
Tests
Comparing
Growth
at
Only
Two
Contrasting
Temperatures:

Species
Better
growth
Worse
growth
Feeding
Ceased
Author
Chinook
13­
16
(
14.5)
21­
24
(
22.5)
Marine
and
Cech,
1998
12
8
Zaugg
and
Wagner,
1973
Sockeye
<
4
and
>
21,
poor
to
no
growth
Donaldson
and
Foster,
1941
4­
7
21
Brett,
1956
Steelhead
and
Rainbow
Trout
22­
25
Behnke,
1992
23
no
growth
Olson
and
Templeton,
1973
16
better
conversion
efficiency
10
Laboratory
Growth
Studies
 
Fluctuating
Temperatures:

Chinook
Salmon
Neilson
and
Green
(
1985)
found
that
in
comparing
fluctuating
to
constant
test
conditions,
that
growth
was
enhanced
through
naturally
cyclic
temperature
regimes,
suggesting
that
food
utilization
in
fluctuating
environments
may
be
higher
(
although
it
should
be
noted
that
this
is
not
a
consistent
finding
among
authors).
Marine
(
1997)
conducted
a
2.5
month
test
on
Sacramento
River
hatchery
stocks
under
satiation
rations
and
a
fluctuating
temperature
regime
(
13­
24
°
C)
that
generally
followed
the
seasonal
pattern
of
the
source
water.
Marine
concluded
that
his
work
suggests
the
upper
optimum
for
growth
is
about
18
°
C
(
approx.
range
17­
20
°
C).
Fish
reared
at
the
next
highest
range
(
21­
24
°
C),
however,
experienced
significantly
less
growth
than
those
reared
in
the
control
group
(
13­
16
°
C).
Hillman
(
1991)
conducted
tests
in
a
controlled
laboratory
stream
environment
with
chinook
salmon.
He
found
that
chinook
salmon
production
was
1.2
times
greater
in
cold
than
warm
water
(
i.
e.,
a
20%
reduction
in
weight
when
reared
at
18­
21
°
C
(
mean
19.5
°
C)
when
compared
with
12­
15
°
C
(
mean
13.5
°
C)
when
fed
twice
per
day
in
experimental
channels).
The
two
works
cited
above
suggest
that
average
temperatures
below
19
°
C
are
necessary
to
support
maximum
Evaluating
Standards
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Protecting
Aquatic
Life
in
Page
59
Washington's
Surface
Water
quality
Standards
growth
rates
in
chinook
salmon,
and
that
the
average
temperature
that
produces
maximum
growth
rates
likely
lies
between
15­
18
°
C
(
median
16.5
°
C).

Coho
Salmon
In
a
controlled
study
by
Everson
(
1973),
test
fish
were
subjected
to
different
fluctuating
temperature
regimes.
The
greatest
growth
occurred
in
coho
at
the
lowest
test
regime
of
12.1­
20.8
°
C
(
median
16.5
°
C).
Everson
also
found
that
juveniles
fed
moderate
rations
and
subjected
to
higher
fluctuating
test
temperatures
did
not
reach
sizes
typical
of
smolts
at
the
time
of
downstream
migration.

Sockeye
Salmon
Biette
and
Geen
(
1980,
as
cited
in
USEPA,
2001)
tested
growth
under
variable
rations
(
4­
6.9%)
and
fluctuating
temperature
regimes
similar
to
what
the
fish
voluntarily
experience
in
a
lake
(
fluctuating
two
times
per
day
from
5­
9
°
C
to
12­
18
°
C)(
approximate
mean
11
°
C).
In
this
work,
sockeye
grew
as
well
or
more
rapidly
under
the
natural
feeding
and
temperature
regime
than
at
constant
temperatures
from
6.2­
15.9
°
C
(
mean
11.05
°
C);
however,
at
satiation
rations
growth
was
better
under
the
constant
temperature
regimes.
Clarke
(
1978)
found
that
under­
yearling
sockeye
salmon
exposed
to
diel
thermocycles
are
able
to
acclimate
their
growth
to
a
temperature
above
the
mean
of
the
cycle.
Specific
growth
in
weight
on
the
7­
13
°
C
(
mean
10
°
C)
cycle
over
42
days
was
equivalent
to
that
on
a
constant
11.4
°
C,
and
on
the
5­
15
°
C
(
mean
10
°
C)
cycle
it
was
equivalent
to
a
constant
13.9
°
C.
Clarke
found
that
growth
was
greater
at
the
5­
15
°
C
cycle
than
at
a
7­
13
°
C
cycle.
In
constant
exposure
testing
on
maximum
rations,
growth
was
linear
over
the
range
of
7.5­
17.5
°
C.
The
work
of
Clarke
suggests
that
maximum
growth
would
be
produced
at
constant
temperatures
of
17.5
°
C.

Steelhead
and
Rainbow
Trout
Grabowski
(
1973)
tested
three
constant
temperatures
(
8,
15,
and
18
°
C)
and
one
fluctuating
temperature
regime
(
8­
18
°
C,
mean
13
°
C)
over
eight
weeks
with
juvenile
steelhead
on
maximum
rations.
The
author
found
that
steelhead
grew
best
at
the
constant
15
°
C
and
second
best
in
the
fluctuating
with
its
mean
of
13
°
C.
When
the
author
plotted
the
data
using
the
midpoint
of
the
fluctuating
test
as
a
surrogate
for
a
constant
test
condition,
it
showed
almost
linear
growth
from
8
to
15
°
C
with
a
steep
drop
as
the
temperature
approached
18
°
C.
The
percentage
weight
gains
at
8
and
18
°
C
were
very
similar
and
both
substantially
lower
than
the
similar
gains
obtained
at
the
mean
of
13
°
C
and
the
constant
15
°
C.

Wurtsbaugh
and
Davis
(
1977)
studied
the
effects
of
ration
size
and
temperature
on
the
growth
of
juvenile
steelhead
trout
through
a
series
of
laboratory
and
field
studies.
In
25­
day
laboratory
tests,
creek
water
was
used
as
a
control,
and
was
heated
3
and
6
°
C
above
its
natural
fluctuating
(
dielly
and
seasonally)
temperature
to
create
two
alternative
test
conditions.
From
this
work,
the
authors
concluded
that
trout
growth
would
be
enhanced
by
average
seasonal
rearing
temperature
increases
to
approximately
16.5
°
C
under
satiation
feeding,
but
if
the
food
of
wild
fish
is
limited,
any
substantial
temperature
increase
would
Page
60
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
result
in
decreased
growth.
By
examining
the
fish
fed
at
rates
more
comparable
to
those
observed
in
a
natural
creek,
the
maximum
growth
rates
observed
occurred
at
mean
temperatures
of
13.3
°
C
in
the
Fall
test
(
1.7%/
day
compared
to
1.3%/
day
at
16.4
°
C)
at
15.2
°
C
in
the
Spring
test
(
1.6%/
day
compared
to
1.4%/
day
at
12.6
°
C),
and
at
16.2
°
C
in
the
summer
test
(
1.2%/
day
compared
to
1.1%/
day
at
a
mean
of
19.5
°
C).
It
is
worthy
of
emphasizing
that
while
the
summer
growth
at
a
mean
of
19.5
°
C
(
range
of
16­
23.9
°
C)
was
only
slightly
less
than
growth
at
16.2
°
C
under
moderate
laboratory
rations
(
7.9
and
6.8%/
day,
respectively),
at
the
next
lowest
ration
tested
(
6.0%/
day)
growth
rates
at
19.5
°
C
fell
sharply
to
0.1%/
day.
The
authors
calculated
that
maintenance
requirements
of
the
fish
increased
more
than
three­
fold
(
2.2­
7.4%/
day)
over
the
temperature
range
of
6.9
to
22.5
°
C.

Dockray
et
al.
(
1996)
conducted
a
90­
day
test
in
which
temperatures
were
allowed
to
fluctuating
with
the
natural
regime
of
the
city
source
water,
which
followed
the
pattern
of
Lake
Ontario,
plus
2
°
C
to
test
the
potential
effects
of
global
warming.
The
authors
found
that
during
the
first
50
days
of
the
90­
day
test
that
the
temperature
increase
of
2
°
C
was
beneficial
to
growth
up
to
daily
average
of
18
°
C,
after
which
further
increases
in
the
daily
temperature
inhibited
long
term
growth.
In
a
follow­
up
study,
Linton
et
al.
(
1998)
noted
that
rainbow
trout
fed
to
satiation
continued
to
feed
and
grow
at
a
mean
temperature
of
20.5
°
C,
that
a
30%
reduction
in
food
intake
occurred
at
22
°
C,
and
that
juvenile
fish
continued
to
feed
near
their
thermal
maximum.
Linton
et
al.
(
1998)
found
that
increasing
the
temperature
regime
by
2
°
C
over
the
natural
(
base)
level
for
Lake
Ontario
trout
resulted
in
increased
spring
and
early
summer
growth
that
was
lost
in
the
latter
part
of
the
summer
due
to
suppression
of
appetite
and
growth.
Mortality
rates
increased
from
6
to
13.1%
in
the
warmer
test
water
during
the
late
summer
in
the
first
summer
of
testing
when
the
mean
monthly
base
temperature
in
August
was
23
°
C.
Mortality
was
almost
nonexistent
through
the
following
summer
which
had
a
mean
August
base
temperature
of
18
°
C.
The
threshold
temperature
for
the
cessation
of
feeding,
and
subsequently
growth,
differed
from
>
20
°
C
to
<
20
°
C
over
the
two
summers,
and
thus
also
fish
size
and
age.
Hokanson
et
al.
(
1997)
found
that
a
constant
exposure
to
17.2
°
C
produced
the
greatest
growth
rates
in
trout
fed
to
satiation
over
a
30­
day
test
period.
Increased
mortality
was
observed
at
temperatures
in
excess
of
this
growth
optimum.
They
also
noted
that
in
fluctuating
temperature
experiments
that
growth
was
accelerated
where
the
mean
temperature
is
below
the
constant
temperature
optimum
(
17.2
°
C),
and
growth
was
retarded
by
mean
fluctuating
temperatures
above
this
optimum.
The
highest
growth
rate
in
the
fluctuating
temperature
environment
occurred
at
a
mean
of
15.5
°
C
(
range
of
11.7­
19.3
°
C).
A
statistically
non­
significant
decrease
occurred
at
a
mean
of
17.3
°
C
(
range
of
13.5­
21.1
°
C).
Through
their
work
the
authors
also
concluded
that
rainbow
trout
acclimate
to
some
value
between
the
mean
and
the
maximum
daily
temperatures.
The
foregoing
works
suggest
that
maximum
growth
will
be
obtained
at
average
daily
temperatures
between
15.5
and
18
°
C.

Cutthroat
Trout
Dickerson
et
al.
(
1999,
and
unpub.
data;
as
cited
in
Dunham,
1999)
found
that
Lahontan
cutthroat
growth
rates
in
a
test
with
temperatures
fluctuating
from
20­
26
°
C
(
mean
of
23
°
C)
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
61
Washington's
Surface
Water
quality
Standards
were
similar
to
groups
of
fish
held
at
a
constant
23
°
C,
and
after
two
weeks
was
not
significantly
different
from
fish
held
at
a
constant
20
°
C.

Table
4.9.
Growth
studies
conducted
in
laboratory
test
waters
with
fluctuating
temperature
exposures.

Species
Temperature
(
range
and
mean)
Results
Feeding
level
Comments
Author
Chinook
13­
16
(
14.5)
17­
20
(
18.5)
21­
24
(
22.5)
Optimal
17­
20,
with
13­
16
having
significantl
y
better
growth
than
21­
24
Satiation
2.5
month
test
following
source
water
regime
Marine,
1997
12­
15
(
13.5)
18­
21
(
19.5)
12­
15
significantl
y
better
than
18­
21
Satiation
18­
21
had
20%
less
growth
and
12­
15
Hillman,
1991
Coho
12.1­
20.8
(
16.5)
Greater
growth
than
higher
regimes
tested
Satiation
Juveniles
fed
moderate
rations
did
not
reach
typical
size
of
smolts
Everson,
1973
Sockeye
5­
9
to
12­
18
(
approx
11)
Variable
from
4
to
6.9%
body
weight
Grew
more
rapidly
under
natural
feeding
regime
and
fluctuation
temperatures
than
at
constant
11.05C,
but
constant
better
under
satiation
feeding.
Biette
and
Geen,
1980
Cutthroat
7­
13(
10)
5­
15(
10)
5­
15(
10)
had
greater
growth
than
7­
13(
10)
Satiation
Clarke,
1978
Steelhea
d
and
Rainbow
8­
18
(
13)
Better
growth
in
a
constant
15
Satiation
Weight
gains
similar
at
8
and
18
and
both
Grabowski,
1973
Page
62
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Trout
than
the
fluctuating
8­
18
(
13)
substantially
lower
than
at
13
and
15.
16.5
average
seasonal
temp
considered
to
enhance
growth
Satiation
(
6.8%/
day
)
Heated
creek
water
(
3
and
6C)
followed
natural
regime
for
25
days
Wurtsbaugh
and
Davis,
1977
16.2
summer
average
seasonal
considered
to
enhance
growth
under
natural
feeding
Restricted
to
rates
seen
in
control
stream
(
1.2%/
day
)
Heated
creek
water
(
3
and
6C)
followed
natural
regime
for
25
days
Wurtsbaugh
and
Davis,
1977
Increases
beneficial
up
to
daily
average
of
18
Satiation
Fluctuated
with
source
water
plus
2C
over
90
days.
Dockray
et
al.,
1996
11.7­
19.3
(
15.5)
13­
21.1
(
17.3)
Highest
growth
rate
in
11.7­
19.3
(
15.5),
with
nonsignificant
decrease
in
13­
21.1
(
17.3)
60
day
test
Hokanson
et
al.,
1997
The
results
of
these
fluctuating
temperature
tests
is
highly
variable
due
at
least
in
part
to
the
high
variability
in
test
methodology.
Better
growth
conditions
varied
from
regular
daily
ranges
of
5­
15
°
C
to
11.7­
19.3
°
C
over
the
course
of
the
studies
(
typically
month­
long
studies),
to
a
monthly
(
described
as
seasonal)
average
of
16.5
°
C,
and
a
highest
daily
mean
of
18
°
C
in
a
seasonally
(
90­
day)
rising
temperature
regime.
This
could
be
simplified
and
viewed
as
a
range
of
10­
16.5
°
C
as
a
monthly
average
and
a
maximum
daily
average
of
18
°
C.
Treating
the
monthly
average
as
either
a
weekly
average
or
a
summer
average,
to
bound
the
potential
best
range,
would
result
in
a
7DADMax
of
13.16­
19.68
°
C
(
16.42)
and
14.64­
21.14
°
C
(
17.89),
respectively.
Thus
the
7DADMax
temperature
that
would
result
in
maximum
growth
would
most
likely
occur
between
13.16­
21.14
°
C
(
17.15).
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
63
Washington's
Surface
Water
quality
Standards
Field
Studies
on
Growth:

Chinook
Salmon
In
studying
growth
in
a
natural
stream,
Bisson
and
Davis
(
1976;
as
cited
in
ODFW,
1992)
reported
that
juvenile
chinook
grew
faster
in
a
stream
where
temperatures
peaked
at
16
°
C
compared
with
a
stream
where
temperatures
peaked
at
20
°
C.

Coho
Salmon
Averett
(
1969;
as
cited
by
Everson,
1973)
proposed
that
August­
September
stream
temperatures
should
fluctuate
between
11­
17
°
C
(
mean
14
°
C)
for
optimal
growth.
Holtby
(
1988)
reviewing
a
long
term
study
of
coho
in
an
experimentally
harvested
watershed
determined
that
coho
had
likely
benefited
from
changes
in
maximum
stream
temperature
from
12
°
C
to
15
°
C
(
which
would
be
approximately
equal
to
a
maximum
weekly
average
of
14.56
°
C).
Similarly,
Thedinga
and
Koski
(
1984)
found
that
smolt
size
and
condition
factor
was
greater
in
years
in
which
stream
temperatures
fluctuating
annually
from
4­
13.5
°
C
than
in
years
with
temperatures
of
near
0
to
11­
12
°
C.
They
associated
greater
smolt
return
rates
with
this
greater
growth.
In
a
field
study
by
Martin
et
al.
(
1984)
the
stream
having
the
lowest
growth
was
the
warmest
surveyed,
and
had
average
monthly
temperatures
of
12­
17
°
C
and
peak
monthly
temperatures
of
21­
26
through
the
summer
months.
The
foregoing
does
not
provide
a
basis
for
estimating
a
maximum
growth
temperature,
but
can
be
used
to
support
the
position
that
weekly
average
temperatures
of
14­
15
°
C
would
be
more
beneficial
to
growth
under
natural
conditions
than
lower
temperature
regimes.
It
also
suggests
that
extreme
daily
fluctuations
and
daily
maximum
temperatures
of
21­
26
°
C
can
be
detrimental
to
growth
even
under
natural
conditions.

Steelhead
and
Rainbow
Trout
De
Leeuw
(
1982)
found
that
stream
temperature
increases
that
raised
the
summer
maximum
temperature
from
12
°
C
to
16.5
°
C
were
associated
with
an
increase
in
growth
rates
in
three
streams
in
British
Columbia,
Canada.

Bisson
and
Davis
(
1976;
as
cited
in
Li
et
al.,
1994)
found
that
streams
with
daily
maximum
temperatures
in
the
range
of
16­
23
°
C
had
greater
standing
crops
of
trout
than
streams
with
warmer
maximum
temperatures
(
26­
31
°
C).
Ebersole
et
al.
(
2001)
found
that
in
tributaries
to
the
Grande
Rhonde
River
in
Oregon,
rainbow
trout
density
declined
as
mean
daily
maximum
temperatures
(
for
the
two
week
period
prior
to
survey)
warmed
above
16
°
C;
reaching
zero
at
approximately
24
°
C.
While
the
decline
in
fish
density
was
highly
variable
in
the
range
of
16
to
20
°
C,
it
was
a
steady
decline
with
very
low
densities
at
temperatures
above
20
°
C.
Page
64
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Cutthroat
Trout
Temperature
increases
in
general
are
not
always
found
to
restrict
cutthroat
populations.
Aho
(
1976)
found
coastal
cutthroat
preferring
an
unshaded
section
of
stream
in
the
Cascade
mountains
of
Oregon.
Density
was
twice
as
high
and
biomass
was
49­
65%
greater
in
the
unshaded
section.
He
suggested
that
earlier
fry
emergence
in
the
warmer
unshaded
section
probably
played
a
role
in
creating
the
greater
fish
weights.
The
highest
weekly
mean
temperature
was
14
°
C
in
the
unshaded
area,
with
the
highest
one
hour
temperature
being
17
°
C.
This
compared
to
a
daily
maximum
temperature
of
14
°
C
in
the
shaded
section.
Cutthroat
were
the
only
fish
species
present
in
the
study
stream
so
any
relationship
to
potential
competition
could
not
be
evaluated.
Martin
(
1985)
studied
the
affect
of
removing
the
vegetative
canopy
along
a
1000
meter
section
of
a
third
order
stream
on
the
Olympic
Peninsula
used
only
by
cutthroat
trout.
Fish
density
increased
in
the
upper
500
meter
treatment
reach
in
comparison
to
the
200
meter
control
section
established
just
above
the
treatment
reach.
Summer
daily
maximum
temperatures
in
this
upper
treatment
ranged
from
13.8
°
C
at
the
boundary
with
the
control
section
to
about
16.1
°
C
at
the
bottom
of
this
upper
treatment
section
500
meters
downstream.
In
the
second
treatment
reach,
which
extended
from
500
to
1000
meters
below
the
control
section,
the
average
fish
weight
increased
but
density
went
down.
Temperatures
in
this
reach
changed
from
16.1
°
C
at
the
upper
boundary
of
the
treatment
reach
to
17.3
°
C
at
the
lower
boundary
of
the
lower
treatment
reach.
The
daily
maximum
temperature
was
15.2
°
C
in
the
midpoint
of
the
upper
treatment
reach
where
fish
density
increased.

Martin
(
1984)
in
studying
a
small
upper
watershed
creek
on
the
Olympic
Peninsula
found
that
higher
water
temperatures
during
the
summer
caused
metabolism
and
thus
food
consumption
to
increase
in
a
population
of
cutthroat
trout.
He
found
that
growth
rates
declined
from
a
spring
high
to
a
low
in
the
winter,
and
was
below
the
maximum
possible
during
periods
of
optimum
water
temperature.
Food
consumption
followed
the
seasonal
trend
in
food
abundance
and
was
limited
by
the
available
food
supply.

Table
4.10.
Summary
results
from
growth
studies
conducted
in
the
field.

Species
Better
growth
Worse
growth
Author
Chinook
16
(
daily
max)
20
Bisson
and
Davis,
1976
Coho
15
(
daily
max)
12
Holtby,
1988
4­
13.5
(
annual
ranges)
0­
12
Size
and
condition
factor
Thedinga
and
Koski,
1984
12­
17
(
average
monthly)
Lowest
growth
stream
Had
peak
temperatures
of
21­
26
Martin
et
al.,
1984
Steelhead
and
Rainbow
Trout
16.5
(
daily
max)
12
De
Leeuw,
1982
16­
23
(
daily
max)
26­
31
Greater
standing
crops
Bisson
and
Davis,
1976
Greater
than
16
Density
declined
Ebersole
et
al.,
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
65
Washington's
Surface
Water
quality
Standards
(
mean
daily)
above
16
and
was
zero
at
24.
Steady
decline
above
20
2001
Cutthroat
14
(
weekly
mean)
17
(
daily
max)
Site
with
highest
weekly
mean
of
14
had
twice
the
biomas.
Highest
one
hour
temperature
was
17.
Aho,
1976
15.2
(
difference
along
reach
13.8­
16.1)
(
daily
max)
16.7(
difference
along
reach
16.1­
17.3)
Density
increased
in
reach
at
15.2,
density
declined
but
average
size
increased
at
16.7.
Martin,
1985
In
the
field
studies
cited
above,
daily
maximum
stream
temperatures
from
15­
17
°
C
were
consistently
cited
in
association
with
streams
having
better
standing
crops
of
fish.
This
temperature
range
would
be
approximately
equal
to
a
7DADMax
range
of
14.05­
16.05
°
C.
It
is
important
to
recognize
that
most
of
these
studies
did
not
evaluate
stream
temperatures
just
one
or
two
degrees
higher
than
those
found
to
have
better
growth.
Thus
the
conclusion
is
somewhat
restricted
to
the
statement
that
under
natural
feeding
and
temperature
regimes
salmonid
populations
will
be
fully
supported
at
7DADMax
temperatures
from
14­
16
°
C.

Predation
and
Competition
Involving
Juvenile
Fish:

Chinook
Salmon
In
examining
the
effect
of
temperature
upon
predation
rates
of
juvenile
chinook,
Marine
(
1997)
found
that
striped
bass
predation
upon
the
juvenile
fish
was
very
intense
in
their
intermediate
test
at
17­
20
°
C
(
mean
18.5
°
C).
Hillman
(
1991)
examined
the
interaction
between
chinook
salmon
and
redside
shiner
in
both
the
Wenatchee
River
in
Washington
and
in
controlled
laboratory
streams.
Immigration
of
shiners
began
as
river
temperatures
reached
15
°
C
and
as
their
numbers
increased
the
shiners
were
able
to
push
the
chinook
out
of
the
most
favorable
portions
of
the
river
for
feeding.
Similarly,
in
the
laboratory
tests
Hillman
found
that
chinook
production
was
1.2
times
greater
in
cold
water
(
12­
15
°
C,
mean
13.5
°
C)
than
warm
water
(
18­
21
°
C,
mean
19.5
°
C
 
resulted
in
a
20%
reduction
in
weight)
in
alloparty.
In
sympatry,
shiners
affected
the
distribution,
activity,
and
production
of
chinook
in
warm
water,
but
not
in
cold
water.
Based
on
the
preceding,
average
temperatures
in
the
range
of
13.5­
15
°
C
favors
the
cold
water
chinook
salmon
over
common
non­
salmonid
competitors.
Average
temperatures
of
18.5­
19.5
°
C
would
place
the
chinook
salmon
at
a
significant
disadvantage
for
occupying
and
defending
important
habitat.

Steelhead
and
Rainbow
Trout
Page
66
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Juvenile
competition,
particularly
with
warm
water
and
non­
salmonid
species,
is
a
concern
that
is
sometimes
expressed
in
association
with
warm
summer
rearing
temperatures.
Reeves
et
al.
(
1987)
found
that
steelhead
production
was
decreased
by
54%
in
the
presence
of
redside
shiner
in
waters
in
the
range
of
19­
22
°
C
(
mean
20.1
°
C),
but
was
not
reduced
in
cooler
waters
(
12­
15
°
C,
mean
13.5
°
C).
They
also
noted
that
production
of
steelhead
in
a
fluctuating
19­
22
°
C
waterway
was
less
than
half
of
that
in
a
fluctuating
12­
15
°
C
waterway.
While
warmer
waters
favor
redside
shiner
over
steelhead,
colder
waters
may
favor
other
species.
It
has
been
suggested,
for
example,
that
in
cooler
waters
brook
trout
can
outcompete
juvenile
steelhead.
In
trying
to
explain
the
patterns
of
species
distributions
observed
in
the
upper
Columbia
river
tributaries,
Mullan
et
al.
(
1992;
using
the
work
of
Cherry
et
al.,
1975)
note
that
sympatry
occurs
between
O.
mykiss
and
brook
trout
in
the
range
of
15­
18
°
C
(
mean
16.5
°
C),
that
temperatures
above
18
°
C
favors
O.
mykiss,
and
that
temperatures
less
than
15
°
C
favors
brook
trout.
They
also
note,
however,
that
under
natural
food
rations,
the
level
of
optimal
competition
is
likely
to
be
lower.
Cunjak
and
Green
(
1986)
found
that
rainbow
trout
were
able
to
compete
better
with
brook
trout
at
19
°
C
than
at
either
8
or
13
°
C.
In
tests
of
the
interaction
between
rainbow
trout
and
bull
trout
in
the
Columbia
River
drainage
in
British
Columbia,
Haas
(
2001)
found
that
rainbow
trout
do
particularly
better
than
bull
trout
when
maximum
temperatures
approach
14­
15
°
C.
Trying
to
set
temperature
threshold
recommendations
that
will
prevent
a
competitive
advantage
with
an
introduced
coldwater
salmonid,
however,
is
not
viewed
as
appropriate
given
the
extensive
overlap
of
their
preferred
thermal
ranges.
Based
on
the
preceding,
it
appears
that
average
temperatures
of
20
°
C
or
warmer
place
steelhead
at
a
competitive
disadvantage
over
warm
water
competitors
like
redside
shiner,
while
average
temperature
of
13.5
°
C
or
less
do
not.

Cutthroat
Trout
DeStaso
and
Rahel
(
1994)
found
that
Colorado
cutthroat
trout
competed
nearly
equally
with
brook
trout
at
10
°
C,
but
at
20
°
C
the
brook
trout
were
dominant.
Schroeter
(
1988;
as
cited
in
Dunham,
1999)
found
that
Lahontan
cutthroat
were
equal
competitors
with
brook
trout
at
15
°
C.
Nilsson
and
Northcote
(
1981)
found
interestingly
enough
that
in
sympatry
the
presence
of
more
aggressive
rainbow
trout
actually
resulted
in
higher
growth
rates
in
cutthroat
trout
than
the
growth
rates
shown
by
the
cutthroat
in
allopatry.

Summary
on
Competition
and
Predation
Trying
to
set
temperature
standards
that
will
prevent
competition
with
an
introduced
cold
water
species
is
not
viewed
as
appropriate
give
the
extensive
overlap
of
their
preferred
thermal
ranges
with
Washington's
native
cold
water
species.
It
is
more
appropriate
to
focus
this
discussion
on
temperatures
that
allow
coolwater
or
warmwater
species
to
out­
compete
our
native
salmon
and
trout.
Table
4.11
below
summarizes
the
studies
on
competition
and
predation.
The
literature
suggests
that
as
average
river
temperatures
increase
to
about
15
°
C
competitors
begin
to
gain
a
competitive
edge
that
becomes
very
significant
as
the
average
temperature
reaches
18.5­
20
°
C.
In
converting
these
average
ranges
they
may
best
represent
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
67
Washington's
Surface
Water
quality
Standards
either
daily
average
or
weekly
average
temperatures.
This
results
in
an
estimate
that
as
the
7DADMax
temperature
exceeds
17.60­
18.18
°
C
(
17.89)
coolwater
species
will
begin
to
displace
juvenile
salmon
and
trout
from
the
best
feeding
habitats,
and
that
the
impact
on
growth
would
be
severe
at
22.05­
23.18
°
C
(
22.62)
even
if
such
displacement
did
not
occur.

Table
4.11.
Studies
on
predation
and
competition
with
salmon
and
trout.

Species
Increased
predation
or
competition
Type
of
Study
Author
Chinook
17­
20
(
18.5)
Laboratory
Intense
increase
in
predation
from
striped
bass
Marine,
1997
15
(
average)
Field
Redside
shiner
immigrated
and
began
to
push
out
juvenile
salmon
Hillman,
1991
18­
21
(
19.5)
Laboratory
Redside
shiner
caused
a
20%
reduction
in
weight
compared
to
competition
at
12­
15
(
13.5)
Hillman,
1991
Steelhead
and
Rainbow
Trout
19­
22
(
20.1)
Laboratory
Redside
shiner
caused
a
54%
reduction
in
production
compared
to
competition
at
12­
15(
13.5)
Reeves
et
al.,
1987
Less
than
15
favors
brook
trout
Sympatry
with
brook
trout
15­
18
under
satiation
rations.
Mullan
et
al.,
1992
8­
13
Brook
trout
are
better
able
to
compete
Cunjak
and
Green,
1986
Below
14
(
maximum
temperatures)
Bull
Trout
Haas,
2001
Cutthroat
Trout
20
and
above
Colorado
cutthroat
are
in
sympatry
with
Brook
trout
at
10
DeStaso
and
Rahel,
1994
Lahontan
cutthroat
equal
with
brook
trout
at
15
Schroeter,
1998
Temperature
Preferences
in
the
Laboratory:
Page
68
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Chinook
Salmon
Brett
(
1951;
as
cited
in
Ferguson,
1958)
reportedly
found
that
the
final
preferendum
temperature
for
chinook
is
11.7
°
C.
Sauter
and
Maule
(
2000)
found
that
the
mean
temperature
preference
of
fall
chinook
decreased
from
a
high
of
17.77
to
a
low
of
11.12
°
C
during
smoltification,
while
yearling
spring
chinook
salmon
showed
no
change
in
their
preference
for
a
mean
temperature
of
16.69
°
C.

Coho
Salmon
Piper
et
al.
(
1982)
suggested
that
maximum
temperatures
between
9.4­
14.4
°
C
are
optimal
or
selectively
preferred
by
juvenile
coho;
however,
Bell
(
1986)
suggested
the
preferred
range
for
coho
is
4.4­
9.4
°
C.
Konnecki,
Woody
and
Quinn
(
1995)
studied
two
groups
of
coho
and
found
that
the
preference
temperature
changed
with
the
parental
stock
used.
Coho
from
parental
stock
originating
from
cold
ground
water
supplied
streams
preferred
9.6
°
C
(
range
6­
16
°
C),
while
those
from
stock
originating
from
warmer
streams
preferred
11.6
°
C
(
range
7­
21
°
C).

Chum
Salmon
Juvenile
chum
salmon
reportedly
prefer
temperatures
between
11­
14.6
°
C,
and
are
considered
to
be
optimally
maintained
at
13­
13.5
°
C
(
Kepshire,
1971;
as
cited
in
Brett,
1979).
Ferguson
(
1958)
suggests
that
the
final
temperature
preferendum
for
chum
salmon
is
14.1
°
C.
In
a
1952
study,
Brett
found
that
juvenile
sockeye
avoided
temperatures
above
15
°
C
and
selected
12­
14
°
C
when
provided
an
opportunity.

Pink
Salmon
Bell
(
1986)
and
Bonar
et
al.
(
1989)
suggest
pink
salmon
prefer
waters
in
the
range
of
about
5.6­
14.5
°
C,
while
Ferguson
sets
the
final
preferendum
at
11.7
°
C
and
Brett
(
1952)
determined
pink
salmon
acclimated
to
temperatures
below
15
°
C
select
temperatures
in
the
range
of
12­
14
°
C.

Sockeye
Salmon
Ferguson
(
1958)
used
earlier
work
by
Brett
(
1951)
to
determine
that
the
final
preferendum
for
sockeye
was
14.5
°
C.
Brett
(
1971;
as
cited
in
USEPA,
2001)
found
swimming
capacity,
metabolic
scope,
growth
on
excess
rations,
and
ingestion
were
maximized
at
15
°
C,
and
that
15
°
C
was
also
the
final
preferendum
temperature.

Steelhead
and
Rainbow
Trout
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
69
Washington's
Surface
Water
quality
Standards
Hahn
(
1977)
compared
the
preference
decisions
of
fry
and
yearling
steelhead
exposed
to
three
constant
(
8.5
°
C,
13.5
°
C,
and
18.5
°
C)
and
one
fluctuating
(
8­
19
°
C)
temperature
regime.
Hahn
found
that
as
many
fish
remained
in
the
fluctuating
regime
(
which
has
a
mean
of
13.5
°
C)
as
in
the
constant
13.5
°
C
regime;
twice
as
many
fish
remained
in
the
fluctuating
regime
compared
with
the
constant
18.5
°
C;
and
twice
as
many
remained
in
the
constant
8.5
°
C
as
in
the
fluctuating
regime.
By
inference,
Hahn
concluded
that
twice
as
many
fish
preferred
a
constant
13.5
°
C
to
a
constant
18.5
°
C,
and
twice
as
many
preferred
a
constant
8.5
°
C
to
a
constant
13.5
°
C.
Hahn's
work
suggests
that
juvenile
steelhead
may
have
a
preference
for
water
temperatures
between
8.5­
13.5
°
C.
It
also
suggests
that
the
daily
average
temperature
may
roughly
equal
a
constant
exposure
test
scenario
for
the
purpose
of
translating
laboratory
tests
into
water
quality
criteria.

Ferguson
(
1958)
cites
13.6
°
C
as
the
final
preferendum
temperature
for
rainbow
trout.
The
work
of
Li
et
al.
(
1993;
and
1994)
and
Li
et
al.
(
1991;
as
cited
in
Spence
et
al.,
1996),
studying
interior
populations
of
rainbow
trout,
suggest
that
while
rainbow
trout
showed
no
avoidance
reactions
when
stream
temperatures
were
below
20
°
C,
they
actively
avoided
staying
in
waters
warmer
than
23­
25
°
C.
McCauley
and
Huggins
(
1975)
found
that
five
large
(
150­
250
grams)
rainbow
trout
had
a
preferred
mean
temperature
of
16.7
°
C,
and
that
the
fish
actively
cycled
between
13.8
°
C
and
18
°
C.
The
work
and
literature
citations
of
Kwain
and
McCauley
(
1978)
(
citing
Huggins
1978
Garside
and
Tait,
1958,
Christie
as
reported
in
Fry,
1971,
and
McCauley
et
al.,
1977)
support
the
position
that
rainbow
trout
temperature
final
preferendum
decline
from
17­
21
°
C
in
first
six
months
of
life
to
eventually
centering
on
13
°
C
as
overyearlings.
McCauley
and
Pond
(
1971)
found
that
underyearling
rainbow
trout
showed
a
preference
for
17­
20
°
C
in
laboratory
tests.

Summary
of
Laboratory
Temperature
Preference
Studies
The
results
of
preference
testing
depends
upon
acclimation
temperature,
length
of
test,
breadth
of
temperature
gradients
used,
and
age
of
fish.
Fish
will
actively
cycle
between
temperature
regimes
during
the
test
period,
which
is
typically
one
hour
in
length,
and
their
time
spent
in
different
temperature
zones
is
used
to
determine
where
they
spent
most
of
their
time
(
preference
is
in
essence
the
mode
of
the
distribution).
While
the
range
of
temperatures
through
which
fish
commonly
cycled
during
the
tests
extended
from
4.4­
21
°
C,
preferred
temperatures
ranged
from
9.6­
17.7
°
C.
The
majority
of
the
preferred
temperatures
were
from
11.6­
16.7
°
C
(
see
Table
4.12,
below).
These
preferred
tests
would
best
relate
to
a
daily
mean
temperature.
A
daily
mean
of
11.6­
16.7
°
C
would
be
roughly
equivalent
to
a
7DADMax
of
14.2­
19.3
°
C
(
16.75).

Table
4.12.
Results
of
laboratory
studies
examining
temperature
preferences.

Species
Preferred
Preferred
Comments
Author
Page
70
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
temperature
Range
Chinook
11.7
Final
preferendum
Brett,
1951
17.7
to
11.12
Fall
Chinook
decrease
in
preference
during
smoltification
Sauter
and
Maule,
2000
16.69
Spring
chinook
Sauter
and
Maule,
2000
Coho
4.4­
9.4
Bell,
1986
9.4­
14.4
9
Piper
et
al.,
1982
9.6
6­
16
Parental
stock
from
colder
water
Konnecki
et
al.,
1995
11.6
7­
21
Parental
stock
from
warmer
water
Konnecki
et
al.,
1995
Chum
11­
14.6
Kepshire,
1971
14.1
Ferguson,
1958
12­
14
Brett,
1952
Pink
5.6­
14.5
Bell,
1986
and
Bonar
et
al.,
1989
11.7
Ferguson,
1958
12­
14
Acclimated
below
15
Brett,
1952
Sockeye
14.5
Used
work
of
Brett,
1951
Ferguson,
1958
15
Final
preferendum
Brett,
1971
Steelhead
and
rainbow
trout
8.5­
13.5
Hahn,
1977
13.6
Final
preferendum
Ferguson,
1958
Actively
avoided
23­
25
Li
et
al.,
1991,
1993,
and
1994
16.7
13.8­
18
Cycled
between
13.8­
18
McCauley
and
Huggins,
1975
17­
21
First
six
months
of
life.
Kwain
and
McCauley,
1978
13
Overyearlings
Kwain
and
McCauley,
1978
17­
20
Underyearlings
McCauley
and
Pond,
1971
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
71
Washington's
Surface
Water
quality
Standards
Swimming
Performance
and
Scope
of
Activity:

Coho
Salmon
In
tests
on
swimming
performance,
Griffiths
and
Alderdice
(
1972)
and
Brett
et
al.(
1958)
found
that
optimum
swimming
performance
in
juvenile
coho
salmon
occurred
at
a
combination
of
acclimation
and
test
temperatures
near
20
°
C.
Griffiths
and
Alderdice
reported
that
above
20
°
C
swimming
performance
experienced
a
marked
reduction.
Delacy
et
al.
(
1956)
and
Paulik
et
al.
(
1957)
also
studied
the
swimming
performance
of
coho
salmon.
They
found
that
coho
could
recover
to
approximately
to
31%
after
a
one­
hour
rest
and
to
70%
within
three
hours
of
exhaustive
swimming
effort,
and
recover
fully
overnight.
They
also
noted
that
eggs
taken
from
coho
salmon
repeatedly
fatigued
in
their
testing
exhibited
normal
fertility
and
survival.
A
significant
swimming
performance
decrement
was
associated
with
pre­
test
activity
of
the
salmon,
and
salmon
that
swam
longer
before
fatigue
required
more
recuperation
time;
suggesting
that
the
salmon
are
highly
susceptible
to
fatigue.
Beamish
(
1978;
as
cited
in
USEPA,
2001),
in
a
literature
review
indicated
that
coho
reach
a
maximum
speed
at
approximately
17­
18
°
C.

Sockeye
Salmon
Brett
(
1971;
as
cited
in
USEPA,
2001)
found
swimming
capacity,
metabolic
scope,
growth
on
excess
rations,
and
ingestion
were
maximized
at
15
°
C,
and
that
15
°
C
was
also
the
final
preferendum
temperature.
In
tests
on
physiological
performance,
juvenile
sockeye
reach
their
maximum
swimming
speed
at
15
°
C
(
Brett
and
Glass,
1973;
Brett
et
al.,
1958).
Brett
et
al.
(
1958)
noted
that
the
capacity
of
young
sockeye
to
stem
a
normal
river
current
of
1.0
ft/
sec.
for
more
than
an
hour
is
limited
to
a
relatively
small
temperature
range
of
12.5­
17.5
°
C.
Beamish
(
1979;
as
cited
in
USEPA,
2001)
found
that
sockeye
reached
maximum
speed
at
15
°
C.

Mcdonald
et
al.
(
2000)
used
field
studies
to
conclude
that
optimal
swimming
performance
occurs
at
17
°
C
and
that
a
20%
reduction
occurs
at
21
°
C.
Brett
(
1983)
in
discussing
the
energetic
needs
of
sockeye
suggested
that
the
protected
environment
of
a
growth­
metabolism
tank
or
hatchery
pond
may
not
correspond
well
with
the
search­
attack­
avoid­
escape
patterns
of
real
life.
An
attack
or
escape
episode
involving
20
second
burst
speed
was
shown,
in
terms
of
energy
expended,
to
be
equal
to
that
of
15
minutes
of
active
metabolism
(
maximum
sustained
swimming
speed)
or
about
3
hours
of
basal
metabolism
(
cites
Brett
and
Groves,
1979).
Brett
suggests
the
additional
energy
requirements
of
real
life
cannot
be
disregarded;
nor
can
it
be
conceived
as
incessantly
present.
Hinch
and
Bratty
(
2000)
found
that
Fraser
River
sockeye
used
a
"
burst
and
then
sustained"
swimming
pattern
to
negotiate
obstacles,
and
that
fish
that
maintained
the
"
burst"
phase
too
long
failed
to
negotiate
significant
obstacles
and
often
dropped
back
downstream
failing
to
complete
their
migration.
This
work
showed
two
important
facts,
the
first
being
that
sockeye
may
be
near
the
limits
of
their
performance
while
migrating
through
rapids
and
that
fatigue
can
result
in
the
permanent
loss
of
the
migrants,
and
that
fish
may
use
techniques
to
perform
better
in
the
field
than
in
laboratory
experiments.
That
such
stressors
may
be
additionally
selective
against
individuals
Page
72
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
is
also
likely.
The
work
by
Hinch
and
Rand
(
1998)
and
others
has
shown
that
males
suffer
more
swimming
impairment
then
females
and
that
small
fish
must
expend
more
energy
than
larger
fish.

Steelhead
and
Rainbow
Trout
Cech
and
Myrick
(
1999)
found
that
winter­
run
steelhead
swimming
velocities
increased
slightly
with
an
increase
in
temperature
from
11
°
C
to
15
°
C
and
decreased
slightly
with
an
increase
in
temperature
from
15
°
C
to
19
°
C.
Dickson
and
Kramer
(
1971)
found
that
the
maximum
scope
of
activity
occurred
at
20
°
C
in
rainbow
trout.

Cutthroat
Trout
Dwyer,
and
Kramer
(
1975;
as
cited
in
Dunham,
1999;
and
Gresswell,
1995)
found
the
greatest
scope
for
activity
in
cutthroat
trout
to
be
at
15
°
C.

Table
4.13.
Studies
examining
swimming
speed
and
maximum
scope
for
activity.

Species
Maximum
performance
Prolonged
performance
Author
Coho
20
A
marked
reduction
occurred
above
20
Griffiths
and
Alderdice,
1972
20
Brett
et
al.,
1958
17­
18
Literature
review
Beamish,
1978
Sockeye
15
Brett,
1971
15
Brett
and
Glass,
1973
15
Brett
et
al.,
1958
12.5­
17.5
Capacity
to
stem
normal
current
for
more
than
one
hour
limited
to
range
of
12.5­
17.5
Brett
et
al.,
1958
15
Beamish,
1979
17
Based
on
field
studies.
20%
reduction
at
21
Mcdonald,
2000
Steelhead
and
15
Cech
and
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
73
Washington's
Surface
Water
quality
Standards
rainbow
trout
Myrick,
1999
20
Maximum
scope
for
activity
Dickson
and
Kramer,
1971
Cutthroat
15
Maximum
scope
for
activity
Dwyer
and
Kramer,
1975,

15
Maximum
scope
for
activity
Gresswell,
1995
Maximum
swimming
speed
and
maximum
scope
for
activity
have
been
noted
at
temperatures
ranging
from
15­
20
°
C
(
17.5)
in
salmon
and
trout.
The
short­
term
tests
that
were
used
to
base
most
of
these
estimates
on,
and
the
real
time
effects
of
temperature
(
the
effect
of
temperature
is
essentially
immediate
on
swimming
performance)
suggests
these
estimates
may
best
transfer
to
some
daily
maximum
metric.
The
biological
consequences
of
having
a
shortperiod
of
each
day
not
allowing
for
maximal
performance
is
difficult
to
ascertain.
However,
the
works
of
Brett
(
1958)
and
Mcdonald
(
2000)
were
designed
to
better
transfer
swimming
performance
estimates
to
the
field,
and
suggest
a
more
narrow
temperature
range
(
12.5­
17.5
°
C,
and
17
°
C,
respectively)
of
temperatures.
Based
on
these
factors
it
seems
reasonable
to
assume
that
exposure
to
temperatures
above
17­
17.5
°
C
may
have
detrimental
biological
consequences
in
cases
where
overcoming
predators
and
river
obstacles
is
necessary.
Since
these
temperature
effects
are
not
apparently
affected
by
average
exposures,
it
is
believed
that
a
7DADMax
of
17­
17.5
°
C
should
be
considered
the
upper
temperature
regime
that
will
fully
protect
the
swimming
performance
of
salmon
and
trout
in
riverine
environments.

Field
Distribution
Restrictions
in
Juvenile
Fish:

Chinook
Salmon
In
the
work
of
Torgersen
et
al.
(
1999)
fish
from
portions
of
the
John
Day
River
basin
in
Oregon
were
found
strongly
associated
with
cool
water
segments
in
the
middle
fork
where
temperatures
ranged
from
22­
24
°
C
along
its
length,
but
were
much
less
related
to
temperature
fluctuations
in
the
cooler
north
Fork
where
temperatures
ranged
from
15­
22
°
C
along
its
length.
In
an
unpublished
report
to
the
USEPA
and
the
National
Science
Foundation
(
1999)
Torgersen,
Baxter,
Li,
and
McIntosh
further
discuss
the
spatial
distribution
of
fish
in
the
John
Day.
They
noted
that
the
cross­
over
point
where
cold
water
fish
no
longer
dominate
over
warm
water
fish
occurred
where
afternoon
temperatures
reached
17
°
C.
The
general
range
for
chinook
salmon
has
been
suggested
as
extending
from
0.0­
0.6
°
C
to
25
°
C,
with
an
optimal
or
preferred
range
extending
from
6.7­
10
°
C
to
13.9­
14.4
°
C
(
Bell,
1986;
Piper
et
al.,
1982).
Page
74
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Coho
Salmon
In
trying
to
determine
the
cause
for
coho
losses
from
streams
on
the
west
side
of
Mount
St.
Helens,
Martin
et
al.
(
1986)
found
that
summer
mortality
was
correlated
with
both
high
monthly
average
temperatures
and
maximum
August
diel
fluctuations.
One
creek
experiencing
high
mortality
exceeded
25
°
C
for
10
days
in
one
year
and
30
days
in
the
next.
Bisson
et
al.
(
1988)
found
that
growth
remained
positive
in
streams
that
exceeded
24.5
°
C,
but
that
at
about
22
°
C
the
coho
seek
out
cool
water
areas.
Hall
and
Lantz
(
1969)
found
no
statistically
significant
change
in
coho
abundance
in
a
clear­
cut
test
stream
where
maximum
summer
temperatures
were
increased
from
16.1­
16.6
°
C
to
24­
30
°
C,
even
though
cutthroat
trout
populations
experienced
a
decline
of
75%.
In
a
study
of
salmonid
distributions
in
a
coastal
Oregon
river
drainage,
Frissell,
Nawa,
and
Liss
(
1992)
found
that
cutthroat,
coho,
and
chinook
salmon
dropped
out
in
sequence
as
daily
maximum
temperatures
increased,
with
rainbow
trout
being
the
only
species
found
in
waters
exceeding
23
°
C.
An
abrupt
loss
of
coho
and
chinook
was
found
to
occur
between
21
and
23
°
C,
and
while
small
numbers
of
coho
were
found
in
the
warmer
waters,
they
were
always
found
in
association
with
small
cool
pockets
of
otherwise
warm
reaches.

Chum
Salmon
Juvenile
and
adult
chum
salmon
have
been
observed
in
waters
with
temperatures
ranging
from
0­
25.6
°
C
(
Bell,
1986).
Bonar
et
al.
(
1989)
considered
10.1
°
C
to
be
the
optimal
temperature
for
adults
in
marine
waters.

Pink
Salmon
Pink
salmon
are
noted
as
being
found
in
temperatures
ranging
from
0­
25.6
°
C
(
Bell,
1986),
while
Sheridan
(
1962)
has
stated
that
the
range
in
Alaskan
waters
extends
from
7.2­
18.3
°
C.
Welch
et
al.
(
1995)
determined
that
the
upper
thermal
boundary
for
the
offshore
marine
occurrence
of
pink
salmon
was
10.4
°
C.

Sockeye
Salmon
While
sockeye
salmon
are
noted
to
occur
within
the
range
of
0.6­
21
°
C
(
Piper
et
al.,
1982),
the
range
of
temperatures
under
which
they
perform
optimally
may
be
more
restricted.
The
optimal
temperature
for
juvenile
rearing
was
identified
10­
15
°
C
by
Piper
et
al.
(
1982),
and
10.6­
12.2
°
C
by
Bell
(
1986).

Steelhead
and
Rainbow
Trout
Frissel
et
al.
(
1992)
studied
the
distribution
of
rainbow
trout
and
found
that
while
they
could
be
found
in
water
temperatures
over
23
°
C,
there
was
a
general
threshold
response
for
age
1+
fish
above
22
°
C
and
for
age
2+
fish
above
21
°
C.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
75
Washington's
Surface
Water
quality
Standards
The
work
of
Li
et
al.
(
1993;
and
1994)
and
Li
et
al.
(
1991;
as
cited
in
Spence
et
al.,
1996),
studying
interior
populations
of
rainbow
trout,
suggest
that
while
rainbow
trout
showed
no
avoidance
reactions
when
stream
temperatures
were
below
20
°
C,
they
actively
avoided
staying
in
waters
warmer
than
23­
25
°
C.
While
the
works
of
Li
et
al.
(
1991,
1993,
and
1994)
were
conducted
on
interior
populations
of
trout
(
likely
to
have
been
redband
trout),
Behnke
(
1992)
reported
finding
redband
trout
in
the
desert
basins
of
southern
Oregon
and
northern
Nevada
that
regularly
encounter
temperatures
that
kill
other
trout.
Trout
in
these
intermittent
desert
streams
were
found
actively
feeding
in
water
of
28.3
°
C.

Kaya
et
al.
(
1977)
found
that
daily
maximum
temperatures
exceeding
25
°
C
caused
rainbow
trout
to
move
out
of
the
main
stem
of
the
Firehole
River
in
Montana.
These
fish
would
move
into
tributary
streams
that
averaged
6­
10
°
C
lower
in
temperature.
Behnke
(
1992)
in
a
review
of
the
Kaya
(
1978)
study
concluded
that
thousands
of
years
of
adaptation
to
a
desiccating
environment
have
enabled
the
Oregon
desert
redband
trout
to
feed
at
high
temperatures,
but
60­
70
years
seem
too
few
to
have
allowed
the
planted
rainbow
trout
to
expand
their
functional
feeding
temperature
in
the
Firehole
River.
While
there
are
observations
of
redband
trout
feeding
and
surviving
at
relatively
high
temperatures
for
a
salmonid
(
28
°
C,
Behnke
1992;
27.4
°
C,
Sonski
1986
and
27
°
C,
Bowers
et
al.
1979
as
cited
in
USEPA,
2001),
it
is
unclear
whether
temperatures
were
measured
at
the
location
of
the
stream
where
the
fish
were
found.
These
trout
may
rely
on
microhabitats
or
thermal
refugia
to
maintain
populations
in
desert
environments
(
Ebersole
et
al.
2001).
Thus
it
may
not
be
appropriate
to
use
these
observations
to
conclude
that
such
temperatures
are
not
harmful.
However,
the
basis
for
the
argument
that
interior
redband
may
have
warmer
optimum
limits
then
the
coastal
rainbow
stocks
seems
likely,
even
if
it
is
not
well
established
in
the
literature.
It
is
believed
that
the
upper
range
of
what
is
sometimes
found
as
fully
protective
for
rainbow
trout
should
be
applied
specifically
to
these
interior
non­
anadromous
redband
trout.

Cutthroat
Trout
Bell
(
1986)
used
a
review
of
the
literature
to
conclude
that
the
range
for
cutthroat
trout
was
from
0.6­
22.7
°
C,
and
the
preferred
range
was
from
9.4­
12.7
°
C.
Hall
and
Lantz
(
1969)
found
a
75%
reduction
in
a
cutthroat
trout
population
in
response
to
experimental
logging
of
riparian
canopies
in
three
coastal
streams.
Prior
to
treatment,
these
streams
had
monthly
average
temperatures
ranging
from
6.1­
12.8
°
C
and
maximum
temperatures
ranging
from
16.1­
16.6
°
C.
In
the
treatment
stream
experiencing
the
greatest
canopy
removal,
daily
maximum
temperatures
increased
to
24
and
30
°
C
in
the
two
years
following
treatment.
These
results
compared
well
with
the
work
of
Frissell
et
al.
(
1992)
who
studied
the
distribution
of
salmonids
in
a
small
coastal
river
system
in
southwest
Oregon.
They
found
that
cutthroat,
coho,
and
chinook
salmon
dropped
out
in
sequence
as
maximum
temperatures
increased,
with
rainbow
trout
the
only
species
present
in
waters
exceeding
23
°
C.
Cutthroat
were
absent,
and
coho
salmon
rare
in
segments
exceeding
21
°
C.
While
Varely
and
Gresswell
(
1988;
as
cited
in
Gresswell,
1995)
reported
that
Yellowstone
cutthroat
are
found
in
geothermally
heated
streams
with
ambient
temperatures
of
27
°
C,
they
report
these
fish
are
found
associated
with
cooler
thermal
refuges.
Varley
and
Gresswell
(
1988)
suggested
that
the
optimum
water
temperature
for
Yellowstone
cutthroat
trout
is
from
5.5­
15.5
°
.
Kelly
Page
76
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
(
1993:
as
cited
in
Gresswell,
1995),
however,
reportedly
found
that
cutthroat
were
excluded
from
a
tributary
to
the
Yellowstone
River
because
summer
water
temperatures
often
exceeded
22
°
C.
Dunham
et
al.
(
unpub.;
as
cited
in
Dunham,
1999)
found
Lahontan
cutthroat
appear
to
have
distributional
limits
that
correspond
closely
to
maximum
water
temperatures
of
26
°
C.
Pauley
et
al.
(
1989)
suggest
that
cutthroat
trout
are
not
usually
found
in
waters
where
the
maximum
temperature
exceeds
22
°
C
even
though
they
may
tolerate
brief
periods
of
temperatures
as
high
as
26
°
C.

Table
4.14.
Summary
of
information
on
the
field
distribution
of
salmonid
and
trout
in
relation
to
temperature.

Species
Temperature
for
range
Optimal
or
preferred
range
Author
Chinook
15­
22
Range
of
river
temperatures
where
fish
not
found
in
association
with
refugia
Torgersen
et
al.,
1999(
a)

17
(
daily
max)
Point
where
warm
water
fish
begin
to
dominate
Torgersen
et
al.,
1999(
b)

0.0­
0.6
to
25
6.7­
10
to
13.9­
14.4
General
observed
range
Bell,
1986,
and
Piper
et
al.,
1982
Coho
22
Begin
to
seek
out
refugia
Bisson
et
al.,
1988
24­
30
(
daily
max)
No
change
in
abundance
with
rise
from
16.1­
16.6
to
24­
30
Hall
and
Lantz,
1969
21­
23
(
daily
max)
Abrupt
loss
in
coho
Frissel
et
al.,
1992
Chum
0­
25.6
General
observed
range
Bell,
1986
10.1
Optimal
in
marine
water
Bonar
et
al.,
1989
Pink
0­
25.6
General
observed
range
Bell,
1986
7.2­
18.3
In
Alaska
Sheridan,
1962
10.4
Upper
thermal
boundary
in
marine
water
Welch
et
al.,
1995
Sockeye
0.6­
21
10­
15
General
observed
range
Piper
et
al.,
1982
10.6­
12.2
Bell,
1986
Steelhead
and
rainbow
trout
22
(
age
1+)
21
(
age
2+)
General
threshold
Frissel
et
al.,
1992
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
77
Washington's
Surface
Water
quality
Standards
response
27,
27.4,
28.3
Redband
trout
found
actively
feeding
Bowers,
et
al.,
1979,
Sonski,
1986,
and
Behnke,
1992
25
(
daily
max)
15­
19
Caused
rainbow
trout
to
move
from
main
stem
Kaya
et
al.,
1977
23­
25
20
Redband
actively
avoided
23­
25,
but
not
20
Li
et
al.,
1991,
1993,
and
1994
Cutthroat
trout
0.6­
22.7
9.4­
12.7
General
observed
range
Bell,
1986
16.1­
16.6
(
daily
max)
Increase
from
16.1­
16.6
to
24­
30
resulted
in
a
75%
population
reduction
Hall
and
Lantz,
1969
21
(
daily
max)
Absent
in
segments
exceeding
21
Frissel
et
al.,
1992
27
5­
15.5
Yellowstone
cutthroat
found
in
river
at
27
in
association
with
refugia
Varley
and
Gresswell,
1988
22
Yellowstone
cutthroat
reported
to
be
excluded
above
22
Kelly,
1993
26
(
daily
max)
Lahontan
cutthroat
distributional
limits
Dunham
et
al.,
1999
22
(
daily
max)
Seldom
found
above
Pauley
et
al.,
1989
While
some
authors
report
finding
salmon
and
trout
in
the
range
of
24­
30
°
C,
most
found
that
their
presence
was
dependent
upon
cold
water
refugia.
The
general
pattern
of
the
findings
cited
above
seems
to
suggest
that
daily
maximum
river
temperatures
of
21­
22
will
create
a
distributional
limit
to
finding
salmon
and
trout,
and
that
healthy
populations
are
more
likely
to
be
found
within
the
range
of
10­
19
°
C.
Most
authors,
however,
place
the
optimal
range
below
17
°
C
and
Torgersen
et
al.,
(
1999b)
additionally
found
that
warm
water
species
begin
to
dominate
above
17
°
C.
Thus
based
on
this
line
of
evidence,
a
7DADMax
of
16.05
°
C
should
be
considered
the
limit
for
finding
strong
populations
of
salmon
and
trout,
and
river
temperatures
above
a
7DADMax
of
20.05­
21.05
°
C
should
be
considered
to
create
a
thermal
Page
78
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
barrier
to
the
general
summer
distribution
of
salmon
and
trout.
Beyond
this
temperature
barrier,
the
presence
of
salmon
and
trout
may
be
primarily
dependent
on
the
availability
of
cold
water
refugia.
There
is
reason
to
suspect
that
redband
trout
may
have
slightly
higher
temperature
ranges
than
the
other
native
salmon
and
trout.
The
information
is
scant
and
should
be
used
with
caution,
but
it
suggests
that
the
healthy
distribution
of
redband
trout
may
be
better
described
by
18.05
°
C
as
7DADMax,
and
22.05
°
C
may
more
appropriately
describe
the
thermal
barrier
for
redband
populations.

General
Findings
and
Recommendations
Found
in
the
Literature:

Chinook
Salmon
The
Independent
Scientific
Group
(
1996)
concluded
that
juvenile
chinook
salmon
rearing
is
optimal
where
temperatures
are
maintained
in
the
range
of
12­
17
°
C;
and
suggests
15
°
C
is
most
optimal.

Coho
Salmon
Servizi
and
Martens
(
1991)
found
that
susceptibility
to
sediment
toxicity
increased
by
33%
at
18
°
C,
and
Shelbourn
(
1980)
determined
that
hypo­
osmoregulatory
capacity
is
optimized
at
14
°
C.

Steelhead
and
Rainbow
Trout
Bell
(
1986)
recommended
that
temperatures
generally
be
maintained
in
the
range
of
7.3­
14.5
°
C
for
optimal
rearing
of
juvenile
steelhead.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
79
Washington's
Surface
Water
quality
Standards
Conclusion
on
Juvenile
Rearing
of
Salmon
and
Trout:

Table
4.15.
The
multiple
lines
of
evidence
for
juvenile
rearing
of
salmon
and
trout.

Line
of
Evidence
7DADMax
(
°
C)
Midpoint
Comments
Laboratory
Growth
Studies
Conducted
at
Constant
Temperatures
14.18­
18.14
16.16
Based
on
well
controlled
laboratory
tests.

Ranges
Identified
in
Literature
as
Optimal
for
Growth
15­
21
18
Basis
for
estimates
and
intended
metrics
unclear.

Comparison
Test
Regimes
With
Better
Growth
11.8­
15.4
13.6
Maximum
growth
temperatures
were
not
determined,
only
favorable
growth
compared
to
one
other
significantly
warmer
or
cooler
temperature.
Laboratory
Growth
Studies
in
Fluctuating
Temperature
Regimes
13.16­
21.14
17.15
Based
on
well
controlled
laboratory
tests.

Field
Studies
on
Growth
14.05­
16.05
15.05
Regime
with
better
standing
crops.
Specific
temperature
limits
were
not
determined.
Predation
and
Competition
17.60­
18.18
17.89
Displacement
of
juvenile
salmon
and
trout
Temperature
Preferences
in
the
Laboratory
14.2­
19.3
16.75
Mode
of
distribution
of
fish
cycling
between
temperature
zones
Swimming
Performance
and
Scope
for
Activity
17­
17.5
17.25
Based
primarily
on
controlled
laboratory
studies.

Field
Distribution
­
Healthy
16.05
16.05
Basis
and
methodology
generally
unclear
for
most
literature
estimates.
Thus
this
estimate
relies
on
the
general
upper
range
considered
healthy,
and
temperatures
above
which
coldwater
species
begin
to
loose
dominance
Best
estimate
of
threshold
14.78­
18.08
mid.
pt.
16.43
Using
all
nine
lines
of
evidence
cited
above,
the
upper
limit
to
defining
a
healthy
summer
rearing
temperature
would
occur
within
the
range
of
14.78­
18.08
°
C
as
a
7DADMax,
with
an
Page
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Evaluating
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Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
overall
mean
of
16.43
°
C.
While
each
line
of
evidence
has
unique
strengths
and
weaknesses,
the
second
and
third
lines
of
evidence
are
particularly
questionable.
The
second
line
of
evidence
consists
of
technically
unsubstantiated
recommendations
of
specific
authors,
and
the
range
produced
in
the
third
line
of
evidence
is
driven
by
comparisons
tests
that
only
compared
and
contrasted
two
very
different
test
regimes
(
e.
g.,
only
compared
8
to
12)
without
any
effort
to
identify
maximum
growth.
Without
these
two
very
weak
lines
of
evidence,
the
range
within
which
favorable
juvenile
rearing
would
be
expected
changes
to
a
7DADMax
of
15.18­
18.05
°
C
(
16.62
°
C).
Since
both
estimates
would
round
out
to
the
same
½
degree
value,
it
is
recommended
that
a
7DADMax
of
16.5
°
C
be
considered
fully
protective
of
juvenile
rearing.

iv)
Juvenile
Winter
Holding
Chinook
Salmon
Temperature
declines
below
1.1­
5
°
C
can
cause
fish
to
become
dormant
and
move
into
the
substrate.
Once
fish
have
initiated
hiding
behavior,
it
may
take
temperatures
rising
again
to
above
7
°
C
to
bring
them
back
out
(
Chapman
and
Bjornn,
1969).
While
this
natural
behavior
is
a
healthy
response
for
winter
survival,
unseasonably
cold
discharges
causing
water
temperatures
to
unseasonably
fall
below
5­
7
°
C
need
to
be
avoided.
Dropping
the
temperature
to
2.5
°
C
for
even
a
few
days
may
result
in
mortality
to
chinook
that
were
acclimated
to
a
river
temperature
of
15
°
C
(
Brett,
1956).
To
avoid
initiating
unseasonable
hiding
behavior,
to
avoid
lethal
effects,
and
to
encourage
strong
growth
rates
the
daily
low
temperature
should
typically
exceed
7
°
C
during
the
growing
season.

Steelhead
and
Rainbow
Trout
After
steelhead
fry
emerge
from
the
gravel
they
move
to
the
slow
moving
waters
of
the
stream
margins,
shifting
to
faster
and
deeper
waters
as
they
grow
larger
(
Chapman
and
Bjornn,
1969).
Martin
et
al.
(
1991)
and
Wydoski
and
Whitney
(
1979)
note
the
importance
of
cover
to
juvenile
steelhead.
Large
woody
debris,
substrate,
and
turbulence
are
all
identified
as
important
habitat
for
young
juveniles.
Temperature
is
believed
to
affect
the
habitat
selection
and
migration
of
juveniles.
Mullan
et
al.
(
1992)
found
that
fry
emigrate
from
cold
headwater
streams
to
down­
stream
reaches
to
rear
in
warmer
waters.
Chapman
and
Bjornn
(
1969)
found
that
young
steelhead
may
move
downstream
in
the
fall
to
over­
winter
in
larger
streams.
Chapman
and
Bjornn
(
1969)
found
that
5­
5.5
°
C
marked
the
boundary
between
activity
and
inactivity
in
steelhead,
with
fish
entering
the
substrate.
They
found
that
steelhead
emigration
could
be
stopped
by
warming
the
water
from
7.2
°
C
to
11­
12.2
°
C.
Mullan
et
al.
(
1992)
suggest
that
while
steelhead
juveniles
may
reside
in
the
dark
frozen
snow
covered
tributaries
near
0
°
C
for
up
to
5
months,
6
°
C
may
form
the
boundary
that
allows
winter
growth
to
occur.

Some
controlled
field
studies
have
shed
some
light
on
the
winter
rearing
habits
of
rainbow
trout.
Rainbow
trout
actively
feed
at
very
cold
temperatures
(
0­
0.6
°
C)
even
under
ice
Evaluating
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Washington's
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quality
Standards
covered
streams.
Their
winter
feeding
habits
are
primary
controlled
by
the
availability
of
food.
They
consume
dislodged
insects,
and
feed
on
insects
that
emerge
at
more
moderate
temperatures
(
4.4
°
C
or
more).
Winter
mortality
is
likely
due
to
a
combination
of
factors
including
insufficient
food
supply,
and
physical
damage
from
snow
slumps
and
ice
scour
(
Needham
and
Jones,
1959;
Needham
and
Slater,
1944;
Maciolek
and
Needham,
1952;
and
Reimers,
1957).

Cutthroat
Trout
Jokober
et
al.
(
1998)
found
that
westslope
cutthroat
trout
made
extensive
migrations
downstream
to
escape
winter
conditions
that
resulted
in
supercooling
(<
1
°
C)
of
the
water
and
anchor
ice
formation.

v)
Adult
Migration
Barriers
to
Migration
of
Salmon
and
Trout:

Chinook
Salmon
After
spending
3­
4
years
in
the
ocean
mature
chinook
salmon
begin
their
return
migrations
to
freshwaters
to
spawn.
Temperatures
can
create
serious
problems
for
migrating
salmon.
In
addition
to
posing
the
threat
of
direct
lethality
to
adult
spawners,
temperatures
can
create
blockages
that
stop
migrating
fish,
create
conditions
that
result
in
high
mortality
of
spawners
from
disease,
and
reduce
the
overall
fitness
of
migrants.
Since
migrating
salmon
do
not
feed
in
freshwaters
they
must
enter
freshwater
with
sufficient
fat
and
muscle
reserves
to
supply
their
metabolic
requirements
up
to
and
through
the
act
of
spawning.
The
increased
active
and
basal
metabolic
demands
caused
by
traveling
and
holding
in
warmer
waters
uses
up
stored
energy
reserves
at
a
more
rapid
rate.
This
can
result
in
a
decrease
in
the
quality
and
quantity
of
eggs
as
well
as
an
overall
reduction
in
the
fitness
of
the
adult
fish
that
need
to
migrate
and
negotiate
obstacles,
excavate
and
guard
redds,
and
complete
the
act
of
spawning.
Berman
and
Quinn
(
1991)
demonstrated
that
in
the
months
prior
to
spawning,
spring
run
chinook
actively
sought
out
cool
water
refuges
in
the
Yakima
River,
in
Washington.
These
fish
were
able
to
maintain
average
internal
temperatures
2­
5
°
C
below
the
ambient
river
condition,
which
may
have
reduced
their
metabolic
demand
by
12­
20%.
Mcdonald
et
al.
(
2000)
found
that
chinook
salmon
in
the
Fraser
River
in
Canada
suffered
unusually
large
losses
(
25%)
where
mean
daily
river
temperatures
frequently
exceeded
20
°
C
and
reached
a
high
of
23
°
C.
The
most
widespread
concern
with
warm
temperatures
is
from
prespawning
mortality
due
to
an
increased
incidence
of
diseases.
These
diseases
can
directly
kill
or
impair
healthy
fish,
or
act
secondarily
through
infection
of
the
minor
wounds
that
normally
occur
in
migrating
fish.
Disease
is
a
serious
concern
and
is
discussed
and
incorporated
separately
in
the
recommendations
of
this
paper.

Daily
maximum
temperatures
rising
above
21­
22
°
C
are
widely
cited
as
causing
barriers
to
migrating
chinook
salmon
(
Stabler,
1981;
Bumgarner
et
al.,
1997;
Hallock,
Elwell,
and
Fry,
Page
82
Evaluating
Standards
for
Protecting
Aquatic
Life
in
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Water
quality
Standards
1970;
Thompson,
1945,
as
cited
in
Snyder
and
Blahm,
1971;
Don
Ratliff,
1977,
as
cited
in
Stabler,
1981;
Fish
and
Hanavan,
1948,
and
Major
and
Mighell,
1967,
as
cited
in
USEPA,
1971;
and
Alabaster,
1988,
as
cited
by
USEPA,
2001).
Hallock,
Elwell,
and
Fry
(
1970)
suggested
that
maximum
temperatures
of
18.9
°
C
in
association
with
low
dissolved
oxygen
levels
(
5ppm)
created
a
partial
block
of
migrating
chinook
salmon.
However,
some
authors
note
chinook
not
showing
avoidance
for
temperatures
as
high
as
24.4
°
C
(
Gray,
1990;
Dunham,
1968;
as
cited
in
CDWR,
1988).
However,
Thompson
(
1945;
as
cited
in
Snyder
and
Blahm,
1971)
suggested
that
it
was
the
difference
in
temperatures
that
stopped
chinook
from
migrating
from
the
Columbia
to
the
Snake
River.
Differences
were
17.2:
21.7
°
C
and
22.2:
26.1
°
C
when
blockages
occurred,
with
migration
resuming
when
the
difference
approached
1.6
°
C.
Similarly,
Gray
(
1990)
suggested
that
incremental
increases
of
9­
11
°
C
formed
a
barrier
to
migration.
Temperatures
above
20­
21
°
C
are
certainly
stressful
for
chinook
salmon.
Sauter
and
Maule
(
1997)
reported
cessation
of
feeding
as
well
as
thermoregulatory
behavior
in
sub­
yearling
fall
chinook
held
between
18­
20
°
C,
with
exposure
to
20
°
C
for
several
hours
inducing
heat
shock
proteins
(
Sauter
et
al.,
in
review,
and
M.
Hargis,
personal
comm.;
as
cited
in
Sauter
and
Maule,
1997).
In
a
field
study
by
Frissel,
Nawa,
and
Liss
(
1992),
it
was
found
that
maximum
water
temperatures
in
a
coastal
river
system
in
Oregon
were
linked
to
the
presence
or
absence
of
various
species
of
salmonids.
While
it
was
noted
that
cutthroat
were
absent
and
coho
salmon
rare
or
absent
in
segments
exceeding
21
°
C,
chinook
dropped
out
completely
only
at
23
°
C;
although,
their
presence
in
such
waters
was
associated
with
positioning
in
small
cool
pockets
in
otherwise
warm
reaches.
Some
authors
have
suggested
criteria
for
the
protection
of
migrating
chinook
salmon.
Piper
et
al.
(
1982)
considering
this
important
life
stage
suggested
that
7.2­
15.6
°
C
was
necessary
to
protect
upstream
migration
and
maturation.
Bell
(
1973;
as
cited
by
Everest
et
al.,
1985)
suggested
that
temperatures
should
be
within
the
range
of
3.3­
13.3
°
C
for
spring
chinook,
13.9­
20
°
C
for
summer
chinook,
and
10.6­
19.4
°
C
for
fall
chinook.
Support
for
assuming
a
general
20­
21
°
C
threshold
for
salmon
migration
can
also
be
found
in
the
technical
literature
on
lethality
studies.
As
summarized
below,
temperatures
of
20­
22
°
C,
particularly
at
lower
prior
acclimation
temperatures,
can
be
directly
lethal
to
chinook
salmon
(
Brett,
1956;
Brett
et
al.,
1982;
Coutant,
1970;
Beacham
and
Withler,
1991;
Becker,
1973;
Orsi,
1971;
as
cited
in
CDWR,
1988)
with
a
7­
day
exposure.
It
also
appears
from
the
available
evidence
that
adults
may
be
more
sensitive
than
the
juveniles
which
are
most
typically
tested
(
Becker,
1973).
Based
on
the
technical
literature,
it
is
concluded
that
to
prevent
a
risk
of
causing
blockage
of
migrating
chinook
salmon
daily
maximum
temperatures
should
not
exceed
21­
22
°
C,
particularly
when
the
migrating
fish
will
be
acclimated
to
lower
(
2­
6
°
C)
water
temperatures
as
they
travel
upstream.

Many
runs
of
chinook
will
need
to
hold
or
travel
during
the
summer
when
stream
temperatures
are
at
a
maximum
in
the
lower
and
mid­
elevation
rivers.
Therefore,
criteria
to
protect
migration
should
be
set
to
protect
against
chronic
sublethal
effects
as
well
as
acute
effects
such
as
blockages
and
lethality.
While
chinook
adults
can
almost
certainly
withstand
occasional
daily
peak
temperature
cycles
up
to
22
°
C,
the
general
condition
encountered
by
these
migrating
fish
should
approach
more
optimal
conditions.
This
will
avoid
unhealthy
levels
of
stress
and
utilization
of
stored
energy
reserves.
It
is
concluded
that
summer
daily
maximum
temperatures
during
migration
should
generally
remain
within
or
below
the
range
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
83
Washington's
Surface
Water
quality
Standards
identified
previously
as
fully
protective
for
juvenile
chinook
salmon
rearing,
and
single
daily
maximum
temperatures
be
maintained
below
20­
21
°
C
wherever
possible.

Where
fish
may
hold
in
waterbody
segments
for
long
periods
of
time
prior
to
spawning,
as
is
commonly
the
case
for
spring
chinook,
special
protection
may
be
warranted
on
a
casespecific
basis.
To
protect
adult
fish
that
are
holding
throughout
the
summer,
temperature
limits
should
be
established
that
recognizes
these
fish
are
living
off
their
stored
energy
reserves
and
may
be
in
a
ripe
condition
in
the
later
portion
the
summer
or
early
Fall.
Three
physiological
endpoints
seem
especially
relevant
in
guiding
the
selection
of
an
appropriate
summer
holding
criteria.
The
first
is
the
preferendum
values
(
constant
11.7
°
C,
and
a
range
of
6.7­
14.4
°
C),
the
second
is
the
fully
protective
rearing
threshold
(
an
average
of
14­
15
°
C,
and
a
7­
day
average
of
the
daily
maximum
temperatures
of
16.5
°
C)
and
the
third
is
the
temperature
regime
identified
to
protect
eggs
invivo
in
ripe
females
(
weekly
average
temperatures
below
(
13­
15
°
C).
In
consideration
of
these
factors
it
is
concluded
that
where
chinook
are
holding
over
the
summer,
the
average
water
temperatures
should
be
maintained
below
13­
14
°
C
and
the
7­
day
average
of
the
daily
maximum
temperatures
maintained
below
16­
17
°
C.

Coho
Salmon
While
adults
can
migrate
through
waters
warmer
than
considered
fully
protective
for
juvenile
rearing,
the
same
thresholds
which
produce
metabolic
stress
with
juveniles
are
likely
to
produce
stress
in
adults
that
can
lead
to
lethal
and
sublethal
effects.
Beschta
et
al.
(
1987)
suggested
as
a
basis
for
water
quality
criteria
that
upstream
migration
occurs
between
7.2­
15.6
°
C.
Studies
with
other
species
support
the
work
of
Thomas
et
al.
(
1986)
in
showing
that
adults
may
actually
be
somewhat
more
temperature
sensitive
than
juveniles.
Sensitivity
is
enhanced
in
adults
through
the
fact
that
they
do
not
feed
during
their
freshwater
migration
and
must
rely
on
their
stored
fat
and
muscle
reserves
to
see
them
through
the
spawning
process.
The
stress
of
higher
temperatures
not
only
influences
the
health
of
the
spawner,
but
to
some
extent
it
also
can
effect
the
quality
of
unfertilized
eggs
carried
by
the
hen
salmon.
In
tests
evaluating
the
effects
of
holding
ripe
adult
coho
at
warm
temperatures,
Bouck
et
al.
(
1970;
as
cited
in
USEPA,
1971)
found
no
apparent
adverse
effects
to
eggs
in
utero
caused
by
prolonged
exposure
to
16.7
°
C.
Flett
et
al.
(
1996),
however,
found
that
adults
migrating
through
waters
often
warmer
than
20
°
C
experienced
reduced
quality
and
more
rapid
deterioration
of
eggs.
For
these
reasons
it
would
be
prudent
to
maintain
temperatures
close
to
the
range
considered
fully
protective
for
juvenile
coho
(
i.
e.,
7DADMax
temperatures
below
16.5
°
C)
over
the
migration
routes
used
by
adult
spawners.
Page
84
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Chum
Salmon
Studies
were
not
found
that
establish
a
specific
basis
for
setting
adult
upstream
migration
temperature
threshold.
Beschta
et
al.
(
1987)
has
suggested
as
a
basis
for
establishing
water
quality
criteria
that
upstream
migration
is
protected
by
keeping
water
temperatures
in
the
range
of
8.3­
15.6
°
C.
While
adults
can
certainly
withstand
much
higher
temperatures
for
short
periods
of
exposure
without
directly
lethal
effects,
the
potential
of
warm
waters
(
above
14
°
C)
to
reduce
the
number
of
viable
eggs,
as
shown
in
other
fish
species,
should
be
cause
for
some
caution.
Since
chum
salmon
spawn
just
above
tidewater
and
have
very
short
migrations
in
most
watersheds,
their
time
in
migration
and
thus
the
potential
for
sublethal
effects
appears
at
least
somewhat
naturally
mitigated
in
most
cases.
Recommendations
to
prevent
barriers
to
migration
and
to
protect
other
mature
Pacific
salmon
spawners
ripe
with
eggs
should
also
be
applied
for
chum
salmon.

Pink
Salmon
Unlike
most
Pacific
salmon,
the
pink
salmon
that
occur
in
Washington
have
relatively
short
migrations
and
concerns
over
disease
and
over
depleting
energy
reserves
are
less
than
with
the
other
salmon.
However,
migrating
adults
will
pass
through
the
lower
reaches
of
major
rivers
near
the
period
of
maximum
seasonal
temperatures
and
disease
is
still
a
concern
that
should
be
considered
in
setting
any
final
temperature
recommendations
(
discussed
and
incorporated
separately).
It
is
also
worthy
of
consideration
that
pink
salmon
have
lower
maximum
sustained
swimming
speeds
than
sockeye
salmon
(
20%
less)
and
require
more
energy
to
support
the
same
speed
(
30%
higher)
(
Brett,
1982;
as
cited
in
Brett,
1995).
While
no
specific
recommendation
is
warranted
for
pink
salmon
migration,
the
literature
reviewed
herein
suggests
that
caution
should
be
exercised
in
applying
the
upper
end
of
the
optimal
range
to
waters
that
may
support
longer
than
typical
migrations
or
periods
of
holding
by
pink
salmon.

Sockeye
Salmon
Migration
exerts
a
tremendous
strain
on
salmon.
Idler
and
Clemens
(
1959)
found
that
female
sockeye
salmon
in
the
Fraser
River
may
use
between
91.4­
96%
of
their
body
fat
reserves,
and
53­
61%
of
their
protein
reserves
from
the
time
of
entrance
to
completion
of
spawning.
As
noted
by
Brett
(
1983),
migration
and
egg
production
uses
up
most
of
the
energy
stored
from
ocean
feeding
and
leaves
"
all
too
slim
a
safe
margin
of
energy
reserves".

Linley
(
1993;
as
cited
in
Quinn
et
al.,
1997)
reportedly
found
that
populations
with
arduous
migrations
show
lower
levels
of
reproductive
output
(
ovary
weight)
than
populations
with
shorter
migrations.
At
a
constant
16.2
°
C,
depletion
of
fat
reserves
and
reproductive
organ
abnormalities
were
noted
by
Bouk
(
1977),
and
Gilhousen
(
1990)
found
that
high
prespawning
mortalities
were
associated
with
adult
salmon
migrating
through
waters
having
daily
maximum
temperatures
between
17.5­
19
°
C.
Temperatures
above
15.5
°
C
(
as
an
apparent
daily
average
value)
were
also
noted
by
Gilhousen
(
1990)
as
being
linked
to
higher
prespawning
mortality
from
columnaris
disease
in
adult
Fraser
River
sockeye
salmon.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
85
Washington's
Surface
Water
quality
Standards
Mcdonald
et
al.
(
2000)
in
a
comprehensive
study
of
the
migratory
success
of
Fraser
River
sockeye
in
relation
to
temperatures
found
that
during
unusually
warm
years
(
mean
daily
temperatures
above
20
°
C)
hormonal
and
stress
indicators
suggested
that
fish
were
suffering
significant
physical
stress
and
maturation
impairment.
Temperature
conditions
detrimentally
effected
the
ability
of
the
fish
to
migrate
successfully
through
rapids.
Poor
spawning
success,
poor
egg
quality
and
viability,
and
senescent
death
prior
to
spawning
were
all
observed.
The
authors
reference
unpublished
laboratory
work
of
Dr.
Craig
Clarke
showing
that
laboratory
exposure
to
19
°
C
over
a
two­
week
period
significantly
depressed
hormones
controlling
maturation,
but
exposure
to
15
°
C
did
not.
This
was
noted
as
consistent
with
findings
with
rainbow
trout
showing
that
temperatures
of
17
°
C
can
reduce
steroid
synthesis
in
rainbow
trout
testes
(
citing
Manning
and
Kime,
1985).
Mcdonald
et
al.
(
2000)
suggested
that
the
upper
threshold
for
successful
reproduction
of
migrating
sockeye
salmon
occurs
with
mean
daily
temperatures
at
the
lower
Fraser
River
(
Hells
Gate)
between
of
18­
22
°
C.
The
authors
also
note
that
field
studies
suggest
the
temperature
for
optimum
swimming
endurance
is
17
°
C
and
that
a
20%
reduction
occurs
at
21
°
C.
The
authors
conclude
that
migration
blockages,
susceptibility
to
disease,
impaired
maturation
processes,
increases
to
stress
parameters,
reduced
efficiency
of
energy
use,
and
reduced
swimming
performance
all
become
more
hazardous
as
daily
mean
temperatures
exceed
17
°
C.

Paulik
(
1960)
found
that
sockeye
subjected
to
daily
swimming
tests
did
not
live
as
long
as
control
fish,
and
postulated
that
as
migrating
salmon
move
upstream
their
swimming
capacity
declines
such
that
performance
is
progressively
reduced.
DeLacy
et
al.
(
1956)
using
eggs
from
coho
salmon
and
steelhead
trout
found,
however,
that
the
viability
of
sex
products
did
not
seem
to
be
affected
by
repeated
exhaustive
testing.

Welch
et
al.
(
1995)
found
that
the
upper
thermal
limit
to
the
off­
shore
occurrence
of
sockeye
salmon
was
8.9
°
C,
and
8.9
°
C
was
found
to
be
the
maximum
holding
temperature
in
lakes
by
migrating
adults
(
Wydoski
and
Whitney,
1979).

Quinn
and
Adams,
1996;
as
cited
in
Quinn
et
al.,
1997)
note
that
in
the
Columbia
River,
based
on
passage
data
at
Ice
Harbor
Dam,
migration
usually
ceases
at
temperatures
above
21
°
C.
Fish
and
Hanava,
(
1948;
as
cited
by
USEPA,
1971)
found
that
during
an
extremely
warm
year
(
1941)
sockeye
were
observed
congregating
in
small
previously
unused
cold
tributary
creeks
when
the
temperature
in
the
Columbia
rose
to
21.7­
23.9
°
C.
Major
and
Mighell
(
1966)
noted
that
entry
of
sockeye
from
the
Columbia
River
into
the
Okanogan
River,
was
blocked
when
rising
or
stable
daily
average
temperatures
were
above
21.1
°
C,
but
that
migration
would
resume
if
temperatures
were
falling.
Hatch
et
al.
(
1992)
found
that
when
water
temperatures
reached
daily
average
temperatures
of
22.8
°
C,
all
migration
of
sockeye
salmon
ceased,
that
the
bulk
of
the
migration
occurred
below
22.2
°
C,
and
that
surges
of
migration
occurred
when
temperatures
fell
to
below
21.1
°
C.

Constant
or
daily
average
temperatures
in
the
range
of
15.5­
17
°
C
have
been
found
by
numerous
authors
to
cause
excessive
depletion
of
energy
reserves
and
prespawning
losses,
and
17
°
C
has
been
set
as
the
limit
beyond
which
detrimental
physiologic
effects
would
Page
86
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
become
hazardous.
Since
maximum
temperatures
in
the
range
of
17­
19
°
C
have
also
been
identified
in
association
with
increased
losses
in
migrating
sockeye,
caution
should
be
exercised
in
selecting
daily
maximum
values.
Based
on
the
available
research,
temperatures
should
not
exceed
a
maximum
21­
day
average
of
14­
15
°
C,
and
the
7­
day
average
daily
maximum
temperatures
should
not
be
greater
than
16­
17
°
C
to
support
the
migration
of
adult
sockeye
salmon.
This
temperature
range
is
generally
associated
with
an
absence
of
prespawning
mortality
and
will
avoid
high
losses
of
stored
energy
reserves.
To
reduce
the
potential
for
causing
blockages
to
migrating
fish,
the
single
daily
maximum
temperatures
should
not
exceed
21­
22
°
C,
particularly
when
the
migrating
fish
are
acclimated
to
lower
(
2­
6
°
C)
water
temperatures
as
they
move
upstream.

Steelhead
Trout
Most
fish
returning
to
Washington's
streams
are
believed
to
have
been
at
sea
for
2
years.
Fish
that
have
been
at
sea
for
three
years
make
up
18.5­
33%
of
the
returning
fish,
and
only
a
few
are
at
sea
for
4
years
(
1­
3.9%).
The
largest
steelhead
are
generally
those
with
the
longest
oceanic
phase
(
Wydoski
and
Whitney,
1979).

Snyder
and
Blahm
(
1971;
as
cited
in
Monan
et
al.,
1975)
found
that
temperatures
of
23.9
°
C
created
a
barrier
to
the
migration
of
steelhead
trout
from
the
Columbia
to
the
Snake
River
that
remained
until
temperatures
declined
to
nearly
21.1
°
C.
Strickland
(
1967;
as
cited
in
Stabler,
1981)
also
noted
that
steelhead
destined
for
the
Snake
River
do
not
leave
the
relatively
cooler
waters
of
the
Columbia
River
until
the
Snake
cools
to
21
°
C
or
lower.
Fish
and
Hanavan
(
1948;
as
cited
in
Stabler,
1981
and
USEPA,
1971)
reported
that
steelhead
trout
entered
minor
typically
unused
tributaries
and
died
there
when
the
temperatures
in
the
Columbia
River
ranged
from
21.6
to
23.8
°
C.
On
the
Deschutes
River
in
Oregon,
nearly
all
steelhead
reportedly
stopped
migrating
past
Pelton
Dam
when
the
water
in
the
ladder
averaged
between
20­
21
°
C
and
the
water
below
the
dam
was
13­
14
°
C
(
Don
Ratliff,
personal
communication,
as
cited
in
Stabler,
1981).
Stabler
(
1981)
noted
that
Fessler
(
1977)
and
Everest
(
1973)
found
that
steelhead
halt
their
migration
and
will
enter
nonparent
streams
when
water
temperatures
exceed
21
°
C.
While
not
a
study
on
migration,
Nielsen
et
al.
(
1994)
found
temperatures
of
22
°
C
elicited
an
avoidance
reaction
in
steelhead
trout.
They
noted
that
foraging
began
to
decline
when
stream
temperatures
reached
approximately
22
°
C;
although,
it
was
noted
that
juvenile
steelhead
were
seen
actively
feeding
in
surface
waters
with
ambient
temperatures
up
to
24
°
C.
Fish
moved
to
cool
portions
of
stratified
pools
when
temperatures
exceeded
22
°
C,
but
not
at
or
below
22
°
C,
and
would
return
to
their
original
stream
territories
once
ambient
stream
temperatures
fell
to
about
23
°
C.
Based
on
the
above
referenced
studies,
daily
average
temperatures
of
21­
24
are
associated
with
avoidance
behavior
and
migration
blockage
in
steelhead
trout.

Support
for
assuming
21­
22
°
C
creates
significant
enough
stress
in
steelhead
as
to
create
a
potential
barrier
to
migration
is
also
found
in
lethality
studies
of
Coutant
(
1970)
and
Becker
(
1973).
Citing
what
appears
to
be
the
same
study,
these
authors
concluded
that
the
incipient
lethal
temperature
for
migrating
adult
steelhead
was
near
a
constant
21­
22
°
C.
They
noted
Evaluating
Standards
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Aquatic
Life
in
Page
87
Washington's
Surface
Water
quality
Standards
that
adults
appear
to
be
more
susceptible
to
high
temperatures
than
are
juveniles,
which
are
typically
used
in
lethality
studies.

Concerns
over
disease
may
warrant
restricting
temperatures
to
levels
well
below
that
which
would
result
in
direct
barriers
to
migrating
steelhead.
Diseases
of
native
fishes
is
discussed
and
incorporated
separately
in
this
paper.
In
setting
a
criteria
for
the
full
protection
of
migration,
it
is
also
important
to
consider
chronic
and
sublethal
effects
caused
by
warm
waters.
Migrating
spring
and
summer
steelhead
will
be
passing
through
during
the
peak
temperatures
of
summer
as
they
move
upstream
to
holding
areas
where
they
will
wait
until
the
following
stream.
While
often
repeat
spawners,
steelhead
still
rely
on
their
muscle
and
fat
reserves
to
hold
them
over
through
to
the
completion
of
spawning
and
return
to
the
ocean.
In
addition
to
concerns
over
disease,
the
metabolic
demands
of
swimming,
negotiating
obstacles,
and
supplying
the
basal
metabolic
requirements
while
holding
leaves
little
reserves
left
for
digging
redds
and
spawning
once
they
reach
their
spawning
streams.
The
warmer
the
water
the
more
energy
will
be
required
to
survive
until
spawning
is
completed
and
thus
the
greater
chance
that
fitness
will
be
affected
and
that
higher
pre­
and
post­
spawning
mortalities
will
occur.
For
these
reasons
it
would
be
unwise
to
assume
that
any
temperature
regime
that
does
not
form
a
blockage
to
migration
or
cause
direct
lethality
will
fully
protect
migrating
steelhead.

Where
fish
may
hold
in
waterbody
segments
for
long
periods
of
time
prior
to
spawning,
as
is
common
for
spring
and
summer
run
steelhead,
daily
maximum
temperatures
should
not
exceed
the
range
previously
identified
as
fully
protective
for
juvenile
rearing.
At
this
point
in
the
life­
stage
of
the
adults
when
feeding
is
not
occurring,
even
cooler
waters
would
be
preferable.
Based
on
the
preceding,
it
is
estimated
that
to
fully
protect
the
adult
migration
of
steelhead
trout
the
7­
day
average
of
the
daily
maximum
temperatures
should
not
exceed
17­
18
°
C.
Single
daily
maximum
temperatures
should
not
exceed
21­
22
°
C.

Cutthroat
Trout
Gresswell
(
1995)
suggests
that
cutthroat
generally
migrate
when
temperatures
approach
5
°
C
(
citing
Varely
and
Gresswell,
1988,
Byorth,
1990,
and
Thurow
and
King,
1994).
In
one
cutthroat
stream
into
Yellowstone
Lake
maximum
daily
water
temperatures
at
the
time
of
peak
spawning
ranged
from
10­
14.2
°
C
over
a
13
year
period
(
USFWS,
unpubl.
data;
as
cited
in
Gresswell,
1995).

Migrations
of
resident
and
potamadromous
cutthroat
trout
occupy
largely
the
same
habitat
used
for
juvenile
rearing.
Thus
for
these
forms,
there
is
little
specific
basis
for
establishing
a
specific
migration
criteria
higher
than
the
fully
protective
range
identified
for
juvenile
rearing.
The
anadromous
forms
of
cutthroat,
unlike
the
Pacific
salmon,
may
make
numerous
journeys
to
the
marine
waters
and
back
again.
Thus
they
may
need
to
repeatedly
pass
through
any
suboptimal
temperature
regime,
however,
since
they
may
feed
on
their
return
migrations
through
fresh
waters,
they
may
be
subject
to
less
sublethal
stress
effects
than
the
Pacific
salmon.
It
is
recommended
that
any
standard
applied
to
protect
the
anadromous
migration
of
the
Pacific
salmon
also
be
applied
to
cutthroat
trout.
Such
a
temperature
value
Page
88
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
would
likely
be
protective
if
the
7­
day
average
of
the
daily
maximum
temperatures
seldom
exceeded
17­
18
°
C
and
the
single
daily
maximum
temperature
does
not
exceed
21­
22
°
C.

Table
4.16.
Barriers
to
migration
of
salmon
and
trout:

Species
Barrier
to
Migration
Comment
Author
Chinook
21­
22
(
21.5)
Stabler,
1981;
Bumgarner
et
al.,
1997;
Hallock,
Elwell,
and
Fry,
1970;
Thompson,
1945,
Ratliff,
1977,
Fish
and
Hanavan,
1948,
Major
and
Mighell,
1967,
and
Alabaster,
1988
18.9
(
maximum)
Partial
blockage
associated
with
low
(
5ppm)
oxygen
levels
Hallock
et
al.,
1970
17.2:
21.7
and
22.2:
26.1
Temperature
differences
that
caused
blockage
Thompson,
1945
9­
11
(
change)
Temperature
differences
that
formed
blockage
Gray,
1990
Sockeye
21
Migration
ceases
Quinn
and
Adams,
1996
Congregated
in
cold
tributaries
when
main
stem
reached
21.7­
23.9
Fish
and
Hanava,
1948
21.1
Blocked
migration
into
tributary
unless
temperatures
were
falling
Major
and
Mighell,
1966
22.8
(
daily
average)
Bulk
of
migration
occurred
below
22.2
and
surges
occurred
when
temperatures
fell
to
below
21.1
Hatch
et
al.,
1992
23.9
Barrier
until
tributary
temperature
declined
to
nearly
21.1
Snyder
and
Blahm,
1971
21
Barrier
for
entry
into
tributary
Strickland,
1967
Steelhead
21.7­
23.9
(
22.8)
Congregated
in
cold
tributaries
when
main
stem
reached
21.7­
23.9
Fish
and
Hanava,
1948
20­
21
(
20.5)
Average
temperature
causing
blockage
when
fish
were
coming
from
water
of
13­
14
Ratliff,
Personal
communication
21
Stabler,
1981
Temperatures
of
21­
22
°
C
are
commonly
cited
as
creating
barriers
to
migration.
While
some
researchers
have
found
that
fish
will
travel
through
waters
as
warm
as
24
°
C,
this
is
generally
uncommon.
Temperatures
below
21
°
C
have
also
been
noted
as
causing
blockages,
but
again
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
89
Washington's
Surface
Water
quality
Standards
this
is
generally
uncommon
and
has
been
in
association
with
fish
encountering
significant
changes
in
temperature
or
encountering
high
temperatures
in
combination
with
low
(
stressful)
oxygen
levels.
Most
of
the
authors
reviewed
were
not
clear
about
the
metric,
but
most
appeared
to
be
referring
to
the
daily
average
temperature.
Some,
however,
may
have
been
referring
to
the
absolute
temperature
at
the
time
of
measurement,
and
thus
would
be
assessing
something
closer
to
the
daily
maximum
temperature.
To
bound
the
estimate
on
the
temperatures
creating
migratory
blockages,
the
commonly
cited
literature
range
for
blockage
should
be
treated
both
as
potentially
related
to
the
daily
maximum
temperature
or
the
daily
average
temperature.
This
would
result
in
the
estimate
that
the
7DADMax
temperature
threshold
that
would
generally
prevent
migration
blockages
would
occur
within
the
ranges
of
20.05­
21.05
°
C
or
23.6­
24.6
°
C,
respectively.
This
would
create
a
potential
range
within
which
migration
blockages
may
occur
of
a
7DADMax
of
20.05­
24.6
°
C
(
22.1
°
C).
The
acute
nature
of
blockages,
the
undescribed
temperature
metrics
reported
by
most
authors,
and
the
general
trait
of
larger
rivers
such
as
the
Columbia
River
being
more
thermally
stable
suggests
the
upper
range
of
this
estimate
may
underestimate
the
effects
of
temperature
in
the
field.

While
barriers
to
migration
have
a
direct
effect
on
adult
migrants,
warm
temperatures
can
harm
migrants
in
many
other
important
ways.
These
include
prespawning
mortality,
decreased
migratory
performance,
reduced
fecundity
and
egg
viability,
and
increased
disease
rates
that
affect
both
the
spawners
and
their
offspring.
For
this
reason
any
criteria
set
to
protect
adult
migration
should
consider
more
than
just
the
temperature
that
creates
a
barrier
to
migration.

Average
daily
temperatures
of
15.5­
20
°
C
have
been
most
frequently
associated
with
prespawning
mortality,
thermal
stress,
and
reduced
reproductive
success
in
adult
migrants
both
in
laboratory
studies
and
natural
streams.
Concerns
over
the
effect
of
temperature
on
egg­
carrying
females
was
discussed
previously
in
the
section
on
reproductive
success.
Some
examples
are
included
in
this
evaluation
of
migratory
effects
that
represent
ambient
river
studies
conducted
at
the
lower
reaches
of
main
stem
rivers.
This
is
because
these
studies
better
fit
with
the
concept
of
protecting
migratory
pathways
than
studies
done
specifically
on
ripe
females.
The
studies
examined
here
are
typically
expressed
as
a
daily
average
exposure,
which
would
be
approximately
equal
to
a
7DADMax
of
18.1­
22.6
°
C
(
20.4
°
C).
Considering
the
lethal
endpoints
under
consideration
here,
it
is
suggested
that
only
the
lower
portion
of
this
range
should
be
considered
potentially
acceptable.
For
these
reasons,
it
is
concluded
that
to
protect
migrating
fish
from
prespawning
losses
in
main
stem
rivers
the
7DADMax
temperatures
should
not
exceed
18.1­
20.4
°
C
(
19.25).
It
is
important
to
note
that
females
carrying
ripe
eggs
would
be
expected
to
have
egg
losses
at
these
temperatures,
and
so
it
is
important
not
apply
this
temperature
estimate
in
headwater
rivers
where
migrating
fish
are
likely
to
be
carrying
ripe
eggs.
Page
90
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Table
4.17.
Migratory
effects
to
salmon
and
trout
other
than
barriers.

Species
Prespawning
Mortality
or
Egg
Loss
Migration
Criteria
Estimates
Migration
Stressors
Comments
Author
Chinook
20
(
mean)
23
(
max)
River
temperature
associated
with
high
(
25%)
losses
Mcdonald
et
al.,
2000
18­
20
Thermo­
regulatory
behavior
in
sub­
yearling
fall
Chinook
at
18­
20
and
heat
shock
proteins
within
several
hours
at
20
Sauter
and
Maule,
1997
7.2­
15.6
Piper
et
al.,
1982
3.3­
13.3
(
spring)
13.9­
20
(
summer)
10.6­
19.4
(
fall)
Migration
recommendation
for
different
runs
Bell,
1973
Coho
7.2­
15.6
Beschta
et
al.,
1987
>
20
Reduced
quality
and
rapid
deterioration
of
eggs
in
utero
in
migrants
Flett
et
al.,
1996
16.7
(
no
effect)
No
apparent
adverse
effects
to
eggs
in
utero
Bouck
et
al.,
1970
Chum
8.3­
15.6
Beschta
et
al.,
1987
Sockeye
16.2
Depletion
of
fat
reserves
and
reproductive
organ
abnormalities
at
16.2
Bouck,
1977
17.5­
19.5
(
maximum)
High
prespawning
mortalities
in
river.
Temperatures
greater
than
15.5
(
daily
ave)
higher
mortality
from
columnaris
disease
Gilhousen,
1990
20
(
average)
20
(
average)
Mean
daily
river
temperatures
above
20
resulted
in
signs
of
significant
stress,
poor
spawning
success,
prespawning
mortality,
and
poor
egg
quality
Mcdonald
et
al.,
2000
19
(
constant)
Laboratory
exposure
over
two
weeks
at
19
depressed
maturation,
but
15
did
not.
Clarke,
Personal
comm.
18­
22
(
mean
daily)
Threshold
of
Fraser
Rv.
at
mouth
for
successful
reproduction
Mcdonald,
2000
Evaluating
Standards
for
Protecting
Aquatic
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in
Page
91
Washington's
Surface
Water
quality
Standards
Holding
of
Adult
Spawners:
It
is
important
to
recognize
that
many
stocks
of
salmon
do
not
migrate
directly
through
to
their
spawning
streams,
but
instead
hold
in
upper
main
stem
rivers
throughout
the
warmest
period
of
the
summer
until
the
fall
when
they
ascend
the
tributaries
and
begin
spawning.
This
section
has
not
included
an
evaluation
of
the
temperature
requirements
of
these
holding
areas.
Because
adult
fish
do
not
feed
while
they
are
holding,
maintaining
cold
waters
are
expected
to
be
especially
important
to
reduce
metabolic
demands
and
stress
in
areas
used
for
over­
summer
holding.
It
is
concluded
that
the
temperature
determined
previously
for
the
protection
of
juvenile
rearing
(
7DADMax
16.5
°
C)
also
be
considered
for
application
to
these
holding
areas,
at
a
minimum.
Or
alternatively,
temperatures
should
be
maintained
at
levels
determined
previously
to
prevent
prespawning
losses
to
eggs
(
7DADMax
14.2
°
C).

vi)
Lethality
to
Adults
and
Juveniles
Constant
Laboratory
Exposure
Studies:

Chinook
Salmon
Beacham
and
Withler
(
1991)
transferred
juvenile
chinook
salmon
from
14
°
C
saline
water
to
a
series
of
test
temperatures
and
noted
genetic
differences
in
the
resistance
times
of
southern
versus
northern
stocks.
They
also
found
that
55%
mortality
occurred
within
three
days
after
transfers
to
saline
waters
with
temperatures
as
low
as
20.3­
21.5
°
C.
They
found
87%
mortality
occurred
within
2
days
at
22.4
°
C.
In
tests
using
adult
"
jack"
chinook
salmon
the
authors
established
an
upper
incipient
lethal
temperature
of
21­
22
°
C.
Orsi
(
1971;
as
cited
in
CDWR,
1988)
found
that
50%
mortality
in
fingerlings
acclimated
to
15.6
°
C
occurred
within
48
hours
at
21.1
°
C,
however,
by
slowly
acclimating
fish
to
21.1
°
C
the
author
raised
the
lethal
endpoint
to
24.7
°
C.
Becker
(
1973)
noted
that
tests
conducted
with
Jack
chinook
salmon
produced
50%
mortality
at
21­
22
°
C
(
this
was
likely
the
same
tests
conducted
in
Coutant,
1970,
in
which
case
the
acclimation
was
at
best
the
prevailing
temperature
of
the
Columbia
River).
Brett
et
al.
(
1982)
found
that
21.5
°
C
was
the
lethal
limit
of
spring
chinook
acclimated
previously
at
10
°
C
and
that
minor
increases
in
mortality
(
up
to
5%)
occurred
at
20
°
C.
Brett
(
1956)
found
that
acclimations
of
5,
10,
15,
and
20
°
C
produced
50%
mortality
at
test
temperatures
of
21.5,
24.3,
25,
and
25.1
°
C.
Brett
et
al.
(
1982)
established
the
lethal
level
to
be
25
°
C
at
an
acclimation
of
20
°
C.
Snyder
and
Blahm
(
1971),
however,
reported
no
mortality
in
chinook
subjected
to
a
change
from
10
to
21.1
°
C
over
a
3­
day
test.

Some
authors
have
noted
temperatures
that
result
in
almost
instantaneous
lethality
to
chinook
salmon.
Orsi
(
1971;
as
cited
in
CDWR,
1988)
found
that
fingerlings
acclimated
to
21.1
°
C
suffered
complete
mortality
when
exposed
to
31.1
°
C
water
for
4­
6
minutes,
and
50%
mortality
occurred
in
fish
acclimated
to
18.3
°
C
and
exposed
to
28.3
°
C
from
4­
6
minutes.
In
a
study
by
Snyder
and
Blahm
(
1971)
a
temperature
of
26.7
°
C
has
resulted
in
mortalities
beginning
after
just
100
seconds
of
exposure
and
complete
mortality
after
4
minutes,
while
at
Page
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Evaluating
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Standards
32.2
°
C
it
only
takes
4
seconds
for
mortality
to
begin
and
complete
mortality
after
11
seconds.
Further,
Gray
(
1990)
found
that
temperature
plumes
above
25.1
°
C
caused
spasmodic
muscle
contractions
in
passing
chinook
salmon.
Gray
(
1990)
cited
research
showing
that
juvenile
salmonids
are
more
susceptible
to
predation
at
10­
20%
of
the
thermal
dose
causing
loss
of
equilibrium.

Coho
Salmon
In
constant
exposure
tests,
the
upper
lethal
levels
producing
50%
mortality
in
juvenile
coho
was
25
°
C
at
an
acclimation
temperature
of
20
°
C
(
DeHart,
1974;
Brett,
1956),
but
at
an
acclimation
of
5
°
C,
Brett
(
1956)
found
the
lethal
level
declined
to
22.9
°
C.
McGeer,
Baranyi,
and
Iwama
(
1991)
exposed
stocks
from
six
hatcheries
to
a
1
°
C/
hour
increase
in
temperature
and
found
the
point
of
50%
mortality
ranged
from
23.8­
24.4
°
C.
While
all
test
fish
survived
up
to
23
°
C
none
survived
beyond
25.5
°
C.
While
juvenile
coho
are
reasonably
tolerant
to
short­
term
peaks
in
temperature,
adults
may
be
far
less
tolerant.
Using
migrating
adult
fish
taken
during
the
summer
from
the
Columbia
River,
Coutant
(
1970,
and
Becker,
1973)
determined
the
lethal
limit
to
be
21­
22
°
C.

Chum
Salmon
Chum
acclimated
to
cold
waters
(
5
°
C)
have
an
upper
lethal
temperature
of
21.8
°
C,
which
increase
to
22.6
and
23.1
°
C
at
acclimation
temperatures
of
10
and
15
°
C
(
Brett,
1956).
In
work
by
Snyder
and
Blam
(
1971)
it
was
found
that
50%
mortality
occurred
in
less
than
50
minutes
to
a
test
population
transferred
from
15.6
°
C
to
26.7
°
C.
A
transfer
from
15.6
°
C
to
29.4
°
C
resulted
in
50%
mortality
in
only
60
seconds,
and
at
32.2
°
C
it
only
required
15
seconds
to
cause
100%
mortality.
Lethal
low
temperatures
range
from
6.5
°
C
and
4.7
°
C
at
acclimation
temperatures
of
20
°
C
and
15
°
C,
to
0.5
°
C
at
a
10
°
C
acclimation.

Pink
Salmon
Brett
(
1952)
found
that
pink
salmon
could
not
be
acclimated
to
24
°
C
and
were
unable
to
survive
for
one
week
at
25
°
C.
Brett
(
1952)
found
that
at
acclimations
of
5
°
C
the
LT50
occurred
within
one
hour
of
exposures
to
22.5
°
C
and
23
°
C.
With
acclimation
to
10
°
C
the
LT50
also
occurred
at
22.5
°
C,
and
at
an
acclimation
of
20
°
C,
it
occurred
at
23.9.

Sockeye
Salmon
Bouck
and
Chapman
(
1975)
found
that
adult
sockeye
could
not
survive
long
periods
at
20
°
C
or
22
°
C,
and
determined
that
the
LT50
for
these
temperatures
occurred
at
11.7
days
and
3.2
days,
respectively.
At
the
lower
acclimation
temperatures
of
5
and
10
°
C,
Brett
found
that
juvenile
sockeye
had
lethal
levels
of
22.2
°
C
and
23.4
°
C
in
a
week­
long
test.
At
acclimations
between
15­
23
°
C,
the
LT50s
for
juvenile
sockeye
were
variable
within
the
range
of
24­
24.8
°
C
(
Brett,
1952;
Beschta
et
al.,
1987;
Servizi
and
Jensen,
1977).
Brett
(
1952)
determined
the
lower
acutely
lethal
temperatures
for
juvenile
sockeye
salmon.
He
found
that
Evaluating
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Page
93
Washington's
Surface
Water
quality
Standards
at
acclimations
of
15
and
20
°
C
the
temperatures
that
produced
50%
mortality
in
a
one
week
test
were
4.1
and
4.7
°
C,
and
that
fish
acclimated
to
10
°
C
had
a
lower
lethal
level
of
3.1
°
C.

Steelhead
and
Rainbow
Trout
In
evaluating
the
effect
of
high
water
temperature
on
steelhead,
Nielsen
et
al.
(
1994)
found
that
the
upper
lethal
temperature
is
approximately
24
°
C
for
juvenile
steelhead.
Redding
and
Schreck
(
1979)
subjected
juvenile
steelhead
previously
acclimated
to
12
°
C
to
a
rapid
rise
(
6.25
hours)
to
26.5
°
C
where
it
was
maintained
for
the
duration
of
the
test.
All
fish
died
within
20.5
hours.
In
a
separate
test
the
temperature
was
held
at
26
°
C,
and
all
fish
died
within
31
hours.
Coutant
(
1970;
and
Becker,
1973),
however,
examined
the
upper
lethal
temperatures
for
adult
steelhead
taken
at
peak
migrating
temperatures
from
the
Columbia
River
in
Washington.
Coutant
concluded
from
his
work
that
the
incipient
lethal
temperature
for
migrating
adult
steelhead
was
closer
to
21­
22
°
C.
Most
laboratory
studies
use
juveniles
of
the
larger
species
of
fish
due
to
the
difficulty
of
handling
adult
salmon
and
steelhead
in
laboratory
tanks.
It
has
been
noted
previously
that
adult
Pacific
salmon
may
have
lower
incipient
lethal
levels
than
that
for
juveniles,
and
it
could
be
that
this
relationship
may
hold
true
for
steelhead
as
well.

Temperatures
as
low
as
23
°
C
have
been
found
to
produce
50%
mortality
(
LT50)
in
rainbow
trout
with
a
week's
constant
exposure
in
fish
previously
acclimated
to
very
cold
(
4
°
C)
waters
(
Sonski,
1982;
Threader
and
Houston,
1983,
as
cited
in
Taylor
and
Barton,
1992),
with
the
lethal
temperature
rising
to
24
°
C
in
moderately
cold
water
(
6­
11
°
C)
acclimated
fish
(
Black,
1953;
Stauffer
et
al.,
1984;
Bidgood,
1980,
as
cited
in
Taylor
and
Barton,
1992).
However,
at
most
acclimation
temperatures
likely
to
be
encountered
during
the
spring
through
fall
seasons
(
12­
20
°
C)
lethal
levels
are
consistently
in
the
range
of
25­
26
°
C
(
Bidgood
and
Berst,
1969;
Hokanson
et
al.,
1987).
With
cautious
acclimation
to
temperatures
in
the
range
of
23­
24
°
C,
rainbow
trout
may
not
experience
LT50
level
effects
until
temperatures
are
held
for
a
week
at
26
°
C
(
Charlon
et
al.,
1970,
as
cited
in
Grande
and
Anderson,
1991).
Even
with
careful
acclimation,
27
°
C
results
in
high
or
complete
mortality
in
less
than
24
hours
(
Charlon,
Barbier,
and
Bonnet,
1970),
and
temperatures
of
29­
30
°
C
result
in
50%
mortality
in
periods
of
1­
2
hours
(
Kaya,
1978;
Craigie,
1963,
and
Alabaster
and
Welcomme,
1962;
as
cited
in
Taylor
and
Barton,
1992).
Some
authors
conducted
critical
thermal
maximum
(
CTM)
tests
on
rainbow
trout.
In
this
type
of
test,
water
temperatures
are
continuously
increased
at
a
rapid
rate
until
the
test
fish
either
loose
equilibrium
or
die
completely.
CTM
values
at
prior
acclimations
of
10
to
25
°
C
ranged
from
27.6
to
32
°
C,
respectively,
in
testing
by
Myrick
and
Cech
(
2000).

Cutthroat
Trout
Heath
(
1963)
subjected
sea­
run
cutthroat
to
a
cyclic
temperature
regime
of
10­
20
°
C
and
calculated
a
critical
thermal
maximum
(
CTM)
of
29.77.
At
constant
acclimations
of
10,
15,
and
20
°
C,
the
corresponding
CTM
values
were
27.63,
29.06,
and
29.88
°
C.
De
Staso
and
Rahel
(
1994)
reported
at
CTM
of
28
°
C
at
some
unknown
acclimation
temperature.
Pauley
et
al.
(
1989)
cite
research
concluding
that
equilibrium
and
ability
to
swim
is
lost
with
a
rise
in
Page
94
Evaluating
Standards
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Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
temperature
to
28­
30
°
C.
Feldmuth
and
Eriksen
(
1978,
as
cited
in
McIntyre
and
Rieman,
1995)
estimated
that
the
CTM
for
westslope
cutthroat
trout
was
27.1
°
C,
a
value
lower
than
those
estimated
for
brook
trout
(
29.8
°
C),
brown
trout
(
29.6
°
C)
and
rainbow
trout
(
31.6
°
C).
Golden
(
1976)
tested
the
temperature
tolerance
of
zero
age
coastal
cutthroat
stocks
from
western
Oregon.
In
CTM
tests
where
the
temperature
is
rapidly
increased
until
the
fish
die
or
loose
equilibrium,
acclimation
to
10
and
23
°
C
yielded
CTM
values
of
28.03
and
30.62
°
C.
At
a
fluctuating
equilibriums
of
7.8­
10
°
C
and
13­
23
°
C
the
CTM
values
were
27.64
and
30.31
°
C.

Golden
(
1978)
tested
the
lethality
of
significantly
fluctuating
temperature
regimes
with
cutthroat
trout.
Incipient
lethal
levels
(
ILL)
were
25.5
°
C
and
25.7
°
C
for
fish
acclimated
to
23
°
C
and
a
fluctuating
regime
of
13­
23
°
C,
respectively.
Golden
estimated
losses
in
one
week
of
10%
or
less
at
a
fluctuating
cycle
of
13­
27
°
C.

Vigg
and
Koch
(
1980)
measured
the
lethal
limits
of
two
stocks
of
Lahontan
cutthroat
in
three
water
types
and
found
that
alkalinity
profoundly
influences
the
results.
In
waters
with
alkalinity
of
1,487
mg/
l
the
lethal
range
was
18.5­
20.2
°
C,
in
waters
with
alkalinity
of
357
mg/
l
the
range
was
20.2­
21.1
°
C,
and
at
an
alkalinity
of
69
mg/
l
the
lethal
range
was
21.8­
23.0
°
C
for
the
two
species.
Kramer
(
1975;
as
cited
in
Vigg
and
Koch,
1980)
found
they
were
able
to
hold
a
Humboldt
River
strain
of
cutthroat
at
24
°
C
for
two
weeks
without
any
mortality,
but
with
feeding
inhibition.

Dickerson
et
al.
(
1999,
and
unpublished
data
as
cited
in
Dunham,
1999)
tested
the
lethal
tolerance
levels
of
Lahontan
cutthroat.
Survival
was
100%
at
24
°
C
but
declined
to
35%
at
26
°
C.
At
28
°
C
mortality
was
complete
in
48
hours.

Were
it
not
for
the
studies
of
Lahontan
cutthroat
trout
by
Vigg
and
Koch
(
1980),
it
would
be
easy
to
conclude
that
cutthroat
trout
would
only
be
expected
to
have
50%
mortality
(
LT50)
over
a
one
week's
exposure
to
constant
temperatures
above
approximately
24
°
C.
While
it
is
clear
that
in
many
cases
no
mortality
would
occur
as
a
consequence
of
short
term
exposure
to
infrequent
daily
maximum
temperatures
as
high
as
26
°
C,
the
data
of
Vigg
and
Koch
(
1980)
suggest
more
caution
may
be
warranted.
They
calculated
LT50
values
of
20.2­
21.1
°
C
and
22­
23
°
C
using
two
stocks
of
cutthroat
and
two
water
sources
with
alkalinity
levels
comparable
to
what
is
regularly
found
in
Washington.
More
work
is
needed
to
clear
up
the
question
of
whether
native
stocks
demonstrate
lower
lethal
thresholds
in
natural
waters
with
high
alkalinity
levels,
prior
to
assuming
the
data
of
Vigg
and
Koch
(
1980)
to
be
anomalous.

Table
4.18.
The
following
summarizes
the
results
of
laboratory
tests
subjecting
salmon
and
trout
to
constant
test
temperatures.
The
column
on
acclimation
provides
the
temperature
to
which
the
fish
were
acclimated
to
prior
to
being
moved
to
the
test
temperature,
and
the
endpoint
reported
is
the
percent
of
mortality
that
occurred
at
each
lethal
temperature
level
(
LT).

Author
or
Study
Species
Acclimation
Temp.
(
°
C)
Temperatur
e
of
Test
Time
to
Endpoint
Endpoint
Reported
Comment
Evaluating
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Protecting
Aquatic
Life
in
Page
95
Washington's
Surface
Water
quality
Standards
(
°
C)
(
seconds)
Beacham
and
Withler,
1991
Chinook
14
20.9
259200
LT55
LT55
in
3­
days
with
transfers
to
20.3­
21.5
(
mean
20.9
°
C).
14
22.4
172800
LT87
2
days
14
21.5
604800
LT50
Adults
7­
day
21­
22
(
mean
21.5
°
C).
Orsi,
1971
Chinook
15.6
21.1
172800
LT50
2
days
21.1
24.7
172800
LT50
2
days
18.3
28.3
300
LT50
4­
6
minutes
21.1
31.1
300
LT100
4­
6
minutes
Becker,
1973
and
Coutant,
1970
Chinook
Summer
ambient
21.5
604800
LT50
Adults
7­
day
21­
22
(
mean
21.5
°
C)
Brett,
1982
Chinook
10
21.5
604800
LT50
Spring
run
10
20
604800
LT5
Spring
run
20
25
604800
LT50
Spring
run
Snyder
and
Blahm,
1971
Chinook
10
21.1
259200
LT0
3­
days
­
no
mortality
10
26.7
100
LT1
100
seconds
10
26.7
240
LT100
4
minutes
10
32.2
4
LT1
4
seconds
10
32.2
11
LT100
11
seconds
Brett,
1956
Chinook
5
21.5
604800
LT50
10
24.3
604800
LT50
15
25
604800
LT50
20
25.1
604800
LT50
DeHart,
1974
Coho
20
25
604800
LT50
Brett,
1956
Coho
20
25
604800
LT50
Coho
5
22.9
604800
LT50
Becker,
1973
and
Coutant,
1970
Coho
Summer
ambient
21.5
604800
LT50
Adults
7­
day
21­
22
(
mean
21.5
°
C)
Brett,
1956
Chum
5
21.8
604800
LT50
Chum
10
22.6
604800
LT50
Chum
15
23.1
604800
LT50
Snyder
and
Blam,
1971
Chum
15.6
26.7
3000
LT50
50
minutes
15.6
29.4
60
LT50
60
seconds
15.6
32.2
15
LT100
15
seconds
Brett,
1952
Pink
5
22.5
3600
LT50
1­
hour
LT50
at
both
22.5
and
23
°
C.
10
22.5
604800
LT50
20
23.9
604800
LT50
24
Could
not
be
acclimated
to
24
°
C
or
survive
one­
week
at
25
°
C.
Brett,
1952
Sockeye
5
22.2
604800
LT50
10
23.4
604800
LT50
Bouck
and
Chapman,
1975
Sockeye
20
1010880
LT50
Adults
­
11.7
days
22
276480
LT50
Adults
 
3.2
days
Brett,
1952;
Servizi
and
Jensen,
1977
Sockeye
15­
23
24­
24.8
(
24.4)
604800
LT50
Page
96
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Nielsen,
Lisle,
and
Ozaki,
1994
Steelhea
d
24
604800
LT50
Upper
lethal
temperature
for
juveniles
Redding
and
Schreck,
1979
Steelhea
d
12
26
73800
LT100
Complete
mortality
in
20.5
hours
with
rapid
rise
to
test
temperature
12
26
111600
LT100
Complete
mortality
in
31
hours
with
immediate
transfer
to
test
temperature
Becker,
1973
and
Coutant,
1970
Steelhea
d
Summer
ambient
21.5
604800
LT50
Adults
7­
day
21­
22
(
mean
21.5
°
C)
Sonski,
1982
Rainbow
Trout
4
23
604800
LT50
Threader
and
Houston,
1983
Rainbow
Trout
4
23
604800
LT50
Black,
1953
Rainbow
Trout
6­
11
24
604800
LT50
Stauffer
et
al.,
1984
Rainbow
Trout
6­
11
24
604800
LT50
Bidgood,
1980
Rainbow
Trout
6­
11
24
604800
LT50
Bidgood
and
Berst,
1969
Rainbow
Trout
12­
20
25­
26
604800
LT50
Hokanson
et.
al.,
1987
Rainbow
Trout
12­
20
25­
26
604800
LT50
Charlon
et
al.,
1970
Rainbow
Trout
23­
24
26
604800
LT50
Charlon,
Barbier,
and
Bonnet,
1970
Rainbow
Trout
High
27
86400
LT50­
LT100
High
to
complete
mortality
in
less
than
24
hours
Kaya,
1978
Rainbow
Trout
High
29­
30
3600­
7200
LT50
1­
2
hours
Craigie,
1963
Rainbow
Trout
High
29­
30
3600­
7200
LT50
1­
2
hours
Alabaster
and
Welcomme,
1962
Rainbow
Trout
High
29­
30
3600­
7200
LT50
1­
2
hours
Golden,
1978
Cutthroat
Trout
23
25.5
604800
LT50
13­
23
25.7
604800
LT50
Fluctuating
acclimation
regime
Vigg
and
Koch,
1980
Cutthroat
Trout
­
Lahontan
18.5­
20.2
(
19.35)
604800
LT50
Alkalinity
1,487
mg/
l
 
two
stocks
of
Lahontan
and
two
water
sources
20.2­
21.1
(
20.65)
604800
LT50
Alkalinity
357
mg/
l
 
two
stocks
of
Lahontan
and
two
water
sources
21.8­
23.0
(
22.4)
604800
LT50
Alkalinity
69
mg/
l
 
two
stocks
of
Lahontan
and
two
water
sources
Kramer,
1975
Cutthroat
Trout
24
129600
LT0
No
mortality
in
two
weeks
using
Humboldt
River
strain
of
cutthroat
Dickerson
et
al.,
1999
Cutthroat
Trout
­
Lahontan
24
604800
LT0
Lahontan
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
97
Washington's
Surface
Water
quality
Standards
26
604800
LT35
Lahontan
28
172800
LT100
2­
days
 
complete
mortality
in
Lahontan
Typical
Constant
Laboratory
Exposures
Producing
Fifty­
Percent
Mortality:

The
results
from
constant
temperature
testing
that
resulted
in
fifty­
percent
mortality
in
3­
12
days
of
exposure
(
shown
in
the
table
above)
can
be
used
to
assess
weekly
temperatures
that
will
not
result
in
excess
lethality.
Of
the
38
tests
assessed,
90%
of
the
values
were
between
20.5­
25.6
°
C,
the
inter
quartile
range
was
21.9­
24.6
°
C,
and
the
median
was
23.1
°
C.
The
5th
and
95th
percentiles
for
the
distribution
are
used
herein
to
calculate
expected
thresholds
to
avoid
the
possibility
of
using
anomalous
extreme
data
points.
Converting
from
an
LT50
to
an
LT1
estimated
mortality
rate
(
by
subtracting
2
°
C)
suggests
a
constant
exposure
to
a
temperature
above
18.5­
23.6
°
C
for
a
period
of
a
week
or
more
may
result
in
the
direct
mortality
to
salmon
and
trout.
Converting
the
constant
laboratory
test
exposure
estimate
to
a
field
exposure
regime
is
especially
difficult
for
this
form
of
lethality
data.
Because
the
exposure
times
necessary
to
cause
lethality
reduce
as
temperatures
continue
to
increase
above
the
incipient
lethal
limit
it
is
not
appropriate
to
treat
the
laboratory
results
as
weekly
average
temperatures.
However,
using
the
results
directly
as
7DADMax
thresholds
would
overstate
the
risks
since
the
time
spent
above
the
threshold
will
be
only
a
small
portion
of
each
day
while
the
original
tests
exposed
fish
to
these
temperatures
for
24­
hours
per
day.
Thus
a
more
defensible
estimate
would
be
one
that
falls
between
these
two
approaches
rather
than
using
the
specific
values
contained
in
either.
Treating
the
constant
laboratory
exposure
as
a
weekly
average
temperature
would
result
in
the
estimate
of
a
7DADMax
of
21.68­
26.78
°
C
(
24.23),
and
using
it
directly
would
result
in
the
estimate
of
a
7DADMax
of
18.5­
23.6
°
C
(
21.05).
Using
these
two
approaches
to
bound
the
estimate
results
in
the
expectation
that
mortality
will
occur
as
7DADMax
temperatures
exceed
21.05­
24.23
with
the
best
estimate
expected
to
be
the
midpoint
of
the
distribution
(
22.64
°
C).

Regression
of
Constant
Laboratory
Exposures
Producing
Fifty­
Percent
Mortality:

Although,
the
relationship
between
acclimation
temperatures
and
test
temperatures
resulting
in
50%
mortality
is
strong
within
individual
studies,
the
relationship
is
weak
when
all
of
the
various
studies
are
combined
for
analysis.
This
is
illustrated
in
figure
4.1
below.

Figure
4.1.
Combined
lethality
data
for
all
salmon
and
trout
species
(
based
on
7­
day
LT50
estimates)
from
Table
4.18
above.
Page
98
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Acclimation
Combined
LT50
Estimated
LT1
with
Temperature
(
°
C)
for
all
Salmonids(
°
C)
NAS
Adjustment
(
°
C)
10
22.50
20.50
12
22.93
20.93
14
23.37
21.37
16
23.79
21.79
18
24.23
22.23
20
24.66
22.66
The
regression
equation
in
figure
4.1
above,
can
be
used
to
predict
temperatures
at
which
significant
lethality
(
50%)
would
be
expected
with
a
constant
exposure
for
7­
days
at
various
prior
acclimation
temperatures.
Since
the
results
are
based
on
50%
mortality,
they
should
be
adjusted
to
better
reflect
temperatures
that
would
not
directly
kill
fish.
The
USEPA
recommends
that
states
subtract
2
°
C
from
LT50
results
to
obtain
an
estimate
that
would
result
in
1%
or
less
mortality
(
LT1)
under
the
same
exposure
assumptions.
Fish
acclimate
to
ambient
temperatures
approximately
equal
to
the
average
temperature,
and
often
fish
that
are
migrating
from
the
ocean
or
from
cooler
tributary
systems
will
not
be
fully
acclimated
to
warm
prevailing
temperatures
in
main
stem
rivers.
For
this
reason,
acclimation
temperatures
from
12­
16
°
C
may
best
represent
the
condition
of
migrating
fish
during
the
summer
period
(
yielding
LT1
estimates
of
20.93­
21.79
°
C).
Juvenile
fish
that
reside
within
a
system,
may
be
acclimated
to
warmer
temperatures.
However,
even
resident
fish
will
not
be
as
fully
acclimated
as
those
used
in
tests
to
estimate
an
ultimate
incipient
lethal
temperature
(
the
absolute
highest
LT50
that
can
be
produced
with
slow
and
careful
laboratory
acclimation).
Based
on
the
foregoing,
a
constant
temperature
exposure
for
7
days
would
be
expected
to
cause
no
more
than
LT1
levels
of
mortality
at
temperatures
of
20.93­
21.79
°
C.
Converting
the
constant
laboratory
test
exposure
to
a
field
exposure
regime
is
especially
difficult
for
this
form
of
lethality
data.
Because
the
exposure
times
necessary
to
cause
lethality
reduce
as
temperatures
continue
to
increase
above
the
incipient
lethal
limit
it
is
not
appropriate
to
treat
the
laboratory
results
as
weekly
average
temperatures.
However,
using
the
results
directly
as
7DADMax
thresholds
would
overstate
the
risks
since
the
time
spent
above
the
threshold
will
be
only
a
small
portion
of
each
day
while
the
original
tests
exposed
fish
to
these
temperatures
LT50
Results
for
Salmon
and
Trout
at
Constant
Temperature
Exposures
y
=
0.2153x
+
20.351
R2
=
0.3206
20
21
22
23
24
25
26
27
9
14
19
24
Acclimation
Temperatures
(
C)
Temperature
of
LT50
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
99
Washington's
Surface
Water
quality
Standards
for
24­
hours
per
day.
Thus
the
more
defensible
estimate
would
be
one
that
falls
between
these
two
approaches,
rather
than
containing
values
from
either.
Treating
the
constant
laboratory
exposure
as
a
weekly
average
temperature
would
result
in
the
estimate
of
a
7DADMax
of
24.11­
24.97
°
C
(
24.54)
using
it
directly
would
result
in
the
estimate
of
a
7DADMax
of
20.93­
21.79
°
C
(
21.36).
Using
these
two
approaches
to
bound
the
estimate
results
in
the
estimate
that
mortality
will
occur
as
7DADMax
temperatures
exceed
21.36­
24.54
with
the
best
estimate
expected
to
be
the
mean
of
the
distribution
(
22.95
°
C).

Modeling
Lethality
of
Fluctuating
Temperature
Regimes:

The
concept
of
resistance
time
is
very
important
to
estimating
potential
lethality.
It
is
well
demonstrated
that
it
is
the
time
spent
above
a
lethal
threshold
that
determines
whether
or
not
short
term
lethal
effects
will
occur.
Different
peak
temperatures
(
e.
g.,
22,
24,
27,
30
°
C),
may
all
be
lethal
to
an
organism,
but
the
organism
can
likely
withstand
these
temperatures
for
variable
lengths
of
time.
A
population
of
fish
may
be
able
to
withstand
21
°
C
for
7
days
of
constant
exposure
without
any
mortality,
but
have
50%
of
the
population
die
after
2
days
at
24
°
C.
At
27
°
C
50%
mortality
may
occur
after
less
than
2
hours
of
exposure,
and
at
30
°
C,
complete
mortality
may
occur
in
just
a
few
minutes.

In
considering
the
effect
of
repeated
hot
days,
it
is
important
to
incorporate
the
potential
for
cumulative
effects
over
a
series
of
days.
DeHart
(
1974)
found
that
lethal
effects
occur
in
relation
to
the
area
of
the
temperature
time
curve
that
is
above
a
fish's
incipient
lethal
level
(
ILL);
an
accumulation
of
thermal
effects
occurs
over
periods
of
several
days
when
the
daily
temperature
cycle
fluctuates
above
the
incipient
lethal
level;
and
the
time
above
the
incipient
lethal
level
influences
the
thermal
resistance
time
independent
of
the
lower
temperatures
experienced
in
fluctuating
tests.
In
other
words,
the
ability
of
a
fish
to
resist
a
single
day's
exposure
to
a
lethal
temperature
may
not
be
sufficient,
and
fifteen
minutes
spent
at
4
°
C
over
the
ILL
is
of
more
consequence
than
the
same
time
spent
at
2
°
C
over
the
ILL.

The
results
from
laboratory
lethality
tests
conducted
at
constant
temperatures
can
be
used
to
estimate
lethality
under
fluctuating
temperature
conditions.
This
can
be
done
by
creating
a
simple
model
of
the
time
to
mortality
using
the
laboratory
studies
of
a
duration
of
7­
days
or
less
from
table
4.18
above.
Each
temperature
value
is
assigned
a
lethality
indices
which
represents
the
proportion
of
a
lethal
exposure
that
occurs
within
a
single
day.
Lethal
loading
stress
is
assumed
to
be
cumulative
such
that
the
model
can
predict
the
number
of
consecutive
days
until
50%
mortality
will
occur
under
any
combination
of
diel
temperature
fluctuations.
Because
of
the
great
variability
in
7­
day
LT50
values,
only
a
single
representative
value
is
used
to
represent
this
particular
exposure
period.
A
constant
temperature
of
21.85
°
C
was
chosen
to
represent
an
7­
day
LT50
temperature,
which
represented
the
25th
percentile
of
the
distribution
of
LT50
values.
Alternatively,
23
°
C
was
considered
since
it
represents
both
the
regressed
LT50
value
at
the
lower
expected
acclimation
temperature
and
is
the
median
of
the
distribution
of
LT50
results.
However,
taking
this
alternative
approach
would
have
risked
extrapolating
the
estimate
beyond
50%
of
the
actual
test
endpoints,
and
would
not
well
incorporate
the
observed
pattern
of
adult
fish
showing
greater
lethality
at
similar
temperatures
as
juveniles
who
are
the
subjects
of
most
of
the
tests.
Page
100
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Figure
4.2
below
shows
a
power
curve
regression
of
the
LT50
data
with
a
single
representative
value
included
to
moderate
the
variability
in
the
tests
conducted
for
seven
days.
The
equation
of
the
regression
line
can
be
used
to
predict
fifty
percent
lethality
rates
at
any
given
duration
of
exposure.
This
equation
(
y
=
5E+
28x­
17.402)
is
used
herein
to
model
lethality
under
fluctuating
temperature
environments.

Figure
4.2.
Regression
of
LT50
data
with
a
single
composite
value
for
all
7­
day
tests.

LT50
Results
for
Salmon
and
Trout
Using
a
Single
Representative
7­
Day
Value
­
Power
Curve
y
=
5E+
28x­
17.402
R2
=
0.7541
0
200000
400000
600000
800000
1000000
1200000
19
21
23
25
27
29
31
Test
temperature
Time
to
LT50
(
seconds)

If
it
is
assumed
that
fish
are
acclimated
to
a
series
of
temperatures
(
representing
the
daily
mean
of
a
fluctuating
environment)
then
the
number
of
days
until
fifty­
percent
mortality
would
be
expected
can
be
estimated
(
See
table
4.19
below).
Since
the
7­
day
exposure
tests
were
composited
using
the
25th
percentile
of
the
7­
day
LT50
values,
the
estimates
would
generally
suggest
that
most
(
75%
or
more)
stocks
would
have
a
greater
degree
of
protection.
The
results
of
this
modeled
analysis
suggest
that
at
acclimation
temperatures
in
the
range
of
what
can
be
expected
in
natural
streams
during
the
summer
months
(
15­
20
°
C)
fifty­
percent
mortality
may
occur
to
some
stocks
with
a
7­
day
exposure
at
daily
maximum
temperatures
of
22.13­
22.47
°
C
(
22.3).
This
estimate
is
then
converted
to
a
temperature
range
that
would
be
unlikely
to
result
in
any
mortality
(
an
estimated
LT1)
by
subtracting
2
°
C.
This
results
in
the
estimate
that
direct
mortality
will
be
prevented
if
the
7DADMax
temperature
is
maintained
below
20.13­
20.47
°
C
(
20.3).
Temperature
of
Test
(
°
C)
Time
to
LT50
Endpoint
(
seconds)
29.4
60
28.3
300
26.7
3000
29.5
5400
29.5
5400
29.5
5400
21.1
172800
24.7
172800
20.9
259200
22
276480
21.85
604800
20
1010880
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
101
Washington's
Surface
Water
quality
Standards
Table
4.19.
Modeled
results
showing
the
days
to
LT50
at
various
daily
mean
and
daily
maximum
temperatures
and
assuming
a
4.13
°
C
per
day
change
in
temperature.

Laboratory
Studies
with
Fluctuating
Temperatures:

Coho
Salmon
DeHart
(
1974)
reported
that
a
cyclic
temperature
regime
having
6
hours
a
day
over
25
°
C
produced
50%
mortality
in
1.5
cycles.
In
a
20­
day
test
with
highly
fluctuating
daily
temperatures
(
5­
23
°
C)
having
an
average
of
11
°
C,
however,
Thomas
et
al.
(
1986)
found
no
increase
in
mortality.
Acclimated
age­
0
and
age­
2
fish
did
not
begin
dying
until
the
diel
temperature
range
reached
4­
25
°
C.
Thomas
et
al.
also
reported
that
juvenile
coho
were
able
to
feed
and
grow
in
fluctuating
temperatures
that
approached
their
upper
lethal
limits.
The
studies
of
DeHart
(
1974)
and
Thomas
et
al.
(
1986)
suggest
that
daily
maximum
temperatures
of
25
°
C
produce
a
risk
of
direct
mortality,
the
level
of
that
mortality
may
vary
substantially
but
can
be
as
high
as
50%.
The
levels
of
fluctuations
in
the
Thomas
et
al.
work
were
so
great
as
to
significantly
limit
the
time
fish
spent
above
temperatures
that
can
be
lethal
with
longerterm
exposure.
Converting
the
LT50
estimate
at
25
°
C
to
an
LT1
estimate
by
subtracting
2
°
C
(
USEPA
recommendation),
yields
the
prediction
that
daily
maximum
temperatures
below
23
°
C
should
prevent
direct
lethality
in
coho
salmon
over
short­
term
(
one­
week
or
less)
exposures.
The
safety
of
taking
this
approach
is
also
supported
by
the
direct
evidence
of
Thomas
et
al.
(
1986)
where
daily
maximums
of
23
°
C
(
diel
range
5­
23
°
C)
did
not
result
in
an
increase
in
mortality.
Daily
mean
Daily
Max
Days
to
LT50
15
22.13
6.802
16
22.13
6.81
17
22.13
6.836
18
22.47
6.911
19
22.13
7.12
20
22.13
7.743
Page
102
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Steelhead
and
Rainbow
Trout
Under
fluctuating
temperature
test
conditions,
rainbow
trout
have
been
found
to
experience
50%
mortality
in
a
week
of
daily
cycles
from
21­
27
°
C
(
Lee,
1980).
If
2
°
C
is
subtracted
to
reduce
the
level
of
mortality
from
50%
to
1%
or
less,
this
would
change
the
daily
maximum
temperature
to
25
°
C
to
prevent
direct
mortality.

Cutthroat
Trout
Golden
(
1976)
tested
the
temperature
tolerance
of
zero
age
coastal
cutthroat
stocks
from
western
Oregon.
It
was
found
that
seven
diel
cycles
of
13­
27
°
C
resulted
in
0­
20%
mortality.
However,
by
increasing
the
peak
temperature
in
the
cycle
0.5
°
C
(
13­
27.5
°
C)
mortalities
increased
to
50­
90%
within
1
to
1.5
cycles.
In
CTM
tests
where
the
temperature
is
rapidly
increased
until
the
fish
die
or
loose
equilibrium,
acclimation
to
10
and
23
°
C
yielded
CTM
values
of
28.03
and
30.62
°
C.
At
a
fluctuating
equilibriums
of
7.8­
10
°
C
and
13­
23
°
C
the
CTM
values
were
27.64
and
30.31
°
C.
Golden
(
1978)
tested
the
lethality
of
significantly
fluctuating
temperature
regimes
with
cutthroat
trout.
Incipient
lethal
levels
(
ILL)
were
25.5
°
C
and
25.7
°
C
for
fish
acclimated
to
23
°
C
and
a
fluctuating
regime
of
13­
23
°
C,
respectively.
Golden
estimated
losses
in
one
week
of
10%
or
less
at
a
fluctuating
cycle
of
13­
27
°
C.
Golden
(
1978)
found
that
high
mortality
could
still
occur
with
temperatures
as
high
as
25.5­
25.7
°
C
at
constant
exposures,
even
for
cutthroat
acclimated
to
high
(
23
°
C)
daily
maximum
temperatures.
The
work
of
Golden
(
7976,
1978)
suggests
that
50%
mortality
can
occur
and
temperatures
from
25.5­
27
°
C
even
where
fish
have
been
well
acclimated
to
warm
waters.
After
converting
this
range
to
one
in
which
1%
or
less
(
LT1)
mortality
would
be
expected
(
by
subtracting
2
°
C)
it
would
be
expected
that
direct
mortality
could
be
avoided
by
maintaining
daily
maximum
temperatures
below
23­
25
°
C.
Since
many
of
the
test
fish
were
carefully
acclimated
to
warm
waters
prior
to
testing,
the
upper
end
of
this
range
may
pose
greater
risks
to
migratory
stocks.

Dickerson
et
al.
(
1999,
and
unpublished
data
as
cited
in
Dunham,
1999)
tested
the
lethal
tolerance
levels
of
Lahontan
cutthroat.
Survival
was
100%
at
24
°
C
but
declined
to
35%
at
26
°
C.
At
28
°
C
mortality
was
complete
in
48
hours.
In
a
separate
test
of
fluctuating
(
20­
26
°
C)
and
constant
(
13,
20,
and
23
°
C)
temperatures
no
mortality
occurred.
Dunham
(
1999)
concluded
from
a
review
of
the
literature
that
Lahontan
cutthroat
can
survive
weekly
exposure
to
daily
temperature
fluctuations
of
20­
26
°
C,
including
1­
hour
exposures
to
temperatures
of
up
to
26
°
C.

The
works
of
Dickerson
(
1999)
and
Golden
(
1976,
1978)
can
be
used
to
support
the
position
that
cutthroat
trout
may
be
capable
of
tolerating
daily
maximum
temperatures
as
high
as
23­
26
°
C
(
24.5)
for
up
to
a
week
of
exposure
without
experiencing
direct
mortality.
However,
without
the
non­
indigenous
Lahontan
cutthroat
data
the
estimate
of
lethality
would
change
to
23­
25
°
C
(
24
°
C)
and
should
be
viewed
as
a
more
defensible
estimate
of
lethality
in
Washington's
cutthroat
trout
populations.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
103
Washington's
Surface
Water
quality
Standards
Summary
of
fluctuating
lethality
studies:
In
many
of
the
fluctuating
exposure
studies
reviewed,
the
laboratory
test
water
had
high
diel
fluctuations
(
14
°
C).
These
fluctuations
are
greater
than
what
would
often
occur
in
natural
waters
and
acts
to
significantly
limit
the
amount
of
time
spent
each
day
spent
above
incipient
lethal
levels.
The
studies
also
included
fish
that
were
pre­
acclimated
to
relatively
warm
temperatures,
and
thus
some
natural
migratory
stocks
may
be
at
slightly
higher
risks.
The
works
also
included
a
non­
indigenous
species
of
cutthroat
that
is
widely
found
in
desert
environments
and
which
may
have
different
lethal
thresholds
than
Washington's
native
cutthroat.
Thus
caution
should
be
exercised
in
using
the
results
from
this
line
of
evidence.
Based
on
the
fluctuating
laboratory
tests
cited
above,
daily
maximum
temperatures
above
23­
25
°
C
should
be
considered
capable
of
causing
direct
mortality
to
salmon
and
trout.
Since
even
very
slight
(
0.5
°
C)
temperatures
increases
in
the
daily
maximum
temperature
have
been
shown
to
produce
significant
increases
in
mortality
in
this
range,
these
values
may
best
be
treated
as
single
daily
maximum
thresholds.
However,
the
single
daily
maximum
range
of
23­
25
°
C
can
also
be
converted
to
the
standard
metric
of
a
7DADMax
temperature
(
by
subtracting
0.95
°
C).
This
approach
yields
the
estimate
that
to
avoid
direct
mortality
the
7DADMax
temperature
should
remain
below
22.05­
24.05
°
C
(
23.05).

Field
Studies
of
Lethality:

Chinook
Salmon
The
previously
mentioned
studies
of
lethality
were
predominantly
conducted
in
a
laboratory
environment.
Baker
et
al.
(
1995),
however,
modeled
the
escapement
of
smolts
from
the
lower
Sacramento
River
and
determined
an
upper
lethal
temperature
of
23
°
C.
Burck
(
1993)
used
live­
box
tests
with
juvenile
chinook
salmon,
and
found
that
daily
maximum
temperatures
in
the
range
of
25.5­
26.6
°
C
were
lethal
to
all
of
the
test
fish
after
a
single
daily
cycle
of
exposure
(
daily
minimums
ranged
from
13.3­
16.1
°
C).
Daily
maximums
in
the
range
of
23.8
to
25.5
°
C
(
minimums
ranging
from
11.1­
13.3
°
C)
resulted
in
80%
mortality
over
the
four­
day
test
period.
No
mortality
occurred
in
the
three
controls
that
had
daily
maximum
temperatures
from
14.4­
17.2
°
C.
While
50%
mortality
occurred
in
one
treatment
with
maximums
of
20.5­
21.6
°
C,
no
mortality
occurred
in
four
other
treatments
with
daily
maximums
in
the
range
of
20.0­
22.7
°
C.
We
can
roughly
compare
Burck's
data
with
values
found
in
standardized
laboratory
tests
using
chinook
salmon.
If
we
assume
the
temperature
midway
between
the
daily
maximum
and
the
daily
average
is
the
acclimation
temperature,
then
these
live­
box
tests
generally
demonstrated
that
at
an
acclimation
temperature
of
20.6
°
C
the
lethal
temperature
was
25.6
°
C,
and
at
an
acclimation
of
18.3
°
C
the
lethal
level
was
24.7
°
C.
At
an
acclimation
temperature
of
16.5
°
C,
however,
there
was
inconsistent
but
typically
complete
survival
at
up
to
21.6
°
C.

High
mortality
has
been
observed
in
fish
held
in
rivers
with
daily
maximum
temperatures
of
23.8­
25.5
°
C
over
a
four­
day
period.
Reducing
this
range
by
2
°
C
would
convert
it
to
a
range
for
daily
maximum
temperatures
at
which
mortality
would
not
be
expected.
This
would
yield
the
range
of
21.8­
23.5
°
C
(
22.65).
This
range
was
derived
from
daily
maximum
Page
104
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
temperatures
over
a
four
day
exposure
period.
This
range
can
be
used
to
create
a
range
that
is
based
on
a
7DADMax
temperature
metric.
The
estimate
can
also
be
bounded
by
treating
the
range
as
if
it
were
representing
either
the
7DADMax
temperature
directly
(
7DADMax
21.8­
23.5
°
C)
and
also
as
if
it
represents
the
single
daily
maximum
temperature
(
since
it
may
have
been
the
warmest
day
that
triggered
the
mortality)
and
converting
(
subtracting
0.95
°
C)
that
value
to
a
7DADMax
estimate
(
20.85­
22.55
°
C).
This
approach
suggests
that
direct
mortality
would
be
prevented
by
maintaining
the
7DADMax
temperature
below
20.85­
23.5
°
C
(
22.18).

Steelhead
and
Rainbow
Trout
Sonski
(
1983)
noted
having
success
with
culturing
rainbow
trout
in
ponds
that
that
reached
a
summer
maximum
of
28.9
°
C,
and
Chandrasekaran
and
Subb
Rao
(
1979)
noted
that
rainbow
trout
in
India
were
largely
able
to
survive
in
rearing
ponds
with
months
having
daily
maximum
temperatures
in
the
range
of
26­
29
°
C.
Neither
of
these
works
estimated
lethality
directly
and
cannot
reasonably
be
used
to
suggest
safe
temperature
ranges
for
rainbow
trout.
They
can
however
be
used
to
suggest
that
losses
can
sometimes
be
moderate
(
acceptable
from
a
fish
culturing
standpoint)
at
temperatures
that
reach
highs
of
26­
29
°
C.

Summary
of
field
studies:
Only
two
of
the
four
citations
actually
provide
estimates
of
lethal
temperature
ranges.
These
two
studies
both
can
be
used
to
support
the
estimate
that
lethality
should
be
expected
as
the
7DADMax
temperature
exceeds
20.85­
23.5
°
C
(
22.18).
The
rearing
pond
observations
that
were
also
reviewed
suggest
that
mortality
rates
sometimes
may
not
become
significant
in
cultured
rainbow
trout
even
though
daily
maximum
temperatures
may
reach
26­
29
°
C.

General
Measures
of
Lethal
Stress:

Cutthroat
Trout
Titus
and
Vanicek
(
1988;
as
cited
in
Muoneke
and
Childress,
1994)
reported
that
mortality
of
cutthroat
trout
caught
and
released
by
angling
was
less
than
2%
at
temperatures
below
17
°
C
but
rose
to
49%
as
the
temperature
neared
21
°
C.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
105
Washington's
Surface
Water
quality
Standards
Summary
of
Lines
of
Evidence
on
Lethality:

Table
4.19.
Direct
lethality
to
salmon
and
trout
with
short­
term
exposure
(
7­
days).

Line
of
Evidence
7DADMax
(
°
C)
Midpoint
(
°
C)
Comments
Ninety
percentile
range
of
constant
laboratory
tests
with
1%
or
less
mortality
estimated.
21.05­
24.23
22.64
Based
on
the
5th
and
95th
percentiles
of
the
distribution
of
7­
day
LT50
results
at
acclimations
from
5
to
23
°
C.
Constant
laboratory
tests
regression
of
1%
or
less
mortality
estimate.
21.36­
24.54
22.95
Regression
of
all
of
the
lethality
data
for
acclimations
between
12
and
16
°
C.

Modeled
exposure
to
fluctuating
temperatures
based
on
constant
laboratory
tests.
20.13­
20.47
20.3
Assumes
a
steady
diel
fluctuation
of
4.13
°
C
per
day
for
a
7­
day
period.
LT50
converted
to
an
LT1
by
subtracting
2
°
C.
Fluctuating
laboratory
exposure
studies.
22.05­
24.05
23.05
Generally
based
on
high
diel
fluctuations
and
including
a
non­
indigenous
form
of
cutthroat
trout.
Field
Studies
with
Fluctuating
exposures
20.85­
23.5
22.18
Based
primarily
on
a
study
of
chinook
held
a
range
of
temperature
exposures
in
a
NW
river.
Best
estimate
of
threshold
21.09­
23.36
mid.
pt
22.23
The
absolute
range
is
a
7DADMax
of
20.13­
24.54
°
C,
with
a
mean
range
of
21.09­
23.36
°
C,
and
with
an
overall
midpoint
of
22.23
°
C.
The
lower
end
of
this
range
is
based
on
the
results
of
several
studies
using
different
species
in
a
controlled
laboratory
environment
at
moderate
acclimation
temperatures.
This
gives
it
greater
weight
than
if
only
one
species
was
tested
or
the
results
came
from
only
one
study
at
low
acclimation
temperatures.
Extra
caution
is
warranted
because
actual
exposure
periods
greater
than
7­
days
at
very
near
but
not
exceeding
the
estimated
lethal
threshold
can
exert
a
cumulative
stress
(
lowering
the
temperature
that
would
cause
lethality
over
the
longer
period
of
exposure).
For
these
reasons,
the
lower
half
of
the
potentially
protective
range
(
21.09­
22.23
°
C,
midpoint
of
21.66)
should
be
considered
to
be
a
more
confident
estimate
for
statewide
use.
Therefore,
to
prevent
direct
lethality
it
is
concluded
that
the
7DADMax
should
be
maintained
below
21.66
°
C,
or
single
daily
maximum
(
adding
0.95
for
conversion)
temperatures
maintained
below
22.61
°
C.

Near
Instantaneous
Lethality
Page
106
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
In
the
previous
discussion
the
period
of
exposure
was
selected
to
represent
a
reasonable
chronic
field
exposure
(
i.
e.,
a
7­
day
duration).
Higher
temperatures,
however,
will
exert
lethal
stress
in
much
shorter
periods
of
time,
such
that
at
some
temperature
lethality
will
occur
almost
instantaneously.
Since
wastewater
can
sometimes
be
discharged
at
very
high
temperatures
(
33­
40
°
C)
there
is
a
need
to
ensure
that
lethal
barriers
are
not
permitted
to
occur
in
Washington's
waters.

Data
is
available
on
exposure
periods
from
4
to
almost
4,000
seconds
have
been
used
to
evaluate
the
risk
of
instantaneous
high
water
temperatures.
The
purpose
of
this
analysis
is
to
determine
if
a
limit
should
be
placed
on
the
temperature
of
water
discharged
to
waters
of
the
state.
The
premise
of
such
a
restriction
would
be
that
at
a
given
high
temperature
fish
passing
through
the
plume
of
hot
water
will
be
immediately
killed.

Data
is
available
for
numerous
fish
species
with
the
tests
conducted
a
various
acclimation
temperatures
and
recording
several
important
mortality
endpoints
(
see
table
below).
These
endpoints
ranged
from
when
deaths
began
in
the
test
population
(
herein
described
as
the
LC1)
through
where
50
percent
of
the
population
had
died
(
LC50)
to
where
100
percent
mortality
(
LC100)
occurred.
Also
included
are
the
results
of
a
study
that
recorded
the
points
where
short­
term
exposures
caused
an
identifiable
increase
in
mortality
due
to
increased
predation.

The
complete
data
set
was
examined
by
separating
out
different
acclimation
ranges
and
different
endpoints,
by
only
looking
at
endpoints
with
significant
mortality
occurring
in
less
than
10
minutes,
and
by
lumping
all
the
data
together
to
examine
the
general
endpoint
of
lethality.
This
process
was
used
as
a
form
of
sensitivity
analysis
to
gain
better
confidence
and
understanding
of
the
strength
associated
with
extrapolating
beyond
the
lowest
tested
endpoint
duration
of
4
seconds.
It
is
assumed
here
that
extrapolation
to
a
1
second
exposure
duration
represents
instantaneous
exposure
and
thus
should
be
used
to
define
instantaneous
lethality.
The
power
curve
that
best
represented
the
data
was
used
to
create
the
extrapolation
equation
used
in
the
following
discussion.
The
following
is
intended
to
provide
a
general
understanding
of
the
analyses
that
were
conducted,
but
does
not
represent
an
exhaustive
description.

Separation
by
acclimation
temperature
and
by
test
endpoints
resulted
in
predicted
1
second
LC1
(
deaths
beginning)
to
LC100
(
all
dead)
values
ranging
from
of
32.76
°
C
to
34.32
°
C
at
moderate
acclimation
temperatures
of
15­
15.6
°
C.
The
1
second
endpoints
at
cold
acclimation
temperatures
(
9­
10
°
C)
ranged
from
32.45
°
C
to
34.34
°
C.
Since
sensitivity
to
temperature
is
well
demonstrated
to
be
dependent
upon
prior
acclimation
temperature,
this
analysis
is
believed
to
be
the
most
appropriate.
In
extrapolation
from
a
single
study
using
rainbow
trout
at
an
unknown
acclimation
temperature,
it
was
found
that
increased
predation
rates
may
occur
with
a
1
second
exposure
to
33.47
°
C.
Since
all
of
these
data
sets
were
small,
analyses
were
also
made
that
lumped
together
the
acclimation
temperatures.
This
resulted
in
an
estimated
1
second
LC1
(
deaths
beginning)
to
LC100
(
all
dead)
values
ranging
from
32.59
°
C
to
34.69
°
C.
In
examining
only
the
LC50
endpoints
that
occurred
within
4
minutes
or
less,
it
was
estimated
that
the
1
second
LC50
(
50%
dead)
would
occur
at
34.69
°
C.
And
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
107
Washington's
Surface
Water
quality
Standards
when
examining
all
of
the
available
short
term
mortality
data
together,
to
create
the
largest
data
set
for
extrapolation,
it
was
predicted
that
some
mortality
will
occur
with
a
1
second
exposure
of
33.18
°
C.
The
point
of
this
discussion
is
to
demonstrate
that
the
estimations
cluster
between
32.5
to
34.6
°
C,
a
very
narrow
range.
Taking
the
midpoint
of
this
range
to
further
improve
confidence
in
using
an
extrapolation
would
suggest
that
an
almost
instantaneous
exposure
to
temperatures
of
33.5
°
C
or
greater
will
likely
result
in
at
least
some
mortality
to
passing
fish.
Given
that
100%
mortality
has
been
demonstrated
at
32
°
C
in
11
seconds
at
low
acclimation
temperatures
(
9­
10
°
C)
and
50%
mortality
in
8
seconds,
and
that
mortality
has
been
noted
at
low
acclimation
temperatures
to
begin
in
4
seconds
the
estimate
should
not
be
considered
overly
conservative.
The
fact
that
greater
sensitivity
can
be
expected
at
even
lower
winter
and
early
spring
acclimation
temperatures
(
4­
8
°
C),
and
the
fact
that
lethal
exposures
are
cumulative
with
all
temperatures
above
the
lethal
threshold,
suggests
temperatures
greater
than
33
°
C
should
not
be
allowed
where
that
water
would
result
in
the
entrainment
of
fish
for
a
duration
of
1
second
or
longer.
For
this
reason,
discharges
with
temperatures
greater
than
33
°
C
should
be
evaluated
for
possible
risks
of
creating
a
lethal
barrier
to
organisms.
The
changing
velocity
and
temperature
of
the
discharge
plume
should
be
examined
to
ensure
that
time
spent
at
or
above
33
°
C
does
not
exceed
1
second
(
it
should
generally
be
assume
the
organism
is
entrained
and
is
moving
along
with
the
plume
in
the
analysis).

Figure
4.20.
The
following
is
an
example
of
one
of
the
data
subsets
examined:

Acclim.
Temp.
(
°
C)
Temperature
of
Test
(
°
C)
Time
(
sec.)
Endpoint
of
Test
Predicted
Endpoint
Temperatures
15.6
26.7
2,970
LC50
26.94052
15
28
1,350
LC50
27.505
15.6
29.4
26
LC50
30.51586
15.6
32.2
10
LC50
31.29244
Predicted
2
32.64542
Predicted
1
33.246
The
following
table
summarizes
the
data
used
for
the
analysis
of
lethality.
Median
Mortality
at
Warm
Acclimation
(
15­
15.6C)
­
LC50
Power
curve
2,970
1,350
26
10
y
=
33.246x­
0.0263
R2
=
0.8644
25
26
27
28
29
30
31
32
33
34
35
0
500
1,000
1,500
2,000
2,500
3,000
3,500
Time
(
seconds)
Temperature
(
C)
Page
108
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Table
4.21.
Summary
of
data
used
to
determine
risk
of
near
instantaneous
lethality.

Author
or
Study
Species
Acclimation
Temp.
(
°
C)
Temperature
of
Test
(
°
C)
Time
to
Effect
(
seconds)
Endpoint
Reported
Snyder
and
Blahm,
1971
Chum
15.6
26.7
2,640
LC1
15.6
26.7
2,970
LC50
15.6
26.7
3,960
LC100
15.6
29.4
15
LC1
15.6
29.4
26
LC50
15.6
29.4
60
LC100
15.6
32.2
6
LC1
15.6
32.2
10
LC50
15.6
32.2
15
LC100
10
26.7
100
LC1
10
26.7
240
LC50
10
26.7
2,220
LC100
10
32
4
LC1
10
32
8
LC50
10
32
11
LC100
Coutant,
1972(
a)
Chinook
15
28
1,350
LC50
Rainbow
17­
19
30.5
900
LC50
Lyytikainen,
Koskela,
and
Rissanen
(
1997)
Char
15
29
120
LC50
18
29
240
LC50
Baroudy
and
Elliott,
1994
Char
5
23.3
600
LC50
5
24.1
600
LC50
5
25.7
600
LC50
Groves
and
Mighell,
1970
Salmon
9
30
10
LC16
9
30
35
LC100
Coutant,
1972(
b)
Rainbow
30
33
Increased
Predation
28
120
Increased
Predation
26
1,920
Increased
Predation
c)
Temperature
Influenced
Fish
Diseases
Temperature
affects
both
the
occurrence
and
severity
of
many
diseases
and
infections
important
to
fish
and
other
aquatic
life.
While
some
diseases
are
associated
with
holding
fish
at
very
cold
temperatures,
and
some
are
prevalent
across
the
full
spectrum
of
temperatures
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
109
Washington's
Surface
Water
quality
Standards
occurring
in
Washington's
waters,
many
are
facilitated
by
temperatures
within
or
slightly
above
the
fully
protective
temperature
range
for
the
growth
of
our
native
fishes.
This
discussion
is
limited
to
those
diseases
that
are
known
to
cause
illness
in
populations
of
wild
fish
that
are
facilitated
by
temperatures
at
the
upper
end
of
the
physiologically
protective
range
of
our
indigenous
fish.
The
intent
is
to
identify
where
a
stream
temperature
standard
may
need
to
be
set
below
this
upper
end
to
prevent,
or
at
least
reduce
the
severity
of,
disease
outbreaks
in
natural
fish
populations.
It
must
be
recognized,
however,
that
temperature
standards
by
themselves
will
not
eliminate
the
risk
of
disease
in
fish.
Besides
warm
water
diseases,
native
fishes
are
harmed
by
numerous
very
important
cold
water
fish
diseases,
such
as
bacterial
kidney
disease.
It
is
of
course
reasonable
that
most
fish
pathogens
thrive
in
cold
waters
here
in
the
northwest
since
the
pathogens
themselves
have
evolved
along
with
our
native
fishes.

Fish
diseases
can
be
divided
into
four
types
of
infections:
1)
bacterial,
2)
viral,
3)
fungal,
and
4)
parasitic.
Regardless
of
the
type
of
infection,
temperature
affects
both
the
virulence
of
the
disease
as
well
as
the
immune
system
of
the
host
fish.
When
environmental
conditions
are
optimal
for
the
disease
it
grows
more
rapidly
and
is
often
more
virulent.
If
the
environmental
conditions
are
more
optimal
for
the
disease
then
they
are
for
the
fish,
then
there
is
a
greater
likelihood
that
the
disease
will
be
able
to
overcome
the
host's
defense
systems
and
create
serious
illness
(
Wedemeyer
and
Goodyear,
1984).
Just
as
each
disease
causing
organism
has
an
optimum
temperature
range,
they
also
have
lethal
boundaries
or
limits
of
activity.
If
the
temperature
is
above
or
below
these
thresholds,
the
disease
causing
organism
will
be
unable
to
reproduce
or
grow,
and
thus
will
be
unable
to
affect
disease
in
fish.
Catastrophic
outbreaks
of
many
bacterial
diseases
are
associated
with
water
temperatures
that
are
optimal
for
the
bacterium
but
above
the
optimal
temperature
for
the
infected
fish.

While
some
diseases
will
have
thresholds
of
temperature
above
or
below
which
the
disease
organism
is
unable
to
grow,
most
diseases
are
able
to
grow
at
some
temperature
that
occurs
during
the
year
in
Washington's
waters.
The
primary
goal,
therefore,
cannot
be
to
set
standards
at
which
diseases
organisms
will
not
exist.
A
complicating
factor
in
evaluating
diseases
is
that
most
have
multiple,
sometimes
hundreds,
of
strains.
Individual
strains
can
have
significantly
different
characteristics
for
optimal
infection
and
virulence.
Since
researchers
often
do
not
specify
specific
strains
examined,
this
creates
variability
in
research
results
that
may
very
well
just
be
the
consequence
of
testing
different
strains.
Further,
native
fish
differ
in
susceptibility
to
these
diseases
both
by
subspecies
and
by
individual
stock
(
Li
et
al.,
1987,
and
others).

The
focus
of
this
effort
is
to
identify
pathogens
that:
1)
occur
in
freshwaters;
2)
are
enhanced
by
increasing
water
temperatures
in
the
upper
optimal
range
of
our
native
fishes
(
roughly
14­
18
°
C);
and
3)
have
been
associated
with
serious
outbreaks
of
disease
in
indigenous
wild
fish
populations.

In
reviewing
the
literature,
only
three
diseases
appear
to
meet
all
three
criteria,
these
are
two
parasites
(
Ichthyophthirius
multifillis
and
Ceratomyxa
shasta)
and
columnaris
disease.
While
numerous
other
diseases
are
documented
to
be
influenced
by
warming
temperatures,
these
others
tend
to
be
primarily
problems
associated
with
intensive
fish
culturing
facilities,
Page
110
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
or
are
associated
with
species
and
temperatures
not
common
in
our
natural
waters.
Since
there
is
little
suggestion
that
any
of
these
other
diseases
pose
a
significant
threat
to
the
welfare
of
our
native
fishes,
they
will
be
used
only
broadly
to
discuss
how
temperatures
influence
the
health
of
fish
and
in
support
of
the
findings
for
the
two
specific
diseases
discussed
below.

Ichthyophthiriasis.
Post
(
1987)
notes
that
the
etiological
agent
for
ichthyophthiriasis
is
the
largest
protozoan
found
on
fishes.
Occurring
on
both
hatchery
and
wild
fish
(
Bell,
1986),
ichthyophthiriasis
is
considered
one
of
the
most
prevalent
diseases
of
fishes
(
Post,
1987).
Bell
(
1986)
notes
that
outbreaks
in
fingerlings
often
occur
at
temperatures
above
15.5
°
C,
and
the
optimum
temperature
for
the
organisms
is
25­
27
°
C.
Post
(
1987)
notes
that
temperatures
over
12­
15
°
C
are
more
suitable
for
reproduction,
and
that
disastrous
losses
have
occurred
in
trout
culture
where
water
warms
to
near
20
°
C.

Ceratomyxiasis
Shasta.
An
obligate
parasite
widespread
in
the
Northwestern
U.
S.
where
anadromous
salmonids
return
from
the
ocean
to
spawn.
While
not
important
to
adult
fish,
C.
shasta
may
be
devastating
to
very
young
salmonids
(
Post,
1987)
and
is
a
parasite
of
concern
for
northwest
salmonids
(
Bartholomew,
Rohevec,
and
Fryer,
1989;
and
Conrad
and
Decew
1961;
as
cited
by
Hoffman
and
Bauer,
1971).
Bartholomew,
Rohevec,
and
Fryer
(
1989)
note
it
is
important
because
it
not
only
causes
losses
in
hatchery­
reared
and
wild
juvenile
salmonids
but
also
contributes
to
prespawning
mortality
in
adult
salmon.

Infections
of
Ceratomyxa
shasta
are
temperature
dependent
(
Yamamoto
and
Sanders
1979,
and
Udey
et
al.
1975;
as
cited
in
Bartholomew,
Rohevec,
and
Fryer,
1989).
Udey
et
al.
(
1975)
found
that
rainbow
trout
exposed
to
the
infective
stage
of
C.
shasta
and
held
at
water
temperatures
of
6.7
to
23.3
°
C
had
little
or
no
ability
to
overcome
the
infection,
and
that
the
mean
time
from
exposure
to
death
was
directly
correlated
to
temperature
(
about
155
days
at
6.7
°
C
and
14
days
at
23.3
°
C).
In
rainbow
trout
the
disease
process
was
suppressed
at
3.9
°
C;
however,
when
the
infected
fish
were
subsequently
transferred
to
water
at
17.8
°
C,
many
died.
Mortality
in
exposed
rainbow
trout
was
greater
than
92%
at
temperatures
above
12.2
°
C,
84­
75%
at
9.4­
6.7
°
C,
and
zero
at
3.9
°
C
(
Fryer
and
Pilcher,
1974).
In
juvenile
coho
salmon
mortality
was
92­
100%
at
20.6
°
C
and
above,
57­
59%
at
17.8
°
C,
13­
31%
at
15­
12.2
°
C,
and
4­
0%
at
9.4
°
C
and
lower.

Evidence
suggests
that
only
salmonids
are
susceptible
to
C.
shasta
infections,
but
that
susceptibility
may
vary
within
species.
Juvenile
salmonids
originating
from
waters
containing
the
infective
stage
of
the
parasite
have
been
found
to
be
more
resistant
than
strains
from
areas
free
of
the
infective
stage
(
Johnson
1975,
Zinn
et
al.
1977,
Buchanan
et
al.
1983,
and
Hoffmaster
1985;
as
cited
in
Bartholomew,
Rohevec,
and
Fryer,
1989).
Sanders
et
al.,
(
1970;
as
cited
in
Li
et
al.,
1987)
reported
that
infected
coho
show
an
increasing
susceptibility
as
temperature
rises:
mortality
below
10
°
C
is
2%
at
most,
22%
at
15
°
C,
and
84%
at
20.5
°
C.
In
contrast,
mortality
of
infected
juvenile
steelhead
trout
can
be
as
high
as
80%
and
is
independent
of
temperature.
Ratliff
(
1983)
studied
the
infective
stage
of
Ceratomyxa
shasta
in
the
Deschutes
River
of
central
Oregon
and
found
that
C.
shasta
emanates
from
the
bottoms
reservoirs,
and
that
the
infective
period
began
in
the
spring
when
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
111
Washington's
Surface
Water
quality
Standards
river
temperatures
were
between
6.9
and
8.6
°
C.
The
author
noted
that
based
on
his
observations
C.
shasta
spores
are
more
likely
observed
when
test
fish
are
held
during
the
incubation
period
at
10
°
C
rather
than
at
warmer
temperatures.
Waters
where
infected
fish
have
been
found
do
not
necessarily
contain
the
infective
stage
of
the
parasite
(
Johnson
et
al.
1979;
as
cited
in
Bartholomew,
Rohevec,
and
Fryer,
1989),
and
even
in
the
Columbia
River
Basin
where
infected
fish
migrate
and
distribute
spores
throughout
the
drainage,
the
infective
stage
has
not
been
demonstrated
in
many
tributaries.
The
authors
(
Bartholomew,
Rohevec,
and
Fryer,
1989)
suggest
that
the
geographic
isolation
of
the
disease
supports
the
position
that
presence
of
the
spores
is
insufficient
to
cause
transmission
and
disease,
and
that
a
yet
unidentified
factor
(
perhaps
an
intermediate
host)
is
required
for
the
completion
of
the
life
cycle
of
this
parasite.

Columnaris
Disease.
There
is
little
doubt
that
columnaris
disease
is
the
most
important
warm­
water
disease
for
our
native
salmonid
populations.
Frequent
and
catastrophic
losses
to
natural
populations
throughout
the
Pacific
northwest
are
well
document
throughout
the
literature.
For
this
reason,
the
prevention
of
human­
caused
additional
losses
from
columnaris
disease
should
be
considered
a
critical
element
in
setting
temperature
standards
for
the
state
of
Washington.

In
evaluating
the
research
it
is
important
to
recognize
that
there
are
probably
at
least
1,200
strains
of
columnaris
in
Washington,
and
that
strains
can
be
categorized
as
possessing
low,
medium,
and
high
levels
of
virulence
(
Pacha,
1961).
High
virulence
strains
are
infective
and
capable
of
producing
high
rates
of
mortality
at
low
temperatures,
while
low
virulence
strains
are
problematic
only
at
higher
temperatures
(
Bell,
1986).
Since
only
a
handful
of
authors
categorized
the
strains
they
were
evaluating,
care
must
be
exercised
in
broadly
applying
the
conclusions
of
any
one
study.
Table
4.22
below
summarizes
the
results
noted
in
the
literature.

The
milestones
of
12­
13,
15­
16,
18­
20
°
C
are
cited
extensively
in
both
field
and
laboratory
research,
and
show
generally
consistent
levels
of
increasing
risk
of
columnaris
disease.
In
field
studies
it
has
been
noted
that
as
river
temperatures
rise
to
about
10­
12.8
°
C,
researchers
begin
isolating
columnaris
strains
from
water
(
Fujihara
and
Nakatani,
1970).
Temperatures
above
13
°
C
were
associated
with
the
occurrence
columnaris
in
thirteen
species
of
fish
collected
from
the
Columbia
River
watershed
(
Fujihara
and
Huntgate,
1970).
In
the
Fraser
River
in
Canada
it
was
found
that
prespawning
mortality
of
sockeye
was
eliminated
by
maintaining
average
temperatures
on
the
spawning
grounds
of
12.8
°
C
(
Colgrove
and
Wood,
1966).
This
finding
was
generally
supported
by
the
work
of
Johnson
and
Brice
(
1952,
as
cited
in
Colgrove
and
Wood,
1966)
that
exposed
four
species
of
salmonids
to
columnaris
for
six
months
and
found
that
no
mortalities
developed
when
daily
maximum
temperatures
were
12.8
°
C
or
less.
At
maximum
temperatures
of
15.6­
18.3
°
C,
mortalities
increased
to
0.7­
15.5%
in
three
species
but
remained
zero
in
the
fourth,
and
at
maximum
temperatures
18.3­
21.1
°
C
all
four
species
showed
high
mortalities
(
37.5­
82%).
These
results
are
supported
by
other
researchers
studying
columnaris
under
natural
environmental
conditions.
Page
112
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
The
spread
of
columnaris
throughout
the
Columbia
River
basin
has
been
noted
to
be
linked
to
the
water
temperatures
occurring
during
individual
years.
In
warmer
years,
columnaris
disease
is
widespread
throughout
the
entire
basin,
but
in
cooler
years,
the
first
major
exposure
occurred
at
McNary
Dam
and
the
warmer
tributaries
(
Pacha
and
Ordal,
1970).
A
difference
of
heating
or
cooling
natural
river
water
2.2
°
C
from
its
natural
(
17.7­
21.7
°
C)
condition
was
shown
to
increase
and
decrease
mortality
rates
in
naturally
exposed
fish
(
Fujihara,
Olson,
and
Nakatani,
1971).
While
the
decrease
resulted
in
only
a
modest
decrease
(
4.2%)
in
mortality
(
6.2
versus
10.4%),
the
increase
resulted
in
a
more
substantial
increase
(
19.5%)
in
mortality
(
29.9
versus
10.4%).

At
river
temperatures
of
about
15
°
C
isolation
of
columnaris
cultures
is
typically
quite
successful
(
Ordal
and
Pacha,
1963;
Pacha
and
Ordal,
1970),
with
15
°
C
found
also
to
be
a
demarcation
point
between
high
(
54%)
and
moderate
(
22%)
levels
of
infection
in
the
crowded
spawning
channels
of
the
Columbia
River.
Scrap
fish
collected
from
waters
throughout
the
Columbia
River
basin
were
found
to
exhibit
the
disease
when
warmed
to
16.7
°
C
(
Pacha
and
Ordal,
1970).
At
river
temperatures
above
18.8
°
C
strains
of
low
and
intermediate
virulence
can
be
readily
isolated
from
fish
(
Pacha,
1961;
Bell,
1986).
Major
outbreaks
are
said
to
almost
always
occur
during
periods
in
which
the
water
temperature
was
18.3­
21.1
°
C
(
Pacha,
1961),
and
an
average
mid­
summer
river
temperature
of
20.3
°
C
was
associated
with
a
catastrophic
outbreak
in
sockeye
salmon
in
the
Columbia
River
(
Fish,
1948).

Laboratory
tests
generally
confirm
what
has
been
found
through
these
field
or
controlled
channel
studies.
A
complication
is
created
by
looking
strictly
at
mortality
rates
from
these
laboratory
tests.
Fish
are
generally
injected
with
or
exposed
to
high
doses
of
the
pathogen,
and
some
are
inoculated
with
high
virulent
strains
while
others
with
low
or
intermediate
virulent
strains.
Putting
aside
these
general
problems
with
comparing
laboratory
test
results
we
can
see
the
same
basic
patterns
emerge
as
were
found
in
field
research.
Constant
temperatures
of
less
than
12
°
C
result
in
low
to
no
infections
or
mortalities
(
Fryer
and
Pilcher,
1974;
Post,
1987;
Fujihara
and
Nakatani,
1970;
Ordal
and
Pacha,
1963).
Constant
temperatures
of
15­
18
°
C
have
produced
a
wide
range
of
mortalities,
but
is
most
characterized
by
moderate
(
typically
20­
60%)
to
heavy
(
80­
100%)
mortality.
In
this
range,
authors
that
have
made
the
distinction
have
shown
high
virulence
strains
to
be
commonly
associated
with
the
highest
mortality
(
see
table
4.22
for
references).
At
constant
temperature
exposures
of
20­
23.6
°
C,
mortalities
of
infected
fish
are
consistently
very
high
(
70­
100%)
(
Ordal
and
Rucker,
1944;
Fryer
and
Picher,
1974;
Fish
and
Rucker,
1943;
as
cited
in
Ordal
and
Pacha,
1963;
Holt
et.
al.,
1975).
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
113
Washington's
Surface
Water
quality
Standards
Table
4.22.
Summary
of
literature
findings
for
columnaris
disease.

Approximate
Temperature
Range
or
Direction
Categories
of
Effects
of
Interest
noted
in
the
literature
associated
with
the
temperature
range
Citation
from
the
Literature
<
12
Seldom
infectious;
difficult
to
isolate
in
the
field;
and
low
(
0­
8%)
fatalities
experimentally
infected
fish.
Fryer
and
Pilcher,
1974;
Post,
1987;
Fujihara
and
Nakatani,
1970;
Ordal
and
Pacha,
1963
13
Organism
begins
to
be
isolated
in
the
water
and
fish;
not
associated
with
measurable
prespawning
mortality;
moderate
(
0­
20%)
to
high
(
20­
100%)
mortality
in
experimentally
infected
fish.
Holt
et
al.,
1975;
Fryer
and
Pilcher,
1974;
Fujihara
and
Nakatani,
1970;
Johnson
and
Brice,
1952
as
cited
in
Pacha,
and
Colgrove
and
Wood,
1966;
Fish,
1944;
USEPA,
1976;
Fujihara
and
Huntgate,
1970;
Colgrove
and
Wood,
1966
14.4
Average
river
temperature
not
leading
to
disease.
Colgrove
and
Wood,
1966
<
15­
15.6
Rarely
a
problem;
mortality
from
low
virulence
strains
decline;
incidence
of
disease
in
river
declines
to
22%.
Amend,
1970;
as
cited
in
Austin
and
Austin;
Johnson
and
Brice,
1952
and
Rucker,
1944
as
cited
in
Pacha,
1961;
Garnjobst,
1945,
as
cited
in
Colgrove
and
Wood,
1966;
Fujihara
and
Olson,
1962,
as
cited
in
Colgrove
and
Wood,
1966
15­
16.7
Moderate
(
31­
56%)
mortalities
become
more
consistent.
in
infected
test
fish;
disease
appears
in
Scrap­
fish
after
temperature
elevation;
epizootics
in
aquarium.
Holt
et
al.,
1975;
USEPA,
1976;
Fryer
and
Pilcher,
1974;
Pacha
and
Ordal;
Colgrove
and
Wood,
1966,
as
cited
in
Pacha
and
Ordal,
1970;
Post,
1987
15.6­
16
Moderate
(
60%)
mortality
of
injured
fish;
high
mortality
(
80­
100%)
from
high
virulence
strains;
prespawning
mortality
63­
81%,
two
week
average
temperature
necessary
to
initiate
pathological
effects
in
river;
moderately
high
(
30­
64%)
mortality
in
infected
test
fish.
Post,
1987;
Pacha
and
Ordal,
1970;
Colgrove
and
Wood,
1966;
Fish,
1944;
Ordal
and
Pacha,
1963;
Ordal
and
Rucker,
1944,
as
cited
in
Pacha
and
Ordal,
1970
>
15­
15.6
Easy
to
isolate
in
the
field;
associated
with
seasonal
mortality
in
hatcheries;
mortality
and
morbidity
become
factors
in
natural
waters;
outbreaks
of
high
virulence
strains;
initiated
mortalities
in
migrating
sockeye;
disease
incidence
in
river
becomes
high
(
54%).
Ordal
and
Pacha,
1963;
Fujihara
and
Nakatani,
1970;
Post,
1987;
Bell,
1986;
Colgrove
and
Wood,
as
cited
by
Gilhousen,
1970;
Fujihara
and
Olson,
1962,
as
cited
in
Colgrove
and
Wood,
1966
13­
18
Low
(
0.6­
7.7%),
moderate
(
20­
50%),
and
high
(
60­
100%)
mortality
in
exposed
test
fish.
Known
virulent
strains
resulting
in
highest
mortality.
Fryer
and
Pilcher,
1974;
Johnson
and
Brice,
1952,
as
cited
in
Pacha,
1961;
Pacha
and
Ordal,
1970
15.6­
18.3
Low
(
0.7­
15.5%)
mortality
in
three
species
of
Johnson
and
Brice,
1952,
as
cited
Page
114
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Water
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Standards
salmonids
in
Colgrove
and
Wood,
1966
15.9­
19.9
Fluctuating
river
temp
with
low
(
6.2%)
mortality.
Fujihara,
Olson,
and
Nakatani,
1971
17­
18
Mortality
sometimes
moderate
(
37­
50%)
but
mostly
very
high
(
99­
100%)
in
infected
test
fish;
explosive
infections;
high
mortality
from
all
strain
types.
Fryer
and
Pilcher,
1974;
Holt
et
al.,
1975;
USEPA,
1976;
Pacha
and
Ordal,
1970
17­
21.7
Fluctuating
river
temp
with
10.4%
mortality.
Fujihara,
Olson
and
Nakatani,
1971
<
18
Deaths
greatly
diminish.
Ordal
and
Pacha,
1963
>
18
Mortality
high
from
all
strains,
low
and
moderate
strains
isolated.
Johnson
and
Brice,
1952,
and
Rucker,
1944,
as
cited
in
Pacha,
1961
18­
22
Mortalities
moderate
to
high
(
37.5­
82%)
in
infected
fish;
most
outbreaks;
fish
found
releasing
columnaris;
Epizootics.
Johnson
and
Brice,
1952,
as
cited
in
Colgrove
and
Wood,
1966,
and
as
cited
in
Pacha,
1961;
Fujihara
and
Nakatani,
1970;
Amend,
1970,
as
cited
in
Austin
and
Austin,
1987;
and
Davis,
1922,
Nigrelli
and
Hunter,
1945,
Isom,
1960,
and
Johnson
and
Brice,
1952,
as
cited
in
Pacha,
1961
19.9­
23.9
Fluctuating
river
temp
with
29%
mortality.
Fujihara,
Olson
and
Nakatani,
1971
20­
21
Outbreaks
of
low
virulence
strains;
mortality
high
(
70­
100%)
in
infected
test
fish,
catastrophic
outbreak,
28­
75%
morbidity
in
river
population;
outbreaks
in
physically
stressed
fish;
serious
epidemics.
Bell,
1986;
Ordal
and
Pacha,
1963;
Pacha
and
Ordal,
1970;
Fish,
1948;
Holt
et.
al.,
1948;
Fryer
and
Pilcher,
1974;
Post,
1987;
Fish,
1944
22.2­
23.6
Mortality
100%
in
infected
test
fish;
summer
maximum
associated
with
devastating
outbreak.
Ordal
and
Rucker,
1944,
as
cited
in
Pacha
and
Ordal,
1970;
Fish,
1948;
Fryer
and
Pilcher,
1974
25­
37
Optimum
growth
temperature
for
organism.
Garnjobst,
1945,
as
cited
in
Colgrove
and
Wood,
1966;
Pacha,
1961;
Post,
1987
Based
on
a
review
of
the
available
literature,
it
is
concluded
that
for
columnaris,
Ceratomyxiasis
Shasta,
and
Ichthyophthiriasis,
as
well
as
for
warm­
water
induced
diseases
in
general,
the
following
general
statements
hold
true:

1.
Average
temperatures
below
12­
13
°
C
significantly
and
often
completely
eliminate
both
infection
and
mortality;
2.
Average
temperatures
above
15­
16
°
C
are
associated
with
often
serious
rates
of
infection
and
noticeable
mortality;
and
3.
Average
temperatures
above
18­
20
°
C
are
commonly
associated
with
very
severe
infections
and
often
catastrophic
outbreaks
of
many
fish
diseases.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
115
Washington's
Surface
Water
quality
Standards
Treating
these
ranges
as
either
daily
or
monthly
average
temperature
may
best
bracket
the
exposure
periods
associated
with
the
spread
of
disease
causing
organisms.
Adding
0.58
°
C
and
3.18
°
C,
respectively,
converts
a
maximum
summer
daily
and
monthly
average
temperatures
to
estimated
7DADMax
temperature
metrics.
This
approach
yields
the
following
estimates
(
Table
4­
24):

Table
4.24.
Warm
water
disease
risk
to
salmon
and
trout.

Disease
Incidence
Level
7DADMax
(
°
C)
Midpoint
(
°
C)
Virtual
elimination
of
warmwater
disease
effects
12.58­
16.18
14.38
Avoiding
serious
rates
of
infection
and
mortality
15.58­
19.18
17.38
Severe
infections
and
catastrophic
outbreaks
18.58­
23.18
20.88
To
reduce
the
risks
of
serious
infection
and
mortality
from
warm
water
mediated
bacterial
and
parasitic
diseases,
it
is
concluded
that
the
7­
day
average
of
the
daily
maximum
temperatures
(
7DADMax)
should
not
exceed
15.58­
19.18
°
C
(
17.38).
This
approach
will
provide
safe
harbor
for
resident
species
and
critical
life
stages
as
well
as
serving
to
reduce
or
eliminate
disease­
caused
prespawning
mortalities
in
migrating
fish.
When
average
river
temperatures
exceed
20
°
C,
or
7DADMax
temperatures
exceed
20.88
°
C,
explosive
infection
rates
and
the
risk
of
catastrophic
population­
level
outbreaks
in
natural
populations
become
a
serious
concern.

The
research
evaluated
in
this
paper
has
focused
on
protecting
cold­
water
fish
communities.
Therefore,
the
above
conclusions
may
not
be
applicable,
and
are
therefore
not
recommended
for
application,
to
any
designated
habitat
for
warm
water
fish­
species.
Information
on
the
disease
threats
to
native
warm
water
fishes
was
not
discovered
during
the
review
of
the
available
literature.
For
warm
water
fish
habitat
it
is
recommended
that
temperatures
be
maintained
within
the
range
of
what
otherwise
is
expected
to
produce
healthy
warm
water
fish
communities.
By
maintain
the
general
health
of
these
species
it
will
also
increase
their
resistance
to
potential
diseases.

d)
Smoltification
and
Sea
Water
Adaptation
Smoltification
is
the
name
given
to
the
physiological,
morphological,
and
changes
that
occur
in
anadromous
fish
as
they
prepare
to
leave
fresh
waters
for
life
in
saline
marine
waters
(
Clarke
and
Hirano,
1995;
Wedemeyer,
Saunders,
and
Clarke,
1980;
Sauter
and
Maule,
1997,
1999).
The
greatest
concerns
at
this
stage
of
life
would
be
that:
1)
parr
fail
to
migrate
because
of
temperatures
that
are
too
warm
(
Clarke,
Shlebourn,
and
Brett,
1981),
2)
that
they
will
migrate
but
not
be
fully
capable
of
living
in
marine
waters
(
Marine,
1997),
or
3)
that
delays
in
smoltification
could
require
that
juvenile
fish
remain
in
the
estuarine
environment
longer
and
experience
enhanced
predation.

Pennel
and
Barton
(
1996)
provided
an
excellent
summary
of
the
process
of
smoltification:
In
marine
and
estuarine
waters
the
salinity
in
the
water
surrounding
the
fish
is
greater
than
the
Page
116
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
salinity
of
the
internal
body
fluids.
The
gradient
produced
tends
to
draw
out
the
water
from
the
body
of
the
fish
while
salts
from
the
surrounding
water
diffuse
inward.
The
lost
water
can
be
replaced
by
drinking
the
saline
marine
water
but
salt
must
be
removed
from
the
body
to
prevent
detrimental
accumulation.
The
process
of
removing
sodium
ions
from
the
blood
takes
place
in
through
special
cells
in
the
gills.
Sodium
ions
are
transported
from
the
area
of
high
concentration
of
the
cells
to
the
external
environment
down
a
concentration
gradient
via
sodium
channels.
Energy
to
drive
the
sodium
pump
comes
from
the
conversion
of
adenosine
triphosphate
(
ATP)
to
adenosine
diphosphate
(
ADP),
a
reaction
catalyzed
by
the
enzyme
sodium/
potassium
adenosine
triphosphatase
(
Na+/
K+­
ATPase).
In
order
to
maintain
neutrality,
Cl­
diffuses
directly
from
the
chloride
cell
into
the
apical
space
and
external
environment,
with
the
activity
of
the
chloride
cell
probably
under
hormonal
control
(
Pennel
and
Barton,
1996).
A
major
change
that
occurs
during
smoltification
is
a
dramatic
increase
in
the
number
and
activity
of
chloride
cells
in
the
gills.
An
increase
in
Na+/
K+­
ATPase
activity
indicates
chloride
cell
activity
and
monitoring
this
activity
is
one
of
the
useful
smolt
indicators.
changes
such
as
saltwater
preference
and
migratory
behavior
(
Folmar
and
Dichoff
1980;
as
cited
in
Pennel
and
Barton,
1996)
often
become
evident
as
the
juvenile
fish
become
smolts.
A
performance
monitoring
method
to
evaluate
functional
smolts
is
the
saltwater
challenge
test.
The
saltwater
challenge
test
is
used
to
determine
if
smolts
are
capable
of
regulating
their
blood
Na+.
Fish
are
placed
into
a
known
concentration
of
salt
water
and
blood
samples
are
taken
from
fish
24
hours
later
for
Na+
analysis.
If
the
fish
are
capable
of
regulating
blood
Na+
levels
to
<
170
meq/
l,
they
are
considered
as
functional
smolts
(
Pennel
and
Barton,
1996).

This
ability
to
regulate
and
maintain
the
osmotic
gradient
across
a
fish's
body
surface,
whether
the
fish
is
exposed
to
the
hydrating
conditions
of
fresh
water
or
the
dehydrating
conditions
of
sea
water,
is
referred
to
as
osmoregulation.
While
salmonid
stocks
that
move
to
the
estuarine
environments
early
as
young
fry
or
juveniles
(
chum,
pink,
some
populations
of
coho,
and
to
some
extent
ocean­
type
chinook)
tend
to
develop
a
more
rapid
tolerance
to
saline
waters,
those
that
have
an
extended
period
of
freshwater
residence
as
juveniles
(
coho,
sockeye,
and
stream­
type
chinook)
possess
a
limited
capacity
for
ionic
regulation
in
hyperosmotic
media
prior
to
reaching
the
smolt
stage
(
Clarke
and
Hirano,
1995).

While
the
evidence
suggests
that
smoltification
is
triggered
by
endogenous
rhythms
(
Hoar,
1988;
Wagner,
1974);
growth
(
Ewing
et
al.,
1979),
photoperiod
(
Folmar
et
al.,
1982
Wagner,
1974),
and
temperature
all
critically
influence
the
timing,
extent,
and
success
of
the
transition
process.
Temperature
influences
smoltification
in
at
least
two
primary
ways.
The
first
is
that
juvenile
fish
must
generally
reach
a
critical
size
to
be
capable
of
successfully
smolting
(
Clarke
and
Shelbourn,
1985;
Mahnken
and
Waknitz,
1979).
The
second
is
that
temperature
affects
critical
enzyme
activities
that
control
the
ability
to
excrete
salts
from
the
blood
 
and
thus
to
be
able
to
live
for
an
extended
period
in
saline
waters
(
Zaugg
and
Wagner,
1973;
Wagner,
1974;
Wedemeyer,
Saunders,
and
Clarke,
1980;
Duston,
Saunders,
and
Knox,
1991).

Laboratory
and
field
work
associating
temperature
with
the
smoltification
of
salmonids
has
to
date
focused
on
steelhead
trout,
chinook
salmon
and
coho
salmon,
all
species
that
have
a
Evaluating
Standards
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Protecting
Aquatic
Life
in
Page
117
Washington's
Surface
Water
quality
Standards
considerable
period
of
fresh
water
rearing
prior
to
moving
to
marine
waters.
The
following
summarizes
the
studies
reviewed
on
these
species:

Steelhead
Trout
In
Washington,
migratory
smolts
usually
move
to
sea
during
April
through
June,
with
a
peak
about
mid
April
(
Wydoski
and
Whitney,
1979).
Zaugg
et
al.
(
1972),
Zaugg
and
Wagner
(
1973),
and
Adams,
Zaugg
and
McLain
(
1973)
suggest
that
temperatures
higher
than
about
12­
13
°
C
may
alter
the
juvenile
migratory
behavior
and
physiological
condition
of
steelhead
trout.
In
their
work
these
authors
found
that
fish
held
at
15
°
C
and
20
°
C
experience
high
rates
of
mortality
with
subsequent
transfer
to
sea
water
(
30ppt).
The
authors
also
noted
that
smolts
held
at
10
°
C
were
unable
to
maintain
high
NaK­
ATPase
activity
throughout
the
smolt
season
since
at
the
beginning
of
the
season
the
increase
in
activity
for
this
group
was
similar
to
that
in
the
6.5
°
C
group.
Steelhead
maintained
at
6.5
or
10
º
C,
however,
experienced
an
elevation
of
Na+,
K+­
stimulated
ATPase
activity
and
assumed
the
characteristic
slender,
silvery
appearance
of
smolts.
Adams,
Zaugg,
and
McLain
(
1975)
evaluated
winter
and
summer
steelhead
trout
metamorphosis
at
six
different
growth
temperatures
ranging
from
6
to
15
º
C.
Salt
water
survival
in
35%
sea
water
at
10
º
C
was
used
to
determine
the
extent
of
transformation.
The
highest
temperature
where
a
transformation
was
indicated
was
11.3
º
C.
By
April
fish
reared
at
6
º
C
had
elevated
ATPase
levels
typical
of
smolts
or
migratory
animals
and
showed
92%
survival
in
sea
water.
Ten
and
11.3
º
C­
reared
fish
showed
a
shortlived
elevation
in
ATPase
in
mid­
April
alone
concurrently
with
100%
sea
water
survival
at
that
time.
Only
in
6
º
C­
reared
animals
did
the
salt
water
survival
ability
continue
into
May.
Zaugg
(
1981)
found
exposures
of
fish
to
13
°
C
resulted
in
a
delay
in
migration,
fewer
total
fish
attempting
migration,
and
lower
gill
Na+­
K+
ATPase
activity.
Wagner
(
1974)
found
that
parr­
smolt
transformation
in
steelhead
appears
to
be
based
on
endogenous
rhythm.
Wagner
also
found
a
tendency
for
fish
reared
at
a
normal
temperature
cycle
(
6.9­
18.6
°
C)
to
migrate
in
greater
numbers
than
those
that
experienced
a
constant
temperature
regime
(
12.3
°
C).
This
was
consistent
with
Wagner
(
1971)
where
fish
reared
under
constant
photoperiods
and
normal
temperatures
also
migrated
in
greater
numbers
than
fish
receiving
a
similar
photoperiod
but
a
constant
temperature.
Zaugg
(
2001)
summarized
a
study
done
in
1977
assessing
the
effect
of
barging
and
trucking
on
the
state
of
smoltification
in
Dworshak
steelhead.
Smolts
held
in
water
below
10
°
C
had
no
trouble
in
maintaining
elevated
ATPase
activities,
whether
of
hatchery
or
wild
origin.
However,
barged
two­
year
old
steelhead
smolts
held
in
the
Columbia
River
water
(
12.5­
13.6
°
C)
reverted
to
parr­
like
gill
ATPase
activities
within
approximately
2
weeks
(
from
May
6
to
May
21).
Smolts
held
in
water
below
10
°
C
retained
a
smolt­
level
gill
ATPase
activity
at
least
through
June
8
when
the
experiment
was
terminated.
Zaugg
(
2001)
summarized
the
work
reported
by
Dr.
H.
W.
Lorz
in
December,
1974,
to
the
Annual
Northwest
Fish
Culture
Conference.
Three
groups
of
winter
steelhead
were
reared
at
the
laboratory
under
normal
photoperiod
but
different
constant
temperatures
(
8,
12,
and
16
°
C).
Only
the
fish
reared
at
16
°
C
did
not
show
an
increase
in
ATPase
activity
during
April.
Samples
of
the
fish
were
released
into
a
small
coastal
stream.
The
fish
reared
at
16
°
C
did
not
migrate
at
the
same
rate
as
those
fish
reared
at
lower
temperatures,
with
a
portion
of
these
fish
showing
a
three
week
delay.
However,
measurements
from
downstream
migrants
showed
that
the
ATPase
activity
had
increased
Page
118
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
during
this
three
to
four
week
period.
This
study
supports
the
position
that
there
is
a
close
relationship
between
elevated
gill
Na+
K+
stimulated
ATPase
activity
and
seaward
migratory
movement;
as
well
as
providing
information
on
the
deleterious
effect
of
elevated
temperature
on
parr­
smolt
transformation.
In
a
second
part
of
this
study,
two
groups
of
steelhead
were
maintained
in
laboratory
tanks,
one
with
water
at
8
°
C
and
the
other
at
16
°
C
into
May.
At
the
time
when
elevated
enzyme
activity
was
observed
in
the
fish
held
at
8
°
C,
enzyme
activity
remained
low
and
unchanged
in
fish
held
at
16
°
C.
Fish
held
at
16
°
C
were
then
subjected
to
six
different
temperature
regimes
(
0/
24,
4/
18,
6/
16,
10/
12,
14/
18,
24/
0)
(
time
at
8
°
C
/
time
at
16
°
C).
The
fish
maintained
at
24hr/
day
at
16
°
C
failed
to
develop
elevated
ATPase
activity
as
did
also
the
fish
held
in
4/
18,
6/
16,
10/
12,
14/
18.
Only
the
fish
held
for
24
hours
per
day
at
8
°
C
exhibited
an
increase
in
ATPase
activity,
which
occurred
in
the
5th
week.

Chinook
salmon
Once
juvenile
chinook
grow
to
an
appropriate
size
they
will
migrate
to
the
ocean
(
Ewing
et
al.,
1979;
Clarke
and
Shelbourn,
1985).
Out­
going
migrations
typically
occur
during
the
first
and
second
years
of
life
(
Wydoski
and
Whitney,
1979)
but
since
success
is
related
to
smolt
size,
some
fish
will
not
be
ready
until
their
third
year.
Clarke
and
Shelbourn
(
1985)
found
that
optimum
regulation
of
plasma
sodium
concentrations
in
ocean­
type
fall
chinook
salmon
occurred
with
transfer
from
13.8
º
C
fresh
water
to
10.2
º
C
sea
water.
They
also
noted
that
severe
descaling
in
their
freshwater
holding
tanks
occurred
in
groups
of
smolts
reared
at
16
or
17
°
C,
as
well
as
with
groups
transferred
from
8­
12
°
C
freshwater
to
14
°
C
seawater.
Marine
(
1997)
conducted
laboratory
tests
on
fall
run
chinook
salmon
from
Sacramento
River
hatchery
stocks
to
determine
the
chronic
effects
of
varying
temperatures
from
13
to
24
°
C
on
growth
and
smoltification
patterns
(
Na+­
K+
ATPase
activity
and
seawater
challenges).
The
control
test
was
15
°
C
(
fluctuated
from
13­
16
°
C),
the
intermediate
was
18.5
°
C
(
17­
20
°
C),
and
the
extreme
was
21.5
°
C
(
21­
24
°
C).
The
author
concluded
that
both
acceleration
and
inhibition
of
chinook
smolt
development
may
occur
at
temperatures
above
17
°
C
and
significant
inhibition
of
gill
ATPase
activity
and
associated
reductions
of
hypoosmoregulatory
capacity
may
occur
when
chronic
elevated
temperatures
exceed
20
°
C.
Wedemeyer
(
1980;
as
cited
in
USEPA,
2001)
noted
that
Fall
chinook
undergo
a
greater
desmoltification
rate
at
15
°
C
than
coho
do
at
that
temperature
range.
Clarke,
Shelbourn,
and
Brett
(
1981)
found
that
while
acclimation
to
brackish
water
(
15­
20ppt)
had
little
effect
on
osmoregulatory
performance
of
coho
and
chinook
reared
at
10
°
C,
it
did
maintain
performance
in
chinook
at
15
°
C
to
the
end
of
the
experiment.
The
authors
also
found
a
dramatic
regression
of
hypo­
osmoregulatory
capacity
of
chinook
smolts
held
in
fresh
water
at
15
°
C
from
week
7
to
week
13.
However,
no
such
loss
was
observed
in
fish
rear
in
10
or
15ppt
salinity.
Marine
(
1997)
found
that
smolts
from
the
control
(
13­
16
°
C)
lot
experienced
higher
ATPase
levels
and
broader
time
periods
of
elevation,
and
better
survival
in
seawater
challenges
than
smolts
reared
in
the
warmer
test
lots
(
17­
20
°
C
and
21­
24
°
C).
The
possibility
that
temperatures
above
17
°
C
exerts
stress
on
migrating
smolts
is
further
supported
by
the
work
of
Connor,
Burge,
and
Bennett
(
1999;
as
cited
in
USEPA,
2001)
that
found
subyearling
chinook
move
into
the
main
current
of
the
Columbia
River
to
avoid
increasing
nearshore
temperatures,
with
this
effect
becoming
significant
above
17
°
C.
It
is
further
supported
by
the
work
of
Sauter
and
Maule
(
1997,
1999)
which
found
cessation
in
feeding
behavior
in
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
119
Washington's
Surface
Water
quality
Standards
smolting
fall
chinook
at
temperatures
above
18
°
C
and
a
decreased
temperature
preference
(
from
17.7
to
11.2
°
C).
Exposure
to
water
temperatures
of
20
°
C
for
several
hours
induced
heat
shock
proteins
(
Sauter
et
al.,
in
review,
and
M.
Hargis,
personal
comm.;
as
cited
in
Sauter
and
Maule,
1997).
Zaugg
(
2001)
summarized
studies
done
by
Bill
Muir
of
the
NMFS.
It
was
observed
that
tagged
yearling
spring
chinook
salmon
subjected
to
advanced
photoperiod
schedules
and
a
10­
day
exposure
to
elevated
water
temperatures
(
11­
12
°
C)
prior
to
release
at
the
Dworshak
hatchery
migrated
more
rapidly
and
were
detected
in
greater
numbers
at
downstream
dams
than
controls
or
fish
exposed
only
to
advanced
photoperiods
(
no
increase
in
water
temperature).
Dr.
Zaugg
notes
that
it
appears
that
this
level
of
temperature
increase
might
actually
benefit
chinook
and
coho
salmon,
and
that
steelhead
smolts
appear
to
be
the
most
sensitive
to
elevated
temperatures.
Roper
and
Scarneccia
(
1999;
as
cited
in
USEPA,
2001)
found
that
approximately
50%
of
the
spring
chinook
emigration
occurs
in
the
range
of
12.5­
15
°
C
and
the
upper
tail
of
this
run
is
generally
complete
before
20
°
C
in
exceeded.
Connor,
Burge,
and
Bennett
(
1999;
as
cited
in
USEPA,
2001)
note
that
as
temperatures
increase
above
17
°
C,
sub­
yearling
chinook
rearing
in
nearshore
areas
of
the
Columbia
River
on
their
way
to
the
ocean
can
be
forced
to
enter
the
main
current
to
avoid
increasing
temperatures
along
river
margins.
This
is
an
indication
that
their
may
be
indirect
effects
on
smolts
caused
by
warmer
waters
in
addition
to
the
direct
effect
on
smoltification
patterns.

Coho
salmon
Zaugg
and
McLain
(
1976;
as
cited
by
Zaugg,
2001
and)
found
that
ATPase
reached
a
maximum
one
month
earlier
in
fish
reared
at
10
°
C
than
those
reared
at
6
°
C,
and
that
activity
rose
even
earlier
at
15
°
C
but
was
only
transitory
in
those
held
at
20
°
C.
Cold
water
(
6
°
C)
was
also
found
to
preserve
the
elevated
activity
while
higher
temperatures
(
10
and
15
°
C)
caused
decreases
after
an
initial
accelerated
increase.
Additionally,
Clarke
and
Hirano
(
1995)
noted;
however,
that
diurnal
and
seasonal
temperature
cycles
may
alter
the
timing
of
smolting.
Juvenile
steelhead
trout
reared
on
a
simulated
seasonal
temperature
cycle
(
6.9­
18.6
°
C)
exhibited
greater
migratory
behavior
and
more
pronounced
elevation
of
gill
sodium,
potassium­
activated
Na+
K+­
ATPase
activity
than
those
reared
at
constant
12.3
°
C
(
Zaugg
and
Wagner
1973,
and
Wagner
1974).
Adams
et
al.
(
1975;
as
cited
in
USEPA,
2001)
found
that
15
°
C
was
linked
to
impairment
of
smoltification,
ability
of
smolts
to
migrate,
and
survival
during
smolt
migration
in
coho
salmon.
Zaugg
(
2001)
noted
that
in
coho
salmon
that
had
reverted
to
parr
and
were
subsequently
released
still
migrated
to
the
lower
Columbia
River
very
rapidly,
and
migrants
captured
there
showed
elevated
ATPase
levels.
Although,
ATPase
levels
were
low
at
the
time
of
release
in
June
and
July,
these
fish
were
capable
of
regenerating
smolt
level
enzyme
activities
during
migration
in
the
warm
waters
of
the
Columbia
during
June
and
July.
Migrants
in
the
June
and
July
releases
migrated
more
rapidly
than
those
released
in
May
when
ATPase
activities
were
elevated
in
the
holding
ponds,
and
they
had
the
greatest
rate
of
survival
to
adults.

Discussion
on
Smoltification
Studies:
Page
120
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Zaugg
(
2001)
has
noted
that
there
is
considerable
risk
in
using
the
results
obtained
from
temperature
experiments
conducted
in
the
laboratory
in
predicting
what
may
occur
in
the
natural
environment
with
similar
temperatures.
Fish
held
in
confined
conditions
fail
to
undergo
full
smolt
development.
Once
released
from
the
confined
environments,
normal
smolt
development
occurs
as
downstream
migration
proceeds.
He
noted
that
while
his
earlier
work
suggested
an
upper
limit
of
12­
13
°
C
existed
for
migrating
steelhead
based
on
confined
laboratory
conditions,
we
must
now
use
some
degree
of
caution
in
applying
this
suggested
upper
limit
to
the
natural
waters.
He
noted
that
once
fish
are
actively
migrating
downstream
and
are
in
the
process
of
transformation,
it
may
take
substantially
warmer
waters
to
make
them
revert.
In
a
discussion
on
the
use
of
the
available
research
to
set
temperature
criteria,
Dr.
Zaugg
supported
the
premise
that
it
may
be
most
important
to
ensure
that
temperature
conditions
in
the
tributaries
and
headwater
systems
allow
for
the
initiation
of
the
smoltification
process.
However,
it
must
be
recognized,
however,
that
not
all
migratory
juveniles
will
have
a
long
journey
to
marine
waters,
and
not
all
will
be
able
to
hold
in
an
estuarine
environment
if
smoltification
is
not
satisfactory
upon
arrival
at
the
mouth
of
the
tributary.
Thus
some
caution
must
be
exercised
in
assuming
that
the
process
of
migration
will
be
sufficient
to
allow
smolts
to
successfully
transition
for
life
at
sea.
The
laboratory
results
for
steelhead
trout
strongly
suggest
that
a
constant
exposure
to
temperatures
greater
than
12­
13
°
C
may
hinder
ATPase
activity,
and
that
temperatures
greater
than
15­
16
°
C
may
disallow
development
altogether
in
multiple
species.
These
results
were
obtained
in
tests
conducted
at
constant
temperatures,
and
would
best
be
represented
by
weekly
or
monthly
average
temperatures.
This
position
is
supported
by
the
work
showing
that
steelhead
trout
reared
on
a
simulated
seasonal
temperature
cycle
(
6.9­
18.6
°
C
 
mean
12.79
°
C)
exhibited
greater
migratory
behavior
and
more
pronounced
elevation
of
gill
sodium,
potassiumactivated
Na+
K+­
ATPase
activity
than
those
reared
at
constant
12.3
°
C.
Treating
the
constant
temperatures
as
a
weekly
average
temperature
results
in
the
suggestion
that
a
7DADMax
limit
of
15.18­
16.18
°
C
may
be
necessary
to
allow
normal
ATPase
development
in
smolts,
and
that
a
7DADMax
of
18.18­
19.18
°
C
may
prevent
or
stop
smoltification
entirely.

To
protect
the
smoltification
capability
of
juvenile
salmonids,
it
is
concluded
that
the
7­
day
average
of
the
daily
maximum
temperatures
(
7­
DADM)
should
not
exceed
15.18­
16.18
°
C.
Except
for
the
Columbia
and
Snake
Rivers
this
temperature
threshold
is
viewed
as
appropriate
for
application
throughout
the
entire
juvenile
migration
path.
These
two
major
rivers
are
cited
as
exceptions
because
they
provide
a
significant
migratory
time
and
terminate
in
a
large
estuary;
both
features
that
have
been
well
demonstrated
to
assist
in
the
full
development
of
successful
smolts,
even
under
temperature
conditions
determined
adverse
in
laboratory
testing.

e)
Miscellaneous
Indigenous
Species
With
the
possible
exception
of
two
amphibian
species,
the
tailed
frog
and
the
torrent
salamander,
and
two
fish
species,
the
mountain
whitefish
and
the
smelt,
no
other
aquatic
organisms
have
been
identified
that
appear
as
sensitive
overall
to
temperature
increases
as
the
native
salmonids
and
char.
This
paper
has
focused
on
protecting
species
sensitive
to
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
121
Washington's
Surface
Water
quality
Standards
temperature
increases,
however,
it
is
important
to
remember
that
some
of
our
native
populations
thrive
in
waters
warmer
than
what
is
fully
protective
for
salmonids
(
Wydoski
and
Whitney,
1979;
Black,
1953;
Li
et
al.,
1993;
Reeves
et
al.,
1987;
Cech,
Mitchell,
and
Wragg,
1984,
and
others).

For
the
water
quality
standards
to
be
fully
protective
in
the
ecological
sense,
they
must
recognize
the
continuum
of
temperature
changes
from
upstream
to
downstream
and
between
ecoregions
and
waterbody
forms.
The
water
quality
standards
should
acknowledge
that
certain
waters
that
are
naturally
not
fully
protective
for
salmonids
may
be
ideally
suited
for
other
native
species.
It
is
appropriate
that
some
of
these
naturally
warmer
waters
have
separate
standards
that
consider
more
accurately
the
warm
water
tolerant
communities
that
have
historically
existed.

i)
Sensitive
Amphibians
Three
stream
dwelling
amphibians
are
of
concern
when
it
comes
to
temperature
alterations.
These
are
the
Rhycotriton
species
(
3
species
of
torrent
salamanders),
the
Dicamptodon
species
(
Pacific
and
Copes
giant
salamanders),
and
Ascaphus
truei
(
tailed
frog).

Rhyacotriton
species
are
found
in
temperatures
ranging
from
5.9­
10.9
°
C,
and
eggs
have
been
found
at
8­
9
°
C
then
taken
to
the
lab
and
incubated
successfully
at
8­
9
°
C.
Like
most
amphibians,
Rhyacotriton
can
withstand
high
daily
peak
temperatures.
When
acclimated
in
a
laboratory
to
temperatures
of
13­
14
°
C,
Rhycotriton
has
a
critical
thermal
maximum
(
CTM)
of
27.8­
29
°
C
(
Kelsey,
1998).
Welsh
and
Lind
(
1996)
suggest
that
suitable
temperatures
for
Rhyacotriton
are
from
6.5­
15
°
C,
with
highest
abundance
of
salamanders
occurring
in
a
narrow
range
of
about
8­
13
°
C.
The
authors,
using
unpublished
data
developed
while
conducting
critical
thermal
maximum
testing,
note
that
signs
of
thermal
stress
were
noted
at
17.2
°
C.

Dicamptodon
species
have
larvae
noted
to
develop
at
12­
16
°
C
in
the
field.
Larvae
may
develop
for
2
years
prior
to
becoming
adult
salamanders.
Adults
spawn
in
the
spring
when
waters
are
warming,
but
may
also
spawn
in
the
fall.
While
the
Pacific
giant
salamander
becomes
a
terrestrial
salamander,
the
Copes
giant
salamander
does
not
(
Kelsey,
1998).

Ascaphys
truei
is
typically
found
in
waters
from
4.4­
14
°
C.
Embryonic
development
can
occur
between
5­
18
°
C.
Tadpoles
from
1­
2
years
old
prefer
waters
around
5­
8
°
C,
but
3­
4
year
olds
prefer
waters
of
12­
16
°
C.
A.
truei
larvae
have
a
critical
thermal
maximum
of
28.9­
30.1
°
C
and
adults
of
23­
24
°
C;
when
both
are
acclimated
at
23
°
C.
The
nine
day
LT50
for
tailed
frog
larvae
was
determined
to
be
23
°
C.
Tadpoles
may
develop
from
1
to
4
years,
depending
on
the
individual
stream,
before
metamorphosis
(
Kelsey,
1998).
Hawkins
et
al
(
1988)
found
that
variations
in
tadpole
densities
were
high
at
maximum
stream
temperatures
below
18
°
C
and
below
that
temperature
substrate
size
and
imbeddedness
characteristics
determined
dispersal
patterns.
The
authors
suggest
that
low
densities
in
nonforested
Page
122
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
watersheds
may
be
better
explained
by
the
sensitivity
of
adults
to
desiccation
rather
than
the
effect
of
moderately
warm
water
temperatures.

As
noted
by
Hawkins
et
al.
(
1988)
tailed
frog
(
Ascaphus
truei)
tadpoles
appear
to
be
particularly
well
suited
to
life
in
swift
streams.
The
body
is
streamlined
and
ventrally
flattened,
and
the
mouth
is
modified
into
a
powerful
suctorial
disc.
Tadpoles
use
the
oral
sucker
to
cling
to
rock
surfaces,
move
against
the
flow
of
water,
and
scrape
periphyton
from
streambed
surfaces.
Diatoms
are
apparently
the
main
food
source
for
these
tadpoles
(
Metter,
1964;
as
cited
by
Hawkins
et
al.,
1988).
In
most
West
Coast
streams,
eggs
hatch
in
late
summer
and
tadpoles
transform
to
adults
almost
exactly
two
years
later
(
Metter,
1967;
as
cited
by
Hawkins
et
al.,
1988).

Claussen
(
1973)
found
that
adult
A.
truei
had
an
incipient
lethal
temperature
limit
of
24.1
°
C
after
45
hours
of
exposure
and
25.8
°
C
after
7.3
hours,
and
that
adults
preferred
temperatures
of
16.5­
18
°
C
in
laboratory
tests
conducted
on
wet
sand.
Claussen
reported
the
adult
frogs
were
kept
in
good
health
in
the
laboratory
in
terrariums
kept
at
room
temperature
(
19­
24
°
C)
for
many
months.
Brown
(
1975;
as
cited
by
Hawkins,
1988)
reportedly
found
that
18.5
°
C
was
the
upper
limiting
temperature
for
egg
development.
De
Vlaming
and
Bury
(
1970;
as
cited
by
Hawkins,
1988)
are
reported
to
have
found
that
age
1+
tadpoles
preferred
5­
8
°
C
and
age
2+
tadpoles
preferred
12­
16
°
C,
and
that
both
cohorts
avoided
temperatures
greater
than
22
°
C
and
50%
of
1+
animals
died
within
29
days
at
23
°
C.

Welsh
(
1990)
studied
the
distribution
of
A.
truei
over
a
three
year
period
at
stream
sites
in
the
Pacific
Northwest
and
found
temperature
to
be
an
excellent
predictor
of
tailed
frog
abundance.
Higher
numbers
of
tailed
frogs
occurred
in
streams
with
lower
temperatures
and
the
highest
stream
temperature
at
which
he
had
observed
tailed
frogs
was
14.3
°
C.
Welsh
suggested
his
work
showed
that
while
tailed
frogs
were
found
at
maximum
temperatures
as
high
as
18
°
C,
their
densities
began
increasing
only
at
stream
temperatures
below
about
14.3
°
C
and
were
characteristically
highest
in
waters
with
temperatures
below
12
°
C.

Hawkins
et
al
(
1988)
sampled
small
streams
near
Mt.
St.
Helens,
Washington
between
1985­
1987
and
found
that
streams
with
little
forest
cover
and
maximum
temperatures
near
20
°
C
had
few
A.
truei
while
streams
in
completely
or
partially
forested
basins
with
maximum
stream
temperatures
less
than
18
°
C
had
relatively
high
tadpole
densities.
Variation
in
density
within
streams
with
maximum
temperatures
below
18
°
C
were
not
associated
with
temperature
differences
but
were
instead
associated
with
substrate
size
and
embeddedness
characteristics.
In
the
Hawkins
et
al.
(
1988)
suggest
that
while
their
work
does
not
substantiate
the
opinion
that
adverse
effects
occur
up
to
20
°
C
on
tadpole
density,
temperatures
in
this
range
are
within
the
critical
limits
found
by
other
authors
and
their
results
should
be
used
with
care.
Hawkins,
Feminella,
and
Crisafulli
(
unpublished
status
survey)
found
that
in
the
streams
around
Mt.
St.
Helens,
Washington,
maximum
tadpole
densities
were
associated
with
mean
maximum
summer
water
temperatures
of
16
°
C
and
note
that
since
1988
tadpole
densities
have
been
consistently
higher
in
nonforested
(
warmer)
than
in
forested
(
cooler)
streams.
Crisafulli
(
personal
communication)
noted
that
the
ability
of
tailed
frog
to
thrive
in
the
deforested
areas
of
the
volcano's
blast
zone
may
be
related
to
the
Evaluating
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in
Page
123
Washington's
Surface
Water
quality
Standards
generally
cool
moist
weather
that
is
experienced
in
that
region
of
the
state
which
helps
avoid
desiccation
of
adult
frogs.
The
concern
being
that
work
done
in
other
regions
associating
tailed
frog
abundance
to
stream
temperatures
may
be
measuring
the
association
of
cool
moist
mature
forest
conditions
to
temperature
rather
than
accurately
gauging
the
importance
of
the
specific
stream
temperatures
to
tailed
frog
health.
Since
flow,
sediment,
and
riparian
area
conditions
well
all
tend
to
be
more
favorable
in
the
cooler
headwater
areas,
this
may
confound
efforts
to
gauge
the
importance
of
temperature
compared
with
these
other
commonly
associated
factors.

Larvae
of
all
of
these
stream
breeding
amphibians
rely
primarily
on
the
perennial
flows
of
1st
and
2nd
order
streams
(
mostly
Type
4
and
5,
but
also
some
smaller
Type
3
streams).
Their
rates
of
growth,
and
thus
time
prior
to
metamorphosis,
is
affected
by
temperature
and
food.
Warmer
waters
within
their
tolerance
range
may
speed
up
development
(
Kelsey,
1998).

The
paucity
of
controlled
research
data
on
the
temperature
requirements
of
amphibians,
and
the
potentially
strong
conflicts
among
the
existing
research,
makes
establishing
a
temperature
standard
aimed
at
ensuring
their
protection
somewhat
problematic.
With
lethal
tolerances
as
high
as
most
salmon
and
trout,
the
reported
distributional
associations
with
temperature
must
be
considered
with
caution.
Since
these
amphibians
commonly
occur
high
in
the
drainage,
often
at
and
above
the
upper
limits
of
many
fish
species,
they
are
typically
able
to
associate
with
shallow
groundwater
dominated
seeps
and
springs.
Stream
breeding
amphibians,
particularly
the
torrent
salamanders
and
the
tailed
frog,
may
in
fact
have
lower
optimal
temperatures
than
salmon
and
trout,
but
at
this
point
the
scientific
information
is
not
strong
enough
to
reach
this
conclusion
and
does
not
suggest
that
establishing
a
species­
specific
temperature
limits
are
clearly
warranted.
There
is
clearly
a
need
for
more
controlled
studies
on
the
temperature
requirements
of
our
stream
breeding
amphibians,
and
the
question
of
whether
more
stringent
temperature
standards
needs
to
be
set
for
these
species
should
be
revisited
periodically
to
ensure
that
new
information
and
understandings
are
taken
into
account
in
setting
and
revising
water
quality
standards.

ii)
Other
Sensitive
Fish
Species
Smelt
and
mountain
whitefish
were
identified
through
the
literature
as
possessing
sensitive
temperature
limits.
The
studies
on
smelt
indicate
they
have
a
lower
lethal
temperature
limit
than
do
the
salmonids
and
a
lower
optimum
temperature
preferendum.
Longfin
smelt
(
Spirinchus
thaleichthys)
was
identified
as
having
a
limit
of
occurrence
of
18.3
°
C
by
Wydoski
and
Whitney
(
1979).
This
corresponds
well
to
acute
exposure
testing
using
the
Eulachon
smelt
(
Thaleichthys
pacificus)
by
Snyder
and
Blahm
(
1971)
who
found
a
change
from
10
to
18
°
C
resulted
in
50%
mortality
to
adults
in
less
than
one
hour,
and
50%
mortality
occurring
in
26
minutes
at
an
exposure
of
21
°
C.
While
temperature
increases
from
10
°
C
to
13
°
C
and
15
°
C,
did
not
induce
mortality
over
a
50­
hour
holding
time,
none
of
the
females
exposed
to
the
higher
test
temperatures
deposited
their
eggs.
USEPA
(
1971)
notes
that
in
temperature
studies
on
the
eulachon,
Smith
and
Saalfeld
(
1955;
as
cited
in
USEPA,
1971)
reported
the
fish
entered
the
Columbia
River
when
the
temperature
was
between
2
and
10
°
C
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Evaluating
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Life
in
Washington's
Surface
Water
quality
Standards
but
they
migrate
up
to
and
beyond
the
Cowlitz
River
(
RM
68)
when
the
Columbia
is
approximately
4.4
°
C.
The
smelt
run
was
delayed
five
weeks
from
entering
the
Cowlitz
River
because
of
low
water
temperatures
during
December
1968
and
January
1969
(
Snyder,
1970;
as
cited
in
USEPA,
1971).
Eulachon
eggs
appear
to
be
more
tolerant
than
adults
to
temperature
increases.
The
eggs
can
withstand
a
temperature
of
14
°
C
from
a
base
temperature
of
4
to
8
°
C
without
appreciable
mortalities
(
Parente
and
Ambrogetti,
1970;
as
cited
in
USEPA,
1971),
but
a
3
°
C
increase
halts
maturation
of
adult
females.
In
tests
in
1968
and
again
in
1969,
it
was
observed
that
female
smelt
exposed
to
water
heated
3.9
°
C
above
river
temperatures
were
reluctant
to
spawn.
Adult
female
smelt
are
less
tolerant
of
temperature
changes
than
other
fish.
Bell
(
1986)
in
a
general
review
on
the
temperature
requirements
of
fish
suggested
that
the
range
for
smelt
was
3.8­
12.7
°
C,
and
the
preferred
spawning
range
was
7.2­
8.3
°
C.
The
spawning
range
identified
by
Bell,
closely
matches
that
observed
for
longfin
smelt
in
the
Cedar
River
in
Washington.
Wydoski
and
Whitney
(
1979)
suggest
that
spawning
occurs
primarily
in
late
February
(
with
a
range
of
mid­
January
to
mid­
April)
when
the
river
was
between
4.4­
7.2
°
C.
They
also
note
a
British
Columbia
stock
that
hatches
at
9.4­
10.6
°
C
approximately
25
days
after
spawning.
Given
that
adult
spawners
and
outgoing
juveniles
may
be
in
fresh
waters
as
late
as
March
to
mid­
April,
and
their
temperature
requirements
may
be
more
strict
than
most
salmonids,
the
protection
of
smelt
is
an
important
consideration
in
setting
water
quality
standards.
In
waters
supporting
smelt,
it
is
estimated
that
the
7­
day
average
of
the
daily
maximum
temperatures
should
not
exceed
12­
14
°
C
prior
to
May
1;
with
no
single
daily
maximum
temperature
greater
than
16
°
C
to
fully
protect
reproduction.
This
will
allow
most
stocks
to
spawn
and
have
the
newly
hatched
juveniles
return
to
the
sea
(
or
lake
where
a
landlocked
population)
under
tolerable
temperature
conditions.

Information
on
the
temperature
requirements
of
the
mountain
whitefish
(
Prosopium
williamsoni
Girard)
is
more
limited
to
that
for
the
smelt.
Eaton
et
al.
(
1995)
calculated
that
a
weekly
mean
temperature
of
23.2
º
C
was
the
95th
percentile
of
the
distribution
of
mountain
whitefish
in
field
studies
and
suggested
this
value
as
a
measure
of
the
species
temperature
tolerance.
Wydoski
and
Whitney
(
1979)
and
Daily
(
1971)
note
that
mountain
whitefish
are
found
in
both
streams
and
lakes
throughout
the
state
of
Washington,
generally
in
large
streams
with
average
temperatures
of
8.8
to
11.1
°
C.
Whitefish
are
generally
nocturnal
autumn
spawners
that
spawn
from
October
into
December
and
at
temperatures
below
4.4
to
5.5
°
C
have
been
associated
with
spawning
activity
(
Brown,
1952;
as
cited
by
Daily,
1971).
Mountain
whitefish
eggs
require
low
temperatures
for
optimum
development.
They
hatch
in
about
5
months
at
1.7
°
C
and
in
about
1
month
at
8.9
°
C
(
Sigler
and
Miller
1963,
and
Simon
1946;
as
cited
in
Davis,
1971).
Daily
(
1971)
noted
that
adult
mountain
whitefish
are
reported
to
prefer
pool
and
meadow
areas
of
cool
streams
where
water
depths
exceed
three
feet.
Sigler
(
1951)
and
La
Rivers
(
1962)
are
cited
by
Daily
(
1971)
as
suggesting
high
water
temperatures
limit
whitefish
to
elevations
above
4,500
feet
in
California,
Nevada,
and
Utah.

Northcote
and
Ennis
(
1994)
note
that
mountain
whitefish
are
often
the
most
abundant
sport
fish
species
in
many
rivers
of
western
North
America
(
Sigler
and
Miller
1963,
and
Brown
1971;
as
cited
in
Northcote
and
Ennis,
1994).
Hagen
(
1970;
as
cited
in
Northcote
and
Ennis,
1994)
found
that
in
Phelps
Lake
spawning
began
in
September
at
temperatures
over
11
°
C
but
Evaluating
Standards
for
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Aquatic
Life
in
Page
125
Washington's
Surface
Water
quality
Standards
may
extend
into
November
at
near
7
°
C.
Spawning
starts
in
late
September
in
the
Sheep
River
and
extends
to
mid­
October
at
temperatures
varying
daily
from
8
to
0
°
C
(
Thompson
and
Davies,
1976;
as
cited
in
Northcote
and
Ennis,
1994).
Other
authors
reportedly
found
spawning
at
temperatures
as
high
as
9
°
C
and
as
low
as
near
zero,
but
usually
in
the
3
to
5
°
C
range.
Brown
(
1952;
as
cited
in
Northcote
and
Ennis,
1994)
found
that
in
the
west
Gallatin
River,
Montana,
fish
do
not
spawn
until
water
temperatures
drop
below
5.5
°
C
with
peak
spawning
activity
later
in
higher
elevation
tributaries
at
temperatures
just
over
2
°
C.
Rajagopal
(
1975,
1979;
as
cited
by
Northcote
and
Ennis,
1994)
studied
the
developmental
rate
of
mountain
whitefish
eggs
and
gave
6
°
C
as
the
upper
optimal
temperature
for
successful
development.
At
temperatures
between
9
and
11
°
C
some
hatching
occurred
but
there
were
high
levels
of
mortality
and
abnormality;
all
eggs
died
at
temperatures
of
12
and
15
°
C.

The
information
is
not
sufficient
to
justify
setting
a
separate
criteria
for
mountain
whitefish,
and
since
mountain
whitefish
occur
in
general
along
with
salmon
and
trout
in
our
streams
and
lakes,
any
assignment
of
criteria
to
protect
these
other
salmonids
would
also
be
used
to
protect
mountain
whitefish
populations.
Based
on
the
above
cited
literature
findings,
it
appears
that
mountain
whitefish
have
summer
growth
tolerances
similar
to
the
Pacific
salmon
and
trouts,
but
they
may
have
spawning
requirements
that
are
more
sensitive.
The
generally
late
spawning
habits
of
the
mountain
whitefish
assist
in
their
protection;
however,
human
actions
that
cause
uncharacteristic
seasonal
warming
may
cause
harm
to
mountain
whitefish
during
their
incubation.
In
waters
used
for
the
incubation
of
mountain
whitefish,
average
temperatures
during
incubation
should
be
maintained
at
or
below
6
°
C.
The
strength
of
the
literature
findings
is
not
sufficient
for
setting
a
separate
temperature
threshold
for
mountain
whitefish.
It
can,
however,
be
used
in
support
of
setting
incubation
criteria
for
the
Pacific
salmon
and
trout.
Treating
this
average
temperature
as
either
a
weekly
average
or
season­
wide
average
temperature
helps
bound
the
estimate
for
an
incubation
threshold
and
allows
conversion
to
the
standard
7DADMax
metric
used
in
this
document.
Taking
this
approach
results
in
the
estimate
that
to
fully
protect
mountain
whitefish
incubation
the
7DADMax
temperature
should
be
maintained
below
7.47­
10.64
°
C
(
9.1).

iii)
Stream
Macroinvertebrates
Only
a
relatively
small
number
of
studies
were
found
that
tested
the
thermal
tolerance
of
stream
macroinvertebrates.
Most
of
the
species
examined
have
thermal
limits
higher
than
the
Pacific
salmon
and
trout
species
found
in
Washington
(
Sprague,
1963;
Nebeker
and
Lemke,
1968;
Moulton
et
al.,
1993;
Sherberger
et
al.,
1977,
as
cited
in
Beschta
et
al.,
1987;
Craddock,
1970
and
Hair,
1971,
as
cited
in
USEPA,
1971).
However,
two
sets
of
studies
were
found
that
suggest
individual
species
may
sometimes
as
sensitive,
and
in
some
cases
more
sensitive,
as
our
indigenous
salmonids.

Gaufin
and
Hern
(
1971;
as
cited
in
Moulton
et
al.,
1993)
reported
a
range
of
21.7
to
30.1
º
C
in
mean
lethal
level
(
median
lethal
temperature)
for
six
species
of
caddisflies
held
at
6.4
º
C.
The
lowest
lethal
level
21.7
º
C
was
observed
for
Parapsyche
elsis
Milne,
(
Hydropsychidae)
Page
126
Evaluating
Standards
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Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
an
inhabitant
of
cold,
fast­
flowing
mountain
streams
while
the
highest
lethal
level
(
30.1
º
C)
was
recorded
for
Hydropsyche
sp.
collected
from
a
marsh­
lake
outflow.

USEPA
(
1971)
conducted
96­
hour
lethality
studies
to
examine
the
heat
tolerance
of
late
instar
larvae
of
15
species
of
aquatic
insects
and
one
species
of
amphipod.
USEPA
reported
that
a
marked
difference
in
sensitivity
was
apparent
in
the
different
species.
A
mayfly,
Cinygmula
par
Eaton,
died
at
11.7
°
C
and
was
the
most
sensitive
of
all
the
species
tested.
This
species
is
found
in
very
cold
clear
mountain
streams.
The
freshwater
shrimp,
Gammarus
limnaeus
Smith,
proved
to
be
surprisingly
sensitive
to
temperature
increases,
with
50%
lethality
at
14.5
°
C.
Ephemerella
doddsi
Needham,
a
small,
widely
distributed
mayfly
characteristic
of
cold
turbulent
streams
in
the
Intermountain
Region,
was
also
very
sensitive
with
a
TLm
(
median
lethal
threshold)
value
of
15.4
°
C.
A
lotic
species
of
mayfly,
Hexagenia
limbata
Guerin,
was
much
more
tolerant
than
other
mayflies
tested
with
a
TLm
of
26.6
°
C.
Considerable
difference
existed
between
the
three
stoneflies
tested.
Isogenus
aestivalis
(
Needham
and
Claassen)
was
quite
sensitive,
50%
dying
at
16
°
C,
while
Pteronarcella
badia
(
Hagen)
and
Pteronarcys
californica
Newport,
two
closely
related
species
survived
increases
to
24.6
and
26.6
°
C,
respectively.
Six
species
of
caddis
flies
were
tested
and
clearly
reflected
thermal
differences
in
their
habitat
requirements.
Parapsyche
elsis
Milne,
which
is
largely
restricted
to
cold,
fast
flowing
mountain
streams,
had
a
TLm
of
21.8
°
C
while
Hydropsyche
sp.
Taken
from
a
slow
flowing
stream
draining
a
marshy
lake
was
very
tolerant
with
a
TLm
of
30.1
°
C.
In
long­
term
(
12­
30
days)
thermal
bioassays
of
five
aquatic
insects,
the
stonefly
Pternorcella
badi
was
most
sensitive
with
50%
mortality
occurring
at
18.1­
20.5
°
C
within
24­
30
days
(
its
96­
hour
TLm
was
22.55
°
C).
Ephemerlla
grandis
Eaton
experienced
50%
mortality
in
12
days
at
21.5
°
C
(
its
96­
hour
TLm
was
also
21.5
°
C).
Pteronarcys
californica
Newport
experienced
50%
mortality
in
25
days
at
20
°
C
(
its
96­
hour
TLm
was
27
°
C).
None
of
the
three
most
sensitive
species
tested
in
the
96­
hour
tests
(
C.
par,
E.
doddsi,
and
G.
limnaeus)
were
included
in
these
long­
term
tests.

Summary
of
macroinvertebrate
temperature
requirements:
While
few
studies
have
been
conducted
to
test
the
thermal
limits
of
macroinvertebrate
species,
those
that
have
been
reported
in
the
literature
suggest
that
individual
species
may
be
significantly
more
sensitive
than
the
salmonids
and
other
fish
examined
in
this
review
document.
Tests
reviewed
focused
only
on
mortality
as
an
endpoint,
so
any
sublethal
effects
of
temperature
on
macroinvertebrates
(
such
as
reproductive
impairment)
remain
undocumented.
For
the
lethality
testing,
even
short
term
exposure
(
4­
days)
resulted
in
50
percent
mortality
at
constant
temperatures
from
range
of
11.7­
16
°
C,
with
three
of
the
four
species
having
LT50
values
between
14.5­
16
°
C.
These
sensitive
species
are
all
ones
that
naturally
occur
in
high
mountain
streams.
For
the
purpose
of
this
analysis
the
lowest
value
will
be
assumed
to
be
anomalous,
and
the
range
of
14.5­
16
°
C
will
be
used
to
represent
the
range
for
sensitive
species.
For
the
other
macroinvertebrates
tested,
4­
day
LT50
values
(
21.5­
30.1
°
C)
were
above
those
typically
found
for
salmonid
species.
These
two
groups
of
macroinvertebrates
are
treated
separately
in
this
analysis,
since
one
best
represents
species
that
would
be
found
in
headwater
regions
and
the
other
best
represents
those
which
would
be
found
at
lower
altitudes
and
in
more
main
stem
stream
areas.
Treating
both
groups
of
species
as
if
their
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
127
Washington's
Surface
Water
quality
Standards
corresponding
laboratory
tests
correlate
as
if
they
were
either
daily
average
or
weekly
average
temperatures
(
and
then
converting
to
a
7DADMax
metric)
bounds
the
estimate
for
an
appropriate
field­
based
temperature
criteria.
Conversions
assume
that
summer
diel
temperatures
average
4­
6
°
C
(
similar
to
what
was
previously
assumed
for
salmon
and
trout).
Based
on
the
above
information
and
recommended
conversion
methodology,
a
temperature
threshold
can
be
estimated
that
will
prevent
lethality
in
macroinvertebrates.
Headwater
assemblages
of
macroinvertebrates
should
be
expected
to
experience
50
percent
mortality
if
the
7DADMax
temperature
exceeds
15.08
 
19.18
°
C
(
17.13)
[
the
lower
end
of
the
range
would
decrease
to
12.38
°
C
(
midpoint
15.78)
if
the
most
sensitive
species
tested
were
included].
Main
stem
and
lower
elevation
assemblages
of
macroinvertebrates
should
be
expected
to
experience
50
percent
mortality
if
the
7DADMax
temperature
exceeds
22.08­
33.28
°
C
(
27.68)
 
although,
greater
confidence
should
be
placed
on
the
lower
portion
of
this
range
(
22.08­
27.68
°
C,
with
a
median
of
24.88)
because
the
range
of
macroinvertebrate
species
tested
was
not
limited
to
those
that
would
occur
in
waters
inhabited
by
salmonids
(
typical
mid
to
lower
elevation
rivers
in
Washington).
To
prevent
any
mortality
the
temperatures
associated
with
50
percent
mortality
should
be
decreased
by
2
°
C,
similar
as
was
explained
previously
for
converting
lethality
studies
for
salmonids.
To
protect
macroinvertebrates
from
lethal
effects
in
cold
water
stream
assemblages
the
7DADMax
temperature
should
not
exceed
13.08­
17.18
°
C
(
15.13)
in
headwater
streams,
and
not
exceed
20.08­
25.68
°
C
(
22.88)
in
the
low
lying
streams.
Page
128
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
f)
Miscellaneous
Indigenous
Fish
Species
The
following
graphical
summary
includes
only
those
species
for
which
temperature
information
was
available.
In
many
cases
the
information
pertains
only
to
their
general
habitat
associations
and
life­
history
traits.
Information
is
not
adequate
to
propose
temperature
thresholds
for
any
of
these
species;
however,
the
information
is
useful
in
evaluating
whether
the
temperature
thresholds
recommended
to
protect
the
native
salmonids
and
char
also
appear
protective
of
these
other
species.

Western
Brook
Lamprey
10
(
Spawn
peak,
May)

Pacific
Lamprey
12.8
(
Too
cold
for
incubation)
15.6
(
Successful
incubation)
15­­­
25
(
Successful
incubation)

White
Sturgeon
8.9­­­
17.2
(
Spawn,
May­
July)
10­­­
18
(
Spawning)
14
(
Optimal
Incubation)
14
(
Most
spawning
occurred)
17.2­
17.8
(
summer
occupation)
18
(
Elevated
embryonic
mortality)
18­­­
20
(
Limited
spawning
occurs)
20
(
Complete
embryonic
mortality)

Eulachon
Smelt
2­­­
10
(
River
temp
during
spawning
migration)
<­
13
(
Successful
egg
deposition)
18
(
Rapidly
lethal
to
adults)

Longfin
Smelt
18.3
(
Limit
to
occurrence)
4.4­­­
7.2
(
Spawn,
primarily
February)
9.4­
10.6
(
Eggs
hatch)

Smelt
(
general)
7.2­
8.3
(
Preferred
spawning)
8.3
(
Optimal
hatch)

Mountain
Whitefish
8.8­­­
11.1
(
Average
temp
of
habitat)
<
­
23.2
(
95%
of
distribution)
6
(
Upper
optimal
incubation)

Chiselmouth
17
­
>
(
Spawn
in
summer)

Redside
shiner
6.7­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­­
23.9
(
Range)
10
­
>
(
Spawn,
April­
July)
12.8­­­­­­­­­­­­­­­­­­
20
(
Summer
preference)
19­­­­­­­­
22
(
Fluct.
better
than
12­
15)
25
(
24
hr
LT50
at
9­
11C
acclimation)
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
129
Washington's
Surface
Water
quality
Standards
27.6
(
24
hr
LT50
at
14C
acclimation)
Longnose
Dace
11.7
­­­
>
(
Spawn,
June­
early
July)
12.8­­­
21.2
(
Prefers)
19.4­­­
30
(
Found
over
two
years)

N.
Pikeminnow
16.1­­­
24.4
(
Prefers)
18.3
(
Spawn,
late
May­
July,
hatch)
29.3
(
24
hr
LT50
at
19­
22C
acclimation)

Tui
Chub
12.8­­­
15.6
(
Spawn,
May­
June)

Peamouth
12.2
­
>
(
Spawn,
late
May­
early
June)
26.6
(
24
hr
LT50
at
14C
acclimation)
27
(
24
hr
LT50
at
11.5C
acclimation)

Lake
Chub
13.9
(
In
streams)
18.9
(
Spawn,
Apr­
Jun,
lakes)

Speckled
Dace
17.8
 
18.9
(
Eggs
hatch)

Largescale
sucker
29.4
(
24
hr
LT50
at
19C
acclimation)

Sucker
(
generic
ref.)
11.7­­­
21.7
(
Prefers)

Longnose
Sucker
5
­
>
(
Spawn,
early
spring)

Mtn.
Sucker
12.8­­­
21.1
(
Summer
preference)

11.1­­­
18.9
(
Spawn,
June­
July)

Burbot
1.7
(
Spawn,
Jan­
Feb)
21.2
(
Final
preferendum)

T­
S.
Stickleback
17.8
(
Spawn,
May­
Aug,
hatch)
26
(
LT50
in
6
days)

Shorthead
Sculpin
<
­
15.6
(
Prefers)
23.8
(
Found)

Piute
Sculpin
15
­
>
(
Frequents)
25
(
Found)
12.2
(
Spawn,
May­
June)

Prickly
Sculpin
10­­­
17.8
(
Typical)
24.1
(
24
hr
LT50,
18­
19C
acclimation)
27.8
(
Found)

Margined
Sculpin
12.8­­­
18.9
(
Typical)

Mottled
Sculpin
12.8­­­
18.3
(
Preferred)
Page
130
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
15.6­­­
23.3
(
Found
over
two
years)
16.5
(
Preferred)

21.1
(
Found)
10­­­
15.6
(
Spawn,
Feb­
June,
hatch)

Riffle
Sculpin
<
­
15.6
(
Prefers)
22.2
(
Found)
<
­
27.8
(
Survive
in
lab)
Reticulate
Sculpin
10­­­
17.8
(
Typical)
25.6
(
Found)
Starry
Founder
11.1
(
Ave.
temp.,
spawn
late
Nov­
Feb)

Although
the
temperature
preferences
of
most
of
these
species
appears
to
overlap
that
of
the
upper
end
of
the
fully
protective
temperature
range
for
rainbow
trout,
some
of
these
species
will
thrive
in
waters
warmer
than
what
is
fully
protective
for
rainbow
trout.
Species
such
as
dace
and
redside
shiner
may
actually
rely
on
their
ability
to
tolerate
warmer
water
to
maintain
strong
populations
where
they
exist
in
sympatry
with
rainbow
trout
or
other
salmonids.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
131
Washington's
Surface
Water
quality
Standards
g)
Summary
of
Temperature
Requirements
for
Indigenous
Aquatic
Life
i)
Cold
Water
Species
The
following
table
summarizes
the
individual
conclusions
made
previously
to
protect
the
state's
cold
water
aquatic
habitats.
For
the
two
salmonid
guilds
examined,
the
life­
stages
are
presented
along
with
other
thermal
stressors
that
would
influence
the
health
of
these
lifestages
(
e.
g.,
disease,
and
interactions
with
other
associated
community
and
prey
species).
This
approach
is
useful
in
identifying
what
temperature
criteria
would
be
most
appropriate
to
provide
for
a
fully
protective
thermal
environment.
The
conclusions
are
provided
as
summary
statements
in
Table
4.25
below.

Table
4.25.
Ranges
within
which
lie
temperatures
likely
to
fully
protect
specific
species
and
lifestages

Requirements
by
Species
Guild
and
Life
Stage
7DADMax
Temperature
Range
(
°
C)
Midpoint
of
Range
(
°
C)

Bull
Trout
and
Dolly
Varden
(
Char)
Char
spawning
and
incubation
7.31­
8.32
7.82
Char
juvenile
rearing
12.61­
13.96
13.29
Disease
­
Virtual
elimination
of
warmwater
disease
effects
in
salmon
and
trout
12.58­
16.18
14.38
Macroinvertebrate
lethality
in
headwater
streams
13.08­
17.18
15.13
Char
lethality
(
7­
day
exposure)
20.73­
212.88
21.31
Summary:
Temperatures
(
7DADMax)
should
be
below
7.5­
8
°
C
at
the
time
of
spawning
for
char
and
below
13­
13.5
°
C
outside
of
the
incubation
period.
This
temperature
regime
will
also
provide
full
protection
from
warm
water
disease
and
support
sensitive
headwater
species
of
macroinvertebrates.

Salmon
and
Trout
Waters
Spawning
and
Incubation
Salmon
and
trout
spawning
and
incubation
12.55­
13.92
13.24
Reproduction
of
Smelt
(
prior
to
May
1)
12­
14
13
Summary:
Temperatures
(
7DADMax)
should
be
below
13­
13.5
°
C
at
the
time
of
spawning
for
salmon
and
trout.
This
temperature
should
also
fully
protect
the
reproduction
of
smelt
and
other
non­
salmonid
species.

Juvenile
rearing
Page
132
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Salmon
and
trout
juvenile
rearing
15.18­
18.05
16.62
Juvenile
smoltification
(
typically
ends
prior
to
late
June­
late
August
when
summer
temperatures
peak)
15.18­
16.18
15.68
Macroinvertebrate
lethality
in
headwaters
13.08­
17.18
15.13
Disease
­
Virtual
elimination
of
warmwater
disease
effects
12.58­
16.18
14.38
Disease
­
Avoiding
serious
rates
of
infection
and
mortality
15.58­
19.18
17.38
Macroinvertebrate
lethality
in
main­
stems
20.08­
25.68
22.88
Summary:
Temperatures
(
7DADMax)
should
be
below
16­
16.5
°
C
to
fully
protect
juvenile
rearing
of
salmon
and
trout.
However,
in
lower
portions
of
rivers
where
smoltification
interference
is
less
likely,
the
natural
macroinvertebrate
community
would
not
be
expected
to
include
the
most
sensitive
taxa,
the
waters
tend
to
be
more
productive
for
food
organisms,
and
where
holding
by
ripe
adult
fish
is
naturally
uncharacteristic;
slightly
warmer
temperatures
(
17­
17.5
°
C)
may
still
be
fully
protective
of
the
indigenous
aquatic
community
during
the
summer
rearing
period.
In
waters
supporting
the
eastside
redband
trout,
a
7DADMax
of
17.5­
18
°
C
should
also
be
considered
fully
protective.

Adult
migration
Direct
Lethality
(
7­
day)
to
salmon
and
trout
21.09­
23.36
22.23
Direct
lethality
(
7­
day
exposure)
to
char
(
recognizing
use
by
migratory
populations).
20.73­
21.88
21.31
Barriers
to
migration
in
salmon
and
trout
20.05­
24.6
22.1
Non­
barrier
migratory
effects
18.1­
20.4
19.25
Disease
­
Avoiding
serious
rates
of
infection
and
mortality
15.58­
19.18
17.38
Salmon
and
trout
juvenile
rearing
(
used
here
as
a
general
sign
of
low
thermal
stress)
15.18­
18.05
16.62
Summary:
Adult
migrants
are
likely
to
be
detrimentally
impacted
at
a
7DADMax
above
17­
19
°
C;
and
barriers
to
migration
and
direct
mortality
should
be
expected
when
7DADMax
temperatures
exceed
21.5­
22
°
C.
One
day
maximum
temperatures
for
which
lethality
should
be
expected
to
begin
would
be
22.5­
23
°
C.

Adult
holding
prior
to
spawning
Prespawning
effects
13.48­
14.96
14.22
Disease
 
Virtual
elimination
of
warmwater
disease
effects.
12.58­
16.18
14.38
Summary:
Locations
in
streams
where
adult
migrants
hold
the
week
or
two
just
prior
to
spawning
should
have
temperatures
not
exceeding
a
7DADMax
of
14.5
°
C
to
avoid
prespawning
losses
of
adults
or
potential
offspring.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
133
Washington's
Surface
Water
quality
Standards
ii)
Warm
water
Species
Water
quality
standards
must
be
applied
in
some
fashion
to
all
types
of
waterbodies;
however,
some
waters
will
naturally
have
higher
temperatures
than
what
would
support
healthy
populations
of
salmonids.
A
natural
warm
water
fish
community
in
Washington
would
be
characterized
by
the
presence
of
redside
shiner;
tui
chub;
margined,
mottled,
or
piute
sculpin;
longnose
or
speckled
dace,
sucker,
and
northern
pikeminnow.
These
fish
are
known
to
exist
in
some
of
Washington's
warmest
waters,
where
they
often
out­
compete
introduced
populations
of
rainbow
trout.

Insufficient
research
information
exists
on
these
species
to
allow
a
similar
analysis
as
was
provided
previously
for
the
state's
cold
water
species.
The
following
conclusions
of
authors
who
have
studied
the
thermal
habitat
and
tolerances
of
Washington's
indigenous
warm
water
species
are
provided
to
support
establishing
appropriately
protective
criteria:

Redside
Shiner
(
Richardsonius
balteatus)

Found
in
streams
and
lakes
throughout
Washington,
redside
shiners
are
found
in
ponds,
lakes
and
irrigation
ditches
with
summer
water
temperatures
of
about
12.8
to
20
°
C.
However,
they
have
been
found
at
water
temperatures
as
cold
as
6.7
°
C
and
as
high
as
23.9
°
C.
The
fish
move
about
in
schools
and
tend
to
stay
in
vegetation
when
in
shallow
areas.
They
move
to
near­
shore
areas
in
the
spring
and
remain
there
until
July,
when
they
move
to
the
deep
water
zone;
by
August,
when
the
surface
waters
become
warmer
they
descend
into
deeper
water.
During
September
and
October
some
movement
from
deep
to
shallow
water
occurs
as
the
near­
shore
temperatures
decrease.
Between
October
and
May
the
shiner
probably
stay
in
deep
water.
Spawning
takes
place
in
the
spring
and
early
summer
(
April
­
July).
Movement
to
the
inlet
and
outlet
streams
in
British
Columbia
occurs
when
the
temperatures
first
exceed
10
°
C.
Spawning
occurs
at
night
over
the
gravel
bottom
of
streams
or
in
vegetation
along
the
lake
shoreline
(
Wydoski
and
Whitney,
1979).

Reeves
et
al.
(
1987)
used
laboratory
streams
to
determine
that
the
production
of
redside
shiner
increased
by
30%
when
moved
from
a
temperature
regime
of
12­
15
°
C
to
one
fluctuating
between
19­
22
°
C.
Black
(
1953)
measured
the
upper
lethal
temperature
values
for
fish
captured
from
lakes
in
southern
Okanogan
Valley
of
British
Columbia.
The
upper
temperature
(
°
C)
at
which
50%
of
the
fish
died
in
24
hours
was
estimated
for
redside
Shiner
(
Richardsonius
balteatus)
as
25
°
C
(
9­
11
°
C),
27.6
°
C
(
14
°
C)
(
with
the
approximate
acclimation
temperature
being
given
in
brackets).

Chiselmouth
(
Acrocheilus
alutaceus)

In
Washington
this
species
is
found
in
the
upper
Columbia
River
(
east
of
the
Cascade
Mountains)
and
its
tributaries.
It
inhabits
both
streams
and
lakes.
It
prefers
the
warmer
areas
of
streams
in
moderately
fast
to
fast
water.
Although
chiselmouth
are
also
found
in
lakes,
they
migrate
into
tributary
streams
to
spawn.
Spawning
occurs
in
late
June
and
early
July
in
British
Columbia
when
water
temperatures
exceed
17
°
C.
This
species
may
serve
as
a
major
Page
134
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
link
in
the
food
chain
from
the
primary
production
of
plants
to
piscivorous
fish
(
Wydoski
and
Whitney,
1979).

Longnose
Dace
(
Rhinichthys
cataractae)

Found
throughout
Washington,
this
dace
inhabits
the
swift­
running
water
of
streams.
it
prefers
summer
temperatures
of
12.8­
21.1
°
C.
Spawning
occurs
in
the
late
spring
or
early
summer
(
June
and
early
July)
on
gravel
bottoms
of
shallow
riffles
when
the
water
temperature
reaches
11.7
°
C.
Fry
hatch
in
7­
10
days
at
about
15.6
°
C
(
Wydoski
and
Whitney,
1979).
Stauffer
et
al.
(
1976)
studied
the
distribution
of
longnose
dace
(
Rhinicthys
cataractae).
They
found
14
specimens
at
water
temperatures
of
20.6­
26.7
°
C
one
year,
and
32
specimens
at
19.4­
30
°
C
the
following
year.

Northern
Pikeminnow
(
Ptychocheilus
oregonensis)

In
Washington
it
is
found
in
the
Columbia
River
system
and
coastal
and
Puget
Sound
drainages.
Northern
pikeminnow
inhabit
lakes
and
areas
of
slow
to
moderate
currents
in
streams.
They
prefer
the
highest
temperatures
(
20­
22.8
°
C)
that
occur
in
Lake
Washington.
Spawning
occurs
from
late
May
through
July,
with
a
peak
in
Washington
of
early
July.
The
eggs
hatch
in
7
days
at
18.3
°
C
and
the
young
become
free
swimming
in
14
days
(
Wydoski
and
Whitney,
1979).
Bell
(
1986)
in
a
review
on
the
temperature
requirements
of
fish
stated
that
the
range
for
northern
pikeminnow
was
from
16.1­
24.4
°
C.
The
hatching
temperature
is
around
18.3
°
C.
Black
(
1953)
measured
the
upper
lethal
temperature
values
for
fish
captured
from
lakes
in
southern
Okanogan
Valley
of
British
Columbia.
The
upper
temperature
at
which
50%
of
the
fish
died
in
24
hours
was
estimated
as
29.3
°
C
(
with
the
approximate
acclimation
temperature
being
19­
22
°
C).
Based
on
field
observations,
northern
pikeminnow
have
also
been
reported
to
prefer
temperatures
of
about
16­
22
º
C
(
Dimick
and
Merryfield
1945;
as
cited
in
Vigg
and
Burley,
1991).

Tui
Chub
(
Gila
bicolor)

In
Washington
this
species
is
found
primarily
in
the
central
part
of
the
state,
east
of
the
Columbia
River.
Tui
chub
migrate
to
shallow
water
in
spring,
but
stay
in
the
deeper
water
during
the
winter.
Tui
chub
first
spawn
in
their
third
year
of
life.
Spawning
occurs
in
May
and
June
(
when
water
temperatures
are
between
12.8­
15.6
°
C)
in
areas
with
many
aquatic
plants.
The
eggs
hatch
in
about
2
weeks;
the
larvae
are
well
developed
at
hatching
and
the
yolk
sac
has
already
been
absorbed.
Tui
chub
often
overpopulate
and
compete
with
trout
(
Wydoski
and
Whitney,
1979).

Peamouth
(
Mylocheilus
caurinus)

Peamouth
occur
in
much
of
the
Columbia
River
system
and
are
also
found
in
the
Coastal
and
Puget
Sound
provinces.
It
is
tolerant
of
salt
water.
In
Lake
Washington,
young
peamouth
inhabit
very
shallow
water
in
spring,
summer,
and
fall;
and
deep
water
during
the
winter.
Peamouth
in
Lake
Washington
grew
fastest
when
the
lake
was
most
eutrophic,
but
their
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
135
Washington's
Surface
Water
quality
Standards
growth
declined
after
the
sewage
was
completely
diverted
from
the
lake.
Spawning
occurs
in
late
May
and
early
June
in
streams
and
along
lake
shores
on
a
gravel
and
rubble
bottom
when
water
temperatures
warm
to
about
12.2
°
C.
Eggs
hatch
in
7­
8
days
at
12.2
°
C
(
Wydoski
and
Whitney,
1979).
Black
(
1953)
measured
the
upper
lethal
temperature
values
for
fish
captured
from
lakes
in
southern
Okanogan
Valley
of
British
Columbia.
The
upper
temperature
at
which
50%
of
the
fish
died
in
24
hours
was
estimated
as
for
Peamouth
(
Mylocheilus
caurinus)
as
27
°
C
(
10
°
C)
and
27.1
°
C
(
14
°
C)
(
with
the
approximate
acclimation
temperature
being
given
in
brackets).

Leopard
Dace
(
Rhinichthys
falcatus)

In
Washington
it
is
found
in
the
upper
Columbia
River
and
in
the
Similkameen
River,
which
flows
into
the
Okanogan
and
then
into
the
Columbia.
Leopard
dace
usually
inhabit
slower
and
deeper
water
than
that
preferred
by
long­
nose
dace.
Spawning
probably
occurs
in
July
and
August
(
Wydoski
and
Whitney,
1979).
Black
(
1953)
measured
the
upper
lethal
temperature
values
for
fish
captured
from
lakes
in
southern
Okanogan
Valley
of
British
Columbia.
The
upper
temperature
(
°
C)
at
which
50%
of
the
fish
died
in
24
hours
was
estimated
for
leopard
dace
(
Rhinichthys
falcatus)
as
28.3
°
C
(
with
the
approximate
acclimation
temperature
being
14
°
C).

Speckled
Dace
(
Rhinichthys
osculus)

It
is
common
throughout
Washington.
This
species
inhabits
the
colder
waters
of
streams
with
currents
ranging
from
slow
to
swift,
and
sometimes
is
found
in
lakes.
Spawning
occurs
from
June
through
August,
peaking
in
late
June.
Eggs
hatch
in
6
days
at
17.8
to
18.9
°
C
(
Wydoski
and
Whitney,
1979).
Li
et
al.
(
1993)
studied
stream
enhancement
works
in
the
John
Day
Basin
of
Oregon.
The
authors
noted
that
speckled
dace
continued
to
feed
when
temperatures
reached
as
high
as
33
°
C.
Baltz,
Moyle,
and
Knight
(
1982)
studied
the
distribution
of
riffle
sculpin
(
Cottus
gulosus)
and
speckled
dace
(
Rhinichthys
osculus)
in
a
California
creek.
While
both
species
occupied
the
same
physical
habitat
types
in
the
stream,
riffle
sculpin
were
confined
to
the
upper
reaches
of
the
creek.
In
testing
the
swimming
performance,
metabolic
rates,
and
competitive
interactions
between
these
two
species
they
concluded
that
competitive
interactions
between
dace
and
sculpin
for
preferred
microhabitat
were
mediated
by
temperature
with
sculpin
showing
dominance
in
the
cooler
water
and
dace
showing
dominance
in
warmer
water.
The
highest
daily
maximum
temperatures
in
the
upper
portion
of
the
creek
was
29
°
C
(
though
the
data
would
suggest
that
daily
maximum
temperatures
are
more
commonly
below
24­
26
°
C)
and
in
the
lower
portion
of
the
creek
reached
32
°
C
(
temperatures
approaching
32
°
C
appears
to
be
common
in
the
lower
portion
of
the
stream).
In
studying
prey
selection
the
authors
found
that
high
proportion
of
the
dace
had
empty
stomachs.
The
swimming
performance
of
dace
increased
significantly
with
temperature,
but
performances
at
15
and
20
°
C
were
not
significantly
different
thus
poor
swimming
performance
at
cooler
summer
temperatures
probably
was
not
a
factor
contributing
to
dace
displacement
in
upstream
riffles.

Largescale
Sucker
(
Catostomus
macrocheilus)
Page
136
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Found
throughout
Washington
in
both
lakes
and
streams.
Spawning
occurs
during
April
or
May
(
Wydoski
and
Whitney,
1979).
Black
(
1953)
measured
the
upper
lethal
temperature
values
for
fish
captured
from
lakes
in
southern
Okanogan
Valley
of
British
Columbia.
The
upper
temperature
at
which
50%
of
the
fish
died
in
24
hours
was
estimated
for
large
scale
sucker
(
Catostomus
macrocheilus)
as
29.4
°
C
(
with
the
approximate
acclimation
temperature
being
19
°
C).

Piute
Sculpin
(
Cottus
beldingi)

In
Washington
this
sculpin
is
found
east
of
the
Cascade
Mountains,
in
the
Columbia,
Yakima,
Snake,
and
Walla
Walla
rivers
and
their
tributaries.
It
inhabits
streams
that
have
a
slight
to
moderate
gradient,
and
is
found
in
riffle
areas
among
rubble
and
large
gravel.
It
often
frequents
water
warmer
than
15
°
C
and
has
been
found
at
25
°
C.
Usually
it
is
found
in
the
lowlands
but
has
been
collected
as
high
as
4,000
feet
in
Oregon.
Piute
sculpin
in
Lake
Tahoe
spawn
in
May
and
June
at
temperatures
of
about
12.2
°
C
(
Wydoski
and
Whitney,
1979).

Margined
Sculpin
(
Cottus
marginatus)

In
Washington
this
species
is
found
in
the
Walla
Walla,
Touchet,
and
Tucannon
rivers.
It
has
been
collected
in
streams
with
temperatures
of
12.8­
18.9
°
C,
but
have
also
been
found
as
high
as
23.9
°
C.
In
Oregon
it
is
associated
with
rainbow
trout,
speckled
dace,
longnose
dace,
and
the
Piute
sculpin.
It
may
have
a
similar
life
history
as
the
reticulate
sculpin
(
Wydoski
and
Whitney,
1979).

Summary
on
Temperature
Requirements
of
Indigenous
Warm
Water
Species:

Insufficient
information
exists
to
develop
individual
water
quality
thresholds
for
each
of
these
native
warmwater
species,
so
it
is
recommended
that
they
be
considered
broadly
as
a
community.
The
lack
of
available
experiential
data
for
which
to
base
sound
recommendations
on
combined
with
the
conflicts
that
are
presented
in
what
is
available,
strongly
suggest
that
these
recommendations
may
need
to
be
revisited
periodically
to
determine
if
changes
are
warranted.
Establishing
criteria
to
protect
our
temperature
tolerant
non­
salmonid
fish
species
will
also
provide
protection
for
desirable
introduced
warm
water
sport
fish
species
such
as
bass
and
crappie.
In
general,
Washington's
indigenous
warm
water
fish
communities
thrive
in
waters
that
have
summer
maximum
temperatures
as
high
as
25­
27
°
C,
although
most
seem
to
prefer
waters
below
18­
20
°
C.

Warm
Water
Habitat
Recommendation:
It
is
recommended
that
in
waters
supporting
communities
of
indigenous
warm
water
fish,
the
highest
moving
7­
day
average
of
the
daily
maximum
temperatures
should
not
exceed
20
°
C.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
137
Washington's
Surface
Water
quality
Standards
To
avoid
conflicts
with
the
thermal
requirements
of
cold
water
fish,
it
should
probably
be
explicitly
specified
in
the
regulation
that:
"
The
Warm
Water
Aquatic
Life
category
may
only
be
applied
to
waters
that
do
not
have
self­
reproducing
populations
of,
or
serve
as
migration
corridors
for,
indigenous
salmonids
or
char.
It
is
appropriate
only
where
the
dominant
species
under
natural
conditions
would
be
temperature
tolerant
indigenous
non­
salmonid
species
such
as
dace,
redside
shiner,
chiselmouth,
sucker,
and
northern
pikeminnow;
and
which
may
also
be
serving
as
habitat
for
introduced
warm
water
tolerant
sport­
fish
species
such
as
bass
and
crappie."
Page
138
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
5.
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Aquatic
Life
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Washington's
Surface
Water
quality
Standards
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
175
Washington's
Surface
Water
quality
Standards
Part
III
Ambient
Temperatures
of
Washington's
Streams
and
Rivers
and
Proposal
for
Protecting
the
Spawning
and
Early
Tributary
Rearing
of
Char
Prepared
by:
Andrew
Kolosseus
Water
Quality
Standards
Washington
Department
of
Ecology
(
360)
407­
7543
akol461@
ecy.
wa.
gov
Page
176
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
177
Washington's
Surface
Water
quality
Standards
Ambient
Temperatures
of
Washington's
Streams
Introduction
As
described
in
Part
two
of
this
document,
salmonids
require
colder
temperatures
during
spawning
and
incubation
than
during
rearing.
Spawning
and
incubation
usually
occur
in
the
fall,
winter,
and
spring.
One
of
the
main
decisions
when
establishing
temperature
criteria
is
whether
to
apply
two
criteria
(
one
during
rearing
and
a
colder
one
during
spawning/
incubation)
or
only
one
criterion
that
is
more
reflective
of
rearing
and
relies
on
natural
seasonal
cooling
to
protect
spawning
and
incubation.

In
other
words,
can
a
single
criterion
be
developed
that
will
protect
rearing
and
spawning/
incubation?
This
section
describes
the
physical
characteristics
of
streams
in
Washington
in
order
to
help
make
that
decision.

Comparison
of
Summer
Rearing
Temperatures
and
Spawning/
Incubation
Temperatures
The
crux
of
whether
or
not
to
apply
separate
spawning/
incubation
criteria
is
a
comparison
of
summer
rearing
temperatures
and
the
spawning/
incubation
temperatures.
Before
a
single
rearing
criterion
can
be
used,
the
question
must
be
asked
"
Will
a
single
rearing
criterion
protect
spawning
and
incubation
when
it
occurs?"

In
order
to
answer
this
question,
Ecology
gathered
continuous
temperature
data
from
a
variety
of
sources.
Ecology
obtained
data
from
126
sites
over
various
years
(
a
large
percentage
of
the
data
was
from
2000
and
2001).
The
data
came
from
a
variety
of
sources
(
Ecology,
WDFW,
USFS,
and
USGS)
and
a
variety
of
streams
from
across
the
state.
Given
the
relatively
small
sample
size,
Ecology
made
no
attempt
to
make
the
data
representative
with
respect
to
the
year
the
monitoring
occurred,
elevation,
geography,
stream
temperature,
stream
size,
stream
type,
or
any
other
factor.
Although
the
sites
do
not
proportionately
represent
water
bodies
in
Washington,
they
do
provide
a
broad
sample
of
water
body
types.

Ecology
used
the
WDFW
Salmonid
Stock
Inventory
(
SaSI)
to
determine
when
spawning
occurred.
Sites
with
no
temperature
data
during
the
spawning/
incubation
period
were
excluded.

The
following
chart
shows
how
streams
with
different
summer
maximum
7­
DADMax
temperatures
cooled
down
by
the
time
spawning
began.
The
multiple
lines
of
evidence
approach
(
described
in
Part
2
of
this
document)
showed
that
the
water
temperature
at
spawning
should
be
less
than
12.5­
14
º
C
(
7­
DADM).
During
non­
spawning
and
non­
incubating
times,
the
temperature
should
be
less
than
16­
17.5
º
C
(
7­
DADM).
These
temperatures
fully
protect
salmonids.
Page
178
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
Streams
that
Met
a
Spawning
Criteria
of
12.5,
13,
13.5,
and
14
º
C
(
7­
DADM)

55%
64%
82%
100%

0%
10%
20%
30%
40%
50%
60%
70%
80%
90%

100%
<
13
º
13­
14
14­
15
15­
16
16­
17
17­
18
18­
19
19­
20
>
20
º
Summer
(
Rearing)
Maximum
7­
DADM
Temperature
Percentage
of
Streams
that
Meet
the
Spawning
Criteria
(

7­

DADM)
12.5
º
13
º
13.5
º
14
º
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
179
Washington's
Surface
Water
quality
Standards
Summer
(
Rearing)
Maximum
Temperature
Ranges
(
7­
DADMax
in
º
C)
Number
of
Sites
with
Data
<
13
º
12
13­
14
º
7
14­
15
º
7
15­
16
º
11
16­
17
º
20
17­
18
º
13
18­
19
º
5
19­
20
º
8
>
20
º
23
Looking
at
the
streams
with
a
summer
7­
DADMax
of
15­
16
º
C,
one
can
see
that
55%
of
those
streams
were
12.5
º
C
(
7­
DADMax)
or
less
by
the
time
spawning
occurred,
64%
were
13
º
C
or
less,
82%
were
13.5
º
C
or
less,
and
all
of
the
streams
were
14
º
C
or
less.
As
the
summer
temperatures
decreased,
a
higher
percentage
of
streams
reach
temperatures
of
12.5­
14
º
C
(
7­
DADMax)
during
spawning.
As
the
summer
temperatures
increased,
a
lower
percentage
of
streams
reached
temperatures
of
12.5­
14
º
C
(
7­
DADMax)
during
spawning.

For
streams
with
a
summer
7­
DADMax
of
16­
17
º
C,
one
can
see
that
only
15%
of
those
streams
were
12.5
º
C
(
7­
DADMax)
or
less
by
the
time
spawning
occurred,
30%
were
13
º
C
or
less,
40%
were
13.5
º
C
or
less,
and
55%
were
14
º
C
or
less.

These
data
show
that
a
single
criterion
of
16
º
C
7­
DADMax
would
have
protected
spawning
in
many,
but
not
all
streams.
Before
drawing
conclusions
about
the
protectiveness
of
a
single
criterion,
two
additional
factors
should
be
taken
into
account.
These
factors
are
inter­
annual
variability
(
how
the
maximum
temperature
of
a
water
body
varies
from
year
to
year)
and
spatial
variability
(
how
the
temperature
of
a
water
body
varies
as
it
flows
downstream).

Inter­
Annual
Variability
Most
of
the
data
just
described
was
from
one
or
just
a
few
years.
An
important
issue
to
address
when
setting
temperature
criteria
is
how
maximum
river
temperatures
fluctuate
from
year
to
year,
also
known
as
inter­
annual
variability.
Unfortunately,
there
is
very
little
continuous
temperature
data
over
a
long
period
of
time
(
i.
e.
ten
or
more
years).
It
is
not
known
how
much
of
the
inter­
annual
variability
in
the
historic
record
is
due
to
natural
conditions
(
i.
e.
climate
and
rainfall)
and
how
much
is
due
to
human
activity
(
i.
e.
canopy
shade
reduction).

To
illustrate
inter­
annual
variability,
consider
the
Cispus
River.
The
USGS
had
a
monitoring
station
on
the
Cispus
River
near
Randle,
Washington
from
1952­
1971.
The
Cispus
River
is
in
Lewis
and
Skamania
Counties
and
is
a
tributary
of
the
Cowlitz
River.
Other
rivers
have
Page
180
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
similar,
but
not
identical,
inter
annual
variability.
A
lack
of
data,
especially
on
the
east
side,
makes
a
more
robust,
statistical
look
at
inter­
annual
variability
impossible.
The
following
table
and
chart
show
how
the
maximum
7­
DADMax
river
temperature
varied
from
year
to
year
in
the
Cispus
River:

Year
Maximum
7­
DADMax
Temperature
(
º
C)
Year
Maximum
7­
DADMax
Temperature
(
º
C)
1952
15.6
1963
15.7
1953
14.2
1964
13.4
1954
12.8
1965
15.2
1955
incomplete
1966
14.0
1956
13.6
1967
15.0
1957
14.8
1968
16.0
1958
16.4
1969
14.0
1959
15.3
1970
14.9
1960
15.6
1971
13.5
1961
14.4
1972
incomplete
1962
15.2
Inter­
Annual
Variability
of
the
Cispus
River
10
º
C
11
º
C
12
º
C
13
º
C
14
º
C
15
º
C
16
º
C
17
º
C
18
º
C
1950
1955
1960
1965
1970
1975
Year
Maximum
7­
DADM
Temperature
While
during
the
hottest
years
the
Cispus
River
reached
16
º
C,
during
the
colder
years
the
river
typically
stayed
below
13­
14
º
C.

When
deciding
if
a
one
criterion
standard
is
protective
enough,
it
is
important
to
keep
interannual
variability
in
mind.
While
a
water
body
might
have
a
low
probability
of
protecting
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
181
Washington's
Surface
Water
quality
Standards
spawning/
incubation
during
warm
years,
during
colder
years
it
would
be
have
a
much
higher
probability
of
protecting
spawning/
incubation.

Spatial
Variability
In
general,
streams
warm
as
they
flow
downstream.
The
temperature
criteria
apply
throughout
the
entire
length
of
the
stream,
including
the
furthest
downstream
point.
This
means
that
in
order
to
meet
the
temperature
criteria
at
the
furthest
downstream
point,
upstream
areas
will
have
to
be
cooler
than
the
criterion.
How
much
a
stream
changes
temperature
as
it
flows
downstream
 
spatial
variability
 
is
quite
different
for
each
water
body
and
depends
on
the
characteristics
of
the
individual
water
body.

To
illustrate
spatial
variability,
consider
the
Chiwawa
River,
a
tributary
of
the
Wenatchee
River
in
Chelan
County.
On
August
12,
2001,
the
temperature
along
the
Chiwawa
River
was
measured.
The
temperatures
were
instantaneous
measurements,
not
7­
DADMax,
so
they
should
not
be
compared
with
the
proposed
criteria.
However,
they
are
still
useful
for
illustrating
the
effects
of
spatial
variability.
Other
rivers
have
different
spatial
variability,
but
the
Chiwawa
River
can
be
used
as
an
example.
A
lack
of
data
makes
a
more
robust,
statistical
look
at
spatial
variability
impossible.

The
following
chart
shows
the
river
temperature
generally
increasing
as
the
river
flows
downstream
from
the
headwaters
to
the
mouth.

Instaneous
Temperature
along
the
Chiwawa
River
on
August
12,
2001
10
12
14
16
18
20
22
0
2
4
6
8
10
12
14
16
18
20
22
24
26
28
30
32
34
Distance
from
mouth
(
miles)
Water
Temperature
(
º
C)

(
Headwaters)
(
Mouth)
Page
182
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
As
the
chart
shows,
the
river
stayed
between
14
º
C
and
16
º
C
for
many
miles.
Over
the
last
about
17
miles,
the
river
warmed
steadily.
While
some
of
this
warming
is
probably
due
to
human
influences,
some
of
it
is
also
naturally
occurring.
As
this
example
shows,
in
order
for
the
entire
river
to
meet
a
criterion,
most
of
the
river
would
be
cooler
than
the
criterion.

When
deciding
if
a
one
criterion
standard
is
protective
enough,
it
is
important
to
keep
spatial
variability
in
mind.
While
a
water
body
might
have
a
low
probability
of
protecting
spawning/
incubation
at
the
lowest
downstream
point,
it
would
have
a
higher
probability
of
protecting
spawning/
incubation
further
upstream.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
183
Washington's
Surface
Water
quality
Standards
Proposal
for
Protecting
the
Spawning
and
Early
Tributary
Rearing
of
Char
Introduction
The
existing
water
quality
standards
do
not
have
temperature
criteria
that
fully
protect
char.
This
document
describes
the
proposal
to
designate
certain
waters
as
char
habitat
for
the
purpose
of
applying
a
fully
protective
temperature
criterion.

The
goal
is
to
fully
protect
the
spawning
and
early
juvenile
rearing
of
char.
Ecology
investigated
three
methods
of
identifying
waters
used
by
char:

1.
Known
spawning
and
early
juvenile
rearing
streams.
Unfortunately,
there
is
no
comprehensive
survey
of
known
spawning
and
early
juvenile
rearing
areas
and
finding
spawning
and
rearing
areas
used
by
these
reclusive
fish
has
proved
to
be
very
difficult.
There
are
entire
populations
of
char
where
the
spawning
areas
are
completely
unknown.
Even
in
areas
of
extensive
study,
not
all
the
spawning
and
early
juvenile
rearing
areas
have
been
identified.
Using
this
method
to
identify
streams
for
protecting
spawning
and
early
juvenile
rearing
of
char
would
likely
result
in
many
streams
with
char
populations
not
being
protected
in
the
water
quality
standards
until
they
were
identified.
2.
Entire
watersheds
where
char
are
present.
Every
water
body
in
the
entire
watershed
that
is
accessible
to
char
would
be
protected,
regardless
of
its
likelihood
of
being
a
spawning
or
early
tributary
rearing
water
body.
Even
lower
main
stem
rivers
with
their
very
warm
temperatures
and
low
likelihood
of
providing
suitable
habitat
would
have
to
meet
very
stringent
temperature
requirements.
3.
All
stream
segments
in
watersheds
used
by
char
that
have
the
basic
physical
characteristics
of
known
char
spawning
and
early
tributary
rearing
streams.
This
option
avoids
applying
the
temperature
criteria
to
streams
that
would
not
likely
be
used
by
char,
but
also
does
not
depend
on
actually
proving
in
advance
that
char
are
using
each
of
the
qualifying
streams.
The
physical
characteristics
that
were
found
to
best
define
char
spawning
and
early
tributary
rearing
waters
are
addressed
below.

Data
The
Washington
Department
of
Fish
and
Wildlife
(
WDFW)
has
compiled
information
on
bull
trout
habitat.
They
recently
released
a
database
that
identifies
known
spawning
areas.
This
data
set
combines
the
knowledge
from
biologists
working
for
WDFW,
USFWS,
Tribes,
and
others.
The
data
are
current
as
of
November,
2001.
There
are
no
databases
of
known
early
tributary
rearing
areas.
However,
based
on
what
is
known
of
the
biology
of
char,
they
would
most
typically
be
in
the
same
general
locations
as
the
known
spawning
areas.
It
is
important
to
note
that
where
site
knowledge
demonstrates
that
this
early
tributary
rearing
Page
184
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
occurs
at
locations
away
from
the
spawning
grounds,
then
the
proposed
methodology
will
allow
these
areas
to
be
protected
in
future
rulemakings.

This
database
(
often
referred
to
as
the
"
bullchar"
database)
forms
the
foundation
of
Ecology's
proposal
for
protecting
char.

Development
of
a
Filter
Ecology
studied
the
locations
of
known
spawning
areas
documented
in
the
"
bullchar"
database
and
found
that
their
occurrence
is
largely
restricted
to
a
relatively
narrow
range
of
elevation
and
stream
order.
Ecology
used
this
pattern
of
elevation
and
stream
order
to
deduce
which
streams
would
reasonably
be
expected
to
be
potential
char
habitat.

Stream
Order
The
stream
order
concept
(
Strahler,
1952)
is
a
method
of
classifying
streams.
Headwater
streams
are
assigned
a
stream
order
of
1.
When
two
1st
order
streams
join,
they
form
a
2nd
order
stream.
When
two
2nd
order
streams
join,
they
form
a
3rd
order
stream,
and
so
on.
When
a
lower
order
stream
joins
and
higher
order
stream
(
for
example,
a
1st
order
stream
joins
a
3rd
order
stream),
the
stream
order
does
not
change.
1
Ecology
found
that
most
known
spawning
areas
were
in
1st,
2nd,
and
3rd
order
streams.
Limited
spawning
occurred
in
4th
order
streams.
The
following
table
shows
the
stream
orders
of
the
known
spawning
streams:

Known
Spawning
Streams:

Stream
Order
East
Side
West
Side
Combined
1
18%
24%
21%
2
36%
36%
36%
3
35%
35%
35%
4
10%
5%
8%
5
1%
0%
0%

As
the
table
shows,
there
is
little
difference
between
east
side
streams
and
west
side
streams.
Approximately
92%
of
all
known
spawning
occurs
in
1st,
2nd,
and
3rd
order
streams,
so
1
The
original
stream
order
system
assigns
the
upper­
most
perennial
streams
to
a
stream
order
of
1.
However,
the
WDFW
bull
trout
data,
and
other
commonly
used
data,
are
at
a
scale
where
not
all
perennial
streams
are
identified.
These
data
are
from
the
Washington
Hydrography
Framework
Layer,
and
are
at
the
1:
100,000
scale.
Given
this
limitation,
the
upper­
most
streams
identified
in
the
data
is
assigned
a
stream
order
of
1.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
185
Washington's
Surface
Water
quality
Standards
Ecology
focused
its
efforts
on
those
streams
when
developing
the
system
for
identifying
char
waters.

Elevation
An
analysis
of
all
the
spawning
data
quickly
shows
that
known
spawning
areas
are
concentrated
in
higher
elevation
streams.
For
each
known
spawning
stream,
the
lowest
elevation
was
calculated.
2
The
following
table
provides
summary
information
of
known
spawning
streams
and
their
elevations:

Elevations
(
in
feet)
of
Known
Spawning
Streams
East
Side
West
Side
Number
of
Streams
77
67
Average
Elevation
3136
1395
Maximum
Elevation
4650
3320
Minimum
Elevation
1419
420
Lower
95th
Percentile
1889
676
This
analysis
found
that
94%
of
the
known
spawning
areas
were
above
2000
feet
on
the
east
side
and
above
700
feet
on
the
west
side.
Thus
this
elevation
filter
captures
most
of
the
known
spawning
streams.

System
for
Identifying
Char
Waters
Using
the
information
about
the
stream
order
and
elevation
of
known
spawning
streams,
Ecology
developed
the
following
proposed
system
for
determining
which
water
bodies
should
be
protected
for
char.
This
system
is
used
in
all
of
the
watersheds
with
known
spawning
areas
or
with
suspected
spawning
populations
identified
by
the
USFW
1999
"
Washington
Distinct
Population
Segment
 
Bull
Trout
Subpopulation"
map.

1.
All
known
char
spawning
streams
will
be
protected
The
"
known
char
spawning
streams"
are
those
streams
identified
by
WDFW
in
either
the
"
bullchar"
database
or
in
WDFW's
1998
Salmonid
Stock
Inventory
(
SaSI).
If
other
streams
are
identified
as
known
spawning
streams
during
this
rule­
making
process,
they
will
also
be
included.

2.
All
streams
upstream
of
known
char
spawning
streams
will
be
protected
These
are
the
streams
upstream
of
those
locations
identified
in
(
1)
above.

2
Throughout
this
analysis,
the
lowest
elevation
of
the
stream
or
stream
segment
was
used.
Both
the
known
spawning
streams
and
the
application
of
the
elevation
filter
used
the
lowest
elevation
to
maintain
consistency
and
reduce
any
bias.
Page
186
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
3.
All
3rd
order
streams
and
their
tributaries
will
be
protected
if
they
join
a
4th
order
stream
As
discussed
earlier,
bull
trout
spawning
areas
are
concentrated
in
1st­
3rd
order
streams.
This
part
of
the
system
includes
certain
3rd
order
streams
and
their
tributaries
(
i.
e.
the
1st
and
2nd
order
streams).
It
does
not
include
the
lower
elevation
3rd
order
streams
that
join
a
5th
or
higher
order
river.
The
known
spawning
areas
generally
did
not
include
these
lower
elevation
3rd
order
streams,
so
they
were
not
included
in
the
filter.

Some
1st
and
2nd
order
streams
are
not
protected
by
this
filter.
These
include
1st
and
2nd
order
streams
that
directly
flow
into
a
4th
or
higher
order
streams,
except
at
outlined
below
in
(
4).
The
known
spawning
areas
generally
did
not
include
these
1st
and
2nd
order
streams
that
flow
into
4th
or
higher
order
streams,
so
they
were
not
included
in
the
filter.

4.
All
2nd
order
streams
and
their
tributaries
will
be
protected
if
they
join
a
4th
order
stream
and
they
are
above
a
stream
protected
by
sections
(
1),
(
2),
or
(
3).
This
part
of
the
filter
captures
the
higher
elevation
2nd
and
1st
order
streams
that
are
used
for
char
spawning.

Elevation
Exception:
The
default
system
described
in
(
3)
and
(
4)
above
is
not
applied
to
streams
below
2000
feet
on
the
east
side
of
the
Cascades
or
below
700
feet
on
the
west
side
of
the
Cascades.

Known
spawning
locations
and
all
streams
upstream
of
known
spawning
locations
will
be
protected
regardless
of
their
elevation.
(
In
other
words,
elevation
can
prevent
a
stream
from
being
covered
under
(
3)
and
(
4),
but
not
under
(
1)
or
(
2)).
This
part
of
the
system
reflects
the
fact
that
about
94%
of
known
spawning
streams
are
above
2000
feet
on
the
east
side
and
700
feet
on
the
west
side.

Protected
Streams
Ecology's
proposal
to
apply
the
proposed
system
to
designate
char
waters
results
in
the
protection
of
the
dark
green
streams
in
the
following
map.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
187
Washington's
Surface
Water
quality
Standards
For
more
detailed
maps,
please
visit
Ecology's
website
at
www.
ecy.
wa.
gov/
programs/
wq/
swqs/
bull_
trout.
Copies
of
the
GIS
map
files
that
are
viewable
for
users
with
ArcView
are
available
upon
request
from
Ecology.

Alternative
Proposal
for
Protecting
Char
An
alternative
proposal
for
protecting
char
is
also
being
considered.
Using
the
same
information
about
the
stream
order
and
elevation
of
known
spawning
streams,
the
alternative
proposal
would
protect
char
according
to
the
following
system:

1.
All
known
char
spawning
streams
will
be
protected
2.
All
streams
upstream
of
known
char
spawning
streams
will
be
protected
3.
All
3rd
order
streams
and
their
tributaries
will
be
protected
IF
the
3rd
order
stream
is
above
700
feet
elevation
(
west
side
of
the
Cascades)
or
2000
feet
elevation
(
east
side
of
the
cascades).
If
only
part
of
the
3rd
order
stream
is
above
the
specified
elevation,
only
that
part
of
the
stream
(
and
all
its
tributaries)
will
be
protected.

This
system,
while
similar
to
Ecology's
proposal
detailed
above,
results
in
more
streams
being
protected
for
char.
Page
188
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Washington's
Surface
Water
quality
Standards
The
following
map
shows
the
streams
that
would
be
protected
by
this
alternative.
The
protected
streams
are
in
dark
green.

The
following
map
compares
the
water
bodies
protected
by
Ecology's
primary
proposal
and
the
water
bodies
protected
by
the
alternative
proposal.
The
streams
that
would
be
protected
by
both
proposals
are
in
dark
green,
the
streams
protect
only
by
Ecology's
primary
proposal
are
in
pink,
and
the
streams
protected
only
by
the
alternative
proposal
are
in
gold.
Evaluating
Standards
for
Protecting
Aquatic
Life
in
Page
189
Washington's
Surface
Water
quality
Standards
For
more
detailed
maps,
please
visit
Ecology's
website
at
www.
ecy.
wa.
gov/
programs/
wq/
swqs/
bull_
trout.
Copies
of
the
GIS
map
files
that
are
viewable
for
users
with
ArcView
are
available
upon
request
from
Ecology.