__

.
b!

QL
155
.
S63
no.
8
Biological
Report
82(
11
.126)
December
1989
TR
EL­
82­
4
Species
Profiles:
Life
Histories
and
Environmental
Requirements
of
Coastal
Fishes
and
Invertebrates
(
Pacific
Northwest)

PACIFIC
HERRING
`
2­
11.126
z
Coastal
Ecology
Group
Fish
and
Wildlife
Service
Waterwavs
ExDeriment
Station
U.
S.
Department
of
the
Interior
U.
S.
Army
Corps
of
Engineers
c
Biological
Report
82(
11.126)
TR
EL­
82­
4
December
1989
Species
Profiles:
Life
Histories
and
Environmental
Requirements
of
Coastal
Fishes
and
Invertebrates
(
Pacific
Northwest)

PACIFIC
HERRING
bY
Dennis
R.
Lassuy
Oregon
Cooperative
Fishery
Research
Unit
Department
of
Fisheries
and
Wildlife
Oregon
State
University
Corvallis,
OR
9733
l­
3803
Project
Officer
David
Moran
U.
S.
Fish
and
Wildlife
Service
National
Wetlands
Research
Center
1010
Gause
Boulevard
Slidell,
LA
70458
Performed
for
U.
S.
Army
Corps
of
Engineers
Coastal
Ecology
Group
Waterways
Experiment
Station
Vicksburg,
MS
39180
and
U.
S.
Department
of
the
Interior
Fish
and
Wildlife
Service
Research
and
Development
National
Wetlands
Research
Center
Washington,
DC
20240
This
series
should
be
referenced
as
follows:

U.
S.
Fish
and
Wildlife
Service.
1983­
19
.
Species
profiles:
life
histories
and
environmental
requirements
of
coastal
fishes
and
invertebrates.
U.
S.
Fish
Wildl.
Serv.
Biol.
Rep.
82(
11).
U.
S.
Army
Corps
of
Engineers,
TR
EL­
82­
4.

This
profile
should
be
cited
as
follows:

Lassuy,
D.
R.
1989.
Species
profiles:
life
histories
and
environmental
requirements
of
coastal
fishes
and
invertebrates
(
Pacific
Northwest)­­
Pacific
herring.
U.
S.
Fish
Wildl.
Serv.
Biol.
Rep.
82(
11.126).
U.
S.
Army
Corps
of
Engineers,
TR­
EL­
82­
4.
18
pp.
3
PREFACE
This
species
profile
is
one
of
a
series
on
coastal
aquatic
organisms,
principally
fish,
of
sport,
commercial,
or
ecological
importance.
The
profiles
are
designed
to
provide
coastal
managers,
engineers,
and
biologists
with
a
brief
comprehensive
sketch
of
the
biological
characteristics
and
environmental
requirements
of
the
species
and
to
describe
how
populations
of
the
species
may
be
expected
to
react
to
environmental
changes
caused
by
coastal
development.
Each
profile
has
sections
on
taxonomy,
life
history,
ecological
role,
environmental
requirements,
and
economic
importance,
if
applicable.
A
three­
ring
binder
is
used
for
this
series
so
that
new
profiles
can
be
added
as
they
are
prepared.
This
project
is
jointly
planned
and
financed
by
the
U.
S.
Army
Corps
of
Engineers
and
the
U.
S.
Fish
and
Wildlife
Setvice.

Suggestions
or
questions
regarding
this
report
should
be
directed
to
one
of
the
following
addresses.

Information
Transfer
Specialist
U.
S.
Fish
and
Wildlife
Service
National
Wetlands
Research
Center
NASA­
Slidell
Computer
Complex
1010
Gause
Boulevard
Slide&
LA
70458
or
U.
S.
Army
Engineer
Waterways
Experiment
Station
Attention:
WESER­
C
Post
Office
Box
63
1
Vicksburg,
MS
39180
.
.
.
111
CONVERSION
TABLE
Metric
to
U.
S.
Customary
Multiply
millimeters
(
mm)
centimeters
(
cm)
meters
(
m)
meters
kilometers
(
km)
kilometers
BY
0.03937
0.3937
3.281
0.5468
0.6214
0.5396
To
Obtain
inches
inches
feet
fathoms
statute
miles
nautical
miles
square
meters
(
m2)
10.76
square
kilometers
(
km2)
square
feet
0.3861
square
miles
hectares
(
ha)
2.471
acres
liters
(
1)
0.2642
cubic
meters
(
m3)
gallons
35.31
cubic
feet
cubic
meters
0.0008110
acre­
feet
milligrams
(
mg)
0.00003527
ounces
grams
(
8)
0.03527
ounces
kilograms
(
kg)
2.205
pounds
metric
tons
(
t)
2205.0
pounds
metric
tons
1.102
short
tons
kilocalories
(
kcal)
Celsius
degrees
("
C)
3.968
1.8
("
C)
+
32
U.
S.
Customary
to
Metric
25.40
2.54
0.3048
1.829
1.609
1.852
British
thermal
units
Fahrenheit
degrees
inches
inches
feet
(
ft)
fathoms
statute
miles
(
mi)
nautical
miles
(
nmi)

square
feet
(
ft2)
square
miles
(
mi2)
acres
millimeters
centimeters
meters
meters
kilometers
kilometers
0.0929
square
meters
2.590
square
kilometers
0.4047
hectares
gallons
(
gal)
3.785
liters
cubic
feet
(
ft3)
0.0283
1
cubic
meters
acre­
feet
1233.0
cubic
meters
ounces
(
oz)
28350.0
milligrams
ounces
28.35
grams
pounds
(
lb)
0.4536
kilograms
pounds
0.00045
metric
tons
short
tons
(
ton)
0.9072
metric
tons
British
thermal
units
(
Btu)
0.2520
kilocalories
Fahrenheit
degrees
("
F)
0.5556
("
F
­
32)
Celsius
degrees
iv
&
CONTENTS
PREFACE
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**­
111
CONVERSION
TABLE
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iv
ACKNOWLEDGMENTS
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vi
NOMENCLATURE/
TAXONOMY/
RANGE
...............................
MORPHOLOGY/
IDENTIFICATION
AIDS
...............................
REASON
FOR
INCLUSION
IN
SERIES
.................................
LIFE
HISTORY
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Spawning
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Eggs
and
Larvae
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Juveniles
and
Adults
.............................................
GSI
and
Fecundity
................................................
GROWTH
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THE
FISHERY
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History
and
Products
..............................................
stocks
........................................................
Population
Dynamics
and
Management
..................................
ECOLOGICAL
ROLE
..............................................
Feeding
Habits
...................................................
Sources
of
Mortality
...............................................
ENVIRONMENTAL
REQUIREMENTS
.................................
Salinity
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Temperature
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Temperature
and
Salinity
Interactions
..................................
Substrate
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~
n~~~
inan;
s
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C
O
N
C
E
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N
S
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112
222
446
6
689
10
10
11
12
12
12
12
13
13
13
13
LITERATURE
CITED
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15
A
t
V
ACKNOWLEDGMENTS
Any
review
of
the
Pacific
herring
must
include
a
large
dose
of
gratitude
to
the
years
of
excellent
work
carried
out
by
many
members
of
the
staff
of
the
Pacific
Biological
Station
in
Nanaimo,
British
Columbia.
The
organizers
and
editors
of,
and
contributors
to,
the
Canadian
Journal
of
Fisheries
and
Aquatic
Sciences
(
Vol.
42,
Suppl.
1
of
1985)
that
covered
the
International
Symposium
on
the
Biological
Characteristics
of
Herring
and
Their
Implications
for
Management
are
also
owed
many
thanks.
I
am
grateful
to
Tom
Jow
(
California
Department
of
Fish
and
Game),
Jerry
Butler
(
Oregon
Department
of
Fish
and
Wildlife),
and
Pat
McAllister
and
Dwane
Day
(
Washington
Department
of
Fisheries)
for
supplying
information.
Jerome
Spratt
(
California
Department
of
Fish
and
Game)
and
Dwane
Day
and
Daniel
Pentilla
(
Washington
Department
of
Fisheries)
reviewed
the
manuscript.
Finally,
I
thank
Adrian
Hunter
for
her
dedicated
and
patient
help
in
preparing
the
manuscript.

vi
Figure
1.
Pacific
herring
(
from
Hart
1973).

PACIFIC
HERRING
NOMENCLATURE/
TAXONOMY/
RANGE
Scientific
name
.........
&
pea
harengus
paZZasi
(
Valenciennes
1847)
Common
name
............
Pacific
herring
Class
....................
Osteichthyes
Order
...................
Clupeiformes
Family
.....................
Clupeidae
Geographic
range:
Geographic
distribution
of
this
subspecies
extends
from
northern
Baja
California
well
into
arctic
Alaska
and
the
U.
S.
S.
R.,
Japan,
and
the
Yellow
Sea.
It
is
commercially
caught
throughout
most
of
its
subarctic
range.

MORPHOLOGY/
IDENTIFICATION
AIDS
Body
elongate,
depth
about
4­
4.5
in
standard
length
(
SL),
considerably
compressed
but
variable.
Head
compressed;
mouth
terminal,
moderate
in
size,
directed
moderately
upward,
lower
jaw
extending
to
point
below
eye;
teeth
lacking
on
jaws,
ovate
patch
of
fine
teeth
on
vomer.
Operculum
without
striae.
Fins:
dorsal
(
l),
15­
21;
anal,
13­
21;
pectorals,
about
17;
pelvics,
about
9,
abdominal,
each
with
fleshy
appendage
at
base;
caudal
forked.
Lateral
line
absent.
Scales
large,
cycloid,
38
to
54
along
midside,
modified
along
midventral
line
with
keels
moderately
developed
anterior
to
pelvic
fins
and
strongly
developed
between
pelvic
fins
and
anus.
Vertebrae,
46
to
55.
Gill
rakers,
20
+
45.
Color
bluish
green
to
olive
on
dorsal
surface,
shading
to
silvery
on
ventral
surface,
dusky
on
peritoneum.
Length
to
18
inches.

Recognition:
Silvery,
lacking
black
spots
on
sides
of
body.
Also
without
spines
or
adipose
fin,
no
scales
or
striae
on
head
or
gill
cover,
no
modified
scales
on
side
of
tail
fin,
no
teeth
on
jaws,
keels
along
midventral
line
only
moderately
developed
(
see
Figure
1).

The
above
description
is
based
entirely
on
the
taxonomic
accounts
of
the
Pacific
herring
presented
by
Clemens
and
Wilby
(
1961);
Miller
and
Lea
(
1972);
and
Hart
(
1973).

The
elongate
form
of
the
larva
is
easily
confused
with
other
species.
The
posterior
1
position
of
the
anus
and
the
absence
of
an
adipose
fin
separate
clupeid
larva
from
others
including
sand
lance,
stichaeid,
and
osmerid
larvae.

REASON
FOR
INCLUSION
IN
SERIES
The
Pacific
herring
has
a
long
history
of
exploitation
for
human
consumption
and
reduction
fisheries
for
animal
feeds
and
as
an
item
of
trade.
It
also
provides
food
for
a
wide
variety
of
pelagic,
intertidal,
and
avian
predators.
The
Pacific
herring
is
particularly
susceptible
to
the
influences
of
shoreline
development
because
its
spawning
grounds
are
limited
to
rather
specific
intertidal
and
shallow
subtidal
locations.
This
and
other
life
history
characteristics
also
make
it
susceptible
to
overfishing.
The
larval
stage
is
sometimes
abundantly
found
in
shallow,
nearshore
waters
that
are
susceptible
to
shore­
based
environmental
impacts.

LIFE
HISTORY
Spawning
Pacific
herring,
&
pea
harenguspallasi,
spawn
primarily
on
vegetation
and
substrates
in
intertidal
or
shallow
subtidal
waters
(
Hay
1985).
Substrate
spawning
within
the
genus
CZupea
is
unique
to
C.
harengus
and
occurs
in
both
the
Pacific
and
Atlantic
subspecies
(
Whitehead
1985).
Spawning
grounds
of
Pacific
herring
are
typically
in
sheltered
inlets,
sounds,
bays,
and
estuaries
rather
than
along
open
coastlines
(
Haegele
and
Schweigert
1985a).
Hay
and
Outram
(
1981)
noted
that
the
locations
of
spawning
grounds
were
consistent
from
year
to
year.
The
general
distribution
and
major
spawning
sites
of
Pacific
herring
in
the
Pacific
Northwest
are
shown
in
Figure
2.
Koons
and
Cardwell
(
1981)
provided
a
detailed
map
of
spawning
sites
in
Puget
Sound.

Within
the
range
of
the
species,
there
is
a
latitudinal
cline
in
spawn
timing.
Spawning
may
begin
as
early
as
October
in
California
(
J.
Spratt,
California
Department
of
Fish
and
Game,
Monterrey;
pet­
s.
comm.)
and
continue
as
late
as
July
in
northern
Alaska
(
Haegele
and
Schweigert
1985b).
Spawning
peaks
in
February
and
March
in
the
Pacific
Northwest.
Regardless
of
the
calendar
month,
spawning
is
apparently
timed
to
coincide
with
"
local
spring"
conditions
(
Ware
1985),
a
period
of
increasing
plankton
productivity.

Within
a
season,
spawning
occurs
in
"
waves"
of
several
days
each
separated
by
a
little
over
1
to
several
weeks.
Larger
fish
within
a
stock
tend
to
spawn
first
and
smaller
fish
later
(
Hay
1986).
In
the
actual
spawning
event,
a
rapid
response
in
females
is
triggered
by
the
presence
of
milt
in
the
water
column
(
Stacey
and
Hourston
1982).
Thereafter,
the
behavior
of
males
and
females
within
the
spawning
school
is
simultaneous
and
nearly
identical.
Spawning
waves
are
usually
completed
within
1
or
3
days
and
may
occur
either
during
the
day
or
at
night
(
Hay
1986).
Stacey
and
Hourston
(
1982)
provided
an
excellent
detailed
description
and
illustration
of
the
spawning
sequence.

Ew
and
Larvae
The
eggs
of
Pacific
herring
adhere
to
vegetation
and
other
solid
substrates
and
may
vary
in
density
"
from
a
few
thinly
scattered
eggs
to
more
than
20
layers"
(
Haegele
and
Schweigert
1985b).
Densities
are
highest
in
the
lower
intertidal
and
upper
subtidal
zones.
The
fertilized
eggs
average
1.2­
1.5
mm
in
diameter
(
Hart
1973);
incubation
time
is
about
2­
3
weeks
(
Hay
and
Fulton
1983).

At
hatching,
Pacific
herring
larvae
"
depend
on
endowed
yolk
to
survive"
&
asker
1985).
Yolk­
sac
larvae
move
actively
in
the
wild
(
Westerhagen
and
Rosenthal
1979).
Acuity
of
the
larval
eye
is
low
(
the
minimum
separable
angle
is
about
3
to
4
degrees
in
larvae
<
12
mm
long)
but
is
sufficient
in
larvae
lo­
12
mm
long
to
detect
prey
at
short
distances
(
Blaxter
and
Jones
1967).
The
yolk­
sac
stage
is
generally
completed
within
a
week;
after
that,
condition
factor
(
weightbolume)
begins
to
increase,
coinciding
with
the
onset
of
feeding
(
Westerhagen
and
Rosenthal
1979).

Larval
distribution
depends
on
local
current
patterns
(
Eldridge
1977)
but
may
be
modified
3
Potential
coastal
distribution
of
juvenile
and
adult
WASHINGTON
MILES
KILOMETERS
b­
`::.:,
Crescent
­_

h.
I.:.:.'
City
`
A
CALlFOR
OREGON
Figure
2.
Distribution
of
the
Pacific
berring
in
the
Pacific
Northwest
Region.
Shaded
areas
show
known
spawning
grounds.

3
by
daily
vertical
migrations­­
down
by
day,
up
by
night
(
Hourston
and
Haegele
1980).
Survival
in
these
early
stages
therefore
depends
on
stable
current
patterns
that
promote
larval
retention
in
areas
favorable
to
feeding
and
growth
(
Stevenson
1%
2).

Juveniles
and
Adults
Larval
Pacific
herring
metamorphose
2
to
3
months
after
hatching
(
Hourston
and
Haegele
1980;
Hay
1985)
and
begin
to
school
when
they
reach
lengths
of
25­
40
mm
(
Hart
1973).
During
the
first
summer
after
having
been
spawned,
juveniles
gather
in
large
schools
and
remain
primarily
in
inshore
waters
(
Hay
1985;
Stocker
et
al.
1985).
Juveniles
may
gather
after
their
first
summer
and
move
offshore
until
maturation
(
Stocker
et
al.
1985)
or
they
may
remain
inshore
until
their
first
spawning
(
Hay
1985).
First­
year
juveniles
that
move
offshore
live
mainly
in
waters
with
depths
of
150­
200
m.
Schools
of
immature
fish
(
second
and
third
year)
are
found
in
areas
with
depths
of
100­
150
m
(
Hourston
and
Haegele
1980).
These
offshore
"
juvenile
schools"
appear
to
remain
separated
from
offshore
schools
of
adults
(
Haist
and
Stocker
1985).
Age
at
first
maturity
is
generally
2­
5
years
but
increases
with
increasing
latitude
(
Hay
1985)
and
decreases
with
increasing
exploitation
(
Ware
1985).

Not
all
stocks
of
Pacific
herring
make
this
extensive
offshore
migration.
Many
small
resident
populations
remain
in
coastal
inlets
and
bays
(
Stevenson
1955).
Some
stocks
migrate
offshore
in
the
spring
after
spawning
and
return
from
their
offshore
feeding
grounds
to
inshore
waters
during
the
late
fall
and
early
winter
of
each
year.
The
large
schools
of
adults
may
arrive
inshore
weeks
or
even
months
before
the
spawning
season
(
Hourston
1980).
The
move
from
inshore
"
holding"
areas
to
spawning
sites
may
simply
be
from
deep
water
to
the
adjacent
shallows
(
Hardwick
1973)
or
may
cover
long
distances
in
a
short
time.
On
the
west
coast
of
Vancouver
Island,
BC,
a
tagged
herring
moved
150
km
in
6
days
(
Haegele
and
Schweigert
1985b).
Migratory
and
non­
migratory
stocks
may
mix
while
both
are
inshore
but
apparently
separate
before
spawning
(
Hay
1985).
GSI
and
Fecunddy
The
gonadosomatic
index
(
GSI)
is
an
expression
of
gonadal
weight
as
a
percentage
of
total
body
weight.
It
has
been
found
to
provide
"
a
sensitive
and
quantifiable
estimate
of
maturity"
for
Pacific
herring
(
Hay
and
Outram
1981).
Since
much
of
the
fshery
for
Pacific
herring
is
for
their
eggs,
or
roe,
such
an
index
can
be
extremely
useful
in
correctly
timing
the
fishery
to
maximize
egg
yield.
The
seasonal
pattern
of
gonadal
development
for
male
and
female
Pacific
herring
from
the
lower
eastern
coast
of
Vancouver
Island,
BC
(
adjacent
to
Puget
Sound,
WA)
is
shown
in
Figure
3.
GSI
is
lowest
in
the
months
after
spawning
and
then
begins
to
increase
sharply
in
the
fall.
Large
herring
attain
a
higher
maximum
GSI
than
do
the
smaller
adults
(
Hay
1985).
The
GSI
of
female
Pacific
herring
during
the
spawning
season
was
estimated
at
29%
(
Gunderson
and
Dygert
1988).

Males
begin
gonadal
development
earlier,
develop
faster,
and
reach
a
lower
maximum
GSI
than
females
(
Hay
and
Outram
1981).
Hay
(
1986)
wrote
that
the
"
energy
investment
of
(
MONTH)

Figure
3.
Seasonal
pattern
of
gonadosomatic
index
(
GSI
=
gonad
weight
+
whole
body
weight
X
100%)
in
a
resident
stock
of
Pacific
herring
(
adapted
from
Hay
and
Outram
1981).
Solid
line
is
females;
dashed
line
is
males.
The
pattern
shown
here
may
not
be
the
same
for
migratory
stocks.
Most
populations
have
a
maximum
GSI
above
25%.
c
;
cI
I
,

­

c
Pacific
herring
in
gonadal
development
is
substantial­­
ovaries
usually
exceed
25%
of
the
total
body
weight."
Peak
GSI
in
females
may
be
as
high
as
30%­
32%
(
Hay
and
Outram
1981;
Hay
1986).
Reduction
of
the
number
of
mature
oocytes
by
atresia
prior
to
spawning
was
found
in
experimental
impoundments
and
may
occur
naturally
(
Hay
and
Brett
1988).

The
estimation
of
fecundity
in
Pacific
herring
has
been
related
by
various
researchers
to
length,
weight,
or
age
(
see
Table
1).
A
pattern
of
decreasing
length­
specific
fecundity
with
increasing
latitude
is
widely
reported
(
Katz
1948;
Paulson
and
Smith
1977;
Hay
1985).
However,
exceptions
within
more
restricted
geographic
areas
(
specifically
coastal
British
Columbia)
have
also
been
noted
(
Nagasaki
1958).
Hay
(
1985)
discounted
the
roles
of
GSI
and
egg
size
in
explaining
the
widespread
latitudinal
differences.
He
suggested
instead
that
the
more
southern
stocks
"
have
a
steeper
length­
weight
relationship."
A
similar
explanation
was
given
by
Paulson
and
Smith
(
1977).

Although
herring
from
northern
stocks
are
characterized
by
decreased
size­
specific
fecundity
they
are
also
characterized
by
greatly
increased
maximum
size
(
Katz
1948;
Paulson
and
Smith
1977).
Their
average
and
maximum
Table
1.
Equations
for
the
estimation
of
fecundity
in
Pacific
herring.

Equationa
Source
Location
F=
7.98
(
x~
O­~)
L3.17'
Quisheng
(
1980)
Yellow
Sea
F=
4.2
(
x~
O­~)
SL3n316
Paulson
and
Smith
(
1977)
Prince
William
Sound,
AK
log
F=
3.25
log
SL
Nagasaki
(
1958)
b
Northern
British
Columbia
+
0.08
log
A
­
3.17
F=
2.33
(
x~
O­~)
L3­
028
Hay
(
1985)'
Northern
British
Columbia
F=
555
W"­
782
Ware
(
1985)
Straits
of
Georgia,
1974
F=
III
W1.120
Ware
(
1985)
Straits
of
Georgia,
1980
log
F=
2.16
log
SL
Nagasaki
(
1958)
b
Southern
British
Columbia
+
0.32
log
A
­
0.90
F=
4.19
(
x~
O­~)
L3m3"
Hay
(
1985)
Southern
British
Columbia
F=
­
63920.9
+
496.6
SL
Rabin
and
Barnhart
(
1977)
Humboldt
Bay,
CA
F=
­
56788.4
+
443.4
SL
Hardwick
(
1973)
d
Tomales
Bay,
CA
`
Abbreviations:
F
=
fecundity
(
number
of
eggs),
L
=
length
(
mm),
SL
=
standard
length
(
mm),
A
=
age
(
yrs),
and
W
=
whole
wet
weight
(
g).
bNagasaki
(
1958)
did
not
actually
measure
standard
length
but
approximated
it
by
measuring
from
"
tip
of
snout
to
end
of
silvery
area
on
the
peduncle."
`
Hay
(
1985)
did
not
specify
whether
female
"
length"
was
measured
as
standard,
fork,
total,
or
some
other
measure
of
fish
length.
!
Based
on
data
presented
by
Hardwick
(
1973,
Table
1).

5
fecundities,
therefore,
are
actually
higher.
Female
Pacific
herring
from
Siberian
stocks,
in
fact,
are
reported
to
reach
370
mm
SL
and
have
an
estimated
fecundity
in
excess
of
134,000
eggs
(
Katz
1948).
The
fish
may
be
larger
because
they
are
older.
Older
fish
may
be
more
ubiquitous
in
northern
waters
because
of
sporadic
recruitment
and
the
absence
of
substantial
fisheries.

GROWTH
The
average
length
of
Pacific
herring
at
the
time
of
hatching
is
7.5
mm
(
Hart
1973).
Alderdice
and
Hourston
(
1985)
estimated
a
growth
rate
of
0.48­
0.52
mm/
day
during
the
first
15
days
after
hatching.
Estimations
were
based
on
field
samples
from
Nanoose
Bay,
Vancouver
Island,
BC,
at
ambient
temperatures
of
8.8­
9.1
"
C.
This
rate
is
two
to
three
times
the
growth
rate
observed
by
Boehlert
and
Yoklavich
(
1984)
for
larvae
of
similar
age
raised
in
the
laboratory
at
a
temperature
of
10
"
C
and
salinity
of
15
ppt.
Larvae
metamorphose
into
juveniles
at
a
length
of
25­
40
mm
about
10
weeks
after
hatching
(
Hart
1973;
Hourston
and
Haegele
1980).
Scales
begin
to
appear
at
this
time
and
the
juveniles
develop
the
general
appearance
of
adults.

Haist
and
Stocker
(
1985)
concluded
that
the
growth
rate
of
juvenile
Pacific
herring
was
equally
moderated
by
temperature
conditions
and
density­
dependent
factors.
They
suggested,
however,
that
density­
dependent
effects
on
adult
growth
rate
would
be
evident
only
in
severely
reduced
stocks.
Spratt
(
1981)
found
no
significant
difference
in
the
growth
rates
of
males
and
females.
Size­
at­
age
data
are
summarized
in
Table
2.
Trumble
and
Humphreys
(
1985)
calculated
von
Bertalanffy
growth
equations
for
stocks
from
San
Francisco
Bay
and
the
eastern
Bering
Sea.
Growth
rate
was
higher
in
San
Francisco
Bay
(
K
=
0.59
vs.
K
=
0.18­
0.35),
but
maximum
size
was
greater
in
the
eastern
Bering
Sea
(
Lm
=
299­
314
mm
vs.
Lm
=
208
mm).
This
same
inverse
relation
between
K
and
L=
along
a
latitudinal
gradient
was
noted
for
stocks
in
the
vicinity
of
Puget
Sound
(
Gonyea
and
Trumble
1983).
Growth
was
slowest
and
maximum
size
greatest
in
the
Strait
of
Georgia
(
K
=
0.36,
Lm
=
263
mm).
Growth
rate
was
near
its
highest
(
K
=
0.59)
and
maximum
size
at
its
lowest
(
La
=
197
mm)
in
Case
Inlet.
The
differences,
however,
may
result
because
the
Strait
of
Georgia
stocks
are
migratory
while
the
Case
Inlet
stocks
are
resident.
Less
than
2
degrees
of
latitude
separate
the
two
areas.

Pacific
herring
may
attain
a
total
length
of
18
inches
(
Miller
and
Lea
1972)
and
weight
of
550
g
(
Ware
1985).
Longevity
may
exceed
15
years,
but
few
live
longer
than
9
years
(
Ware
1985).
A
review
by
Gunderson
and
Dygert
(
1988)
listed
longevity
at
10
years.
They
presented
the
size
and
age
of
a
female
at
50%
maturity
as
209
mm
and
3
years.

THE
FISHERY
Native
Americans
have
for
many
centuries
used
nets,
traps,
and
mazes
to
capture
Pacific
herring
for
use
as
a
fresh
or
salted
food
source,
for
trade,
and
for
bait
(
Hourston
and
Haegele
1980;
Trumble
and
Humphreys
1985).
In
the
early
1900'
s,
dry
salted
herring
and
canned
herring
were
important
products
for
human
consumption.
Large
quantities
were
also
reduced
to
fish
meal
and
oil.
Market
demand
or
processing
capacity
usually
limited
catches
(
Hourston
1980).
From
about
a
decade
after
World
War
II
until
the
early
1970'
s,
demand
for
herring
and
for
human
consumption
declined
and
Pacific
herring
supported
only
a
relatively
minor
fishery
in
California
(
Spratt
1981).
To
the
north,
at
the
center
of
the
Pacific
herring's
abundance,
British
Columbia
reduction
fisheries
prospered
until
the
mid­
1960'
s,
when
a
major
decline
occurred.
The
decline
was
apparently
precipitated
by
continued
heavy
fishing
through
a
period
of
several
years
of
poor
recruitment
(
Hourston
1980;
Ware
1985).

The
removal
of
Japanese
import
quotas
in
the
early
1970'
s
opened
a
new
market
to
U.
S.
and
Canadian
herring
fishermen.
The
product
of
this
fishery
was
the
eggs
(
roe)
of
mature
females
for
use
as
kazunoko
(
i.
e.,
caviar),
a
far
more
valuable
product
than
other
uses
of
Pacific
herring.
Consequently,
roe
fisheries
had
6
Table
2.
Size
of
Pacific
herring
at
different
ages.

c
1
Age
(
vears)
Source
and
location
1
2
3
4
5
6
7
8
9
wa
56.5
Wb
22.1
66.0
86.7
106.3
130.6
147.9
164.6
180.1
201.4
Spratt
(
1981),
Tomales
Bay
wb
18.5
57.9
75.9
95.6
116.8
130.5
149.8
156.6
­
Spratt
(
1981),
San
Francisco
Bay
BLc
113.0
164.0
180.0
193.0
207.0
216.0
224.0
231.0
240.0
Spratt
(
1981),
Tomales
Bay
BLC
113.0
161.0
175.0
188.0
200.0
200.0
216.0
219.0
­
Spratt
(
1981),
San
Francisco
Bay
Ld
90.3
153.6
197.9
232.0
255.4
278.4
291.9
­
­
Naumenko
(
1979),
eastern
Bering
Sea
90.0
112.5
136.0
155.5
­
­
­
Haist
&
Stocker
(
1985),
Strait
of
Georgia
`
W
=
mean
whole
wet
weight
(
g)
for
males
and
females
combined
from
Haist
and
Stocker
bW
(
1985,
Table
1).
=
expected
whole
wet
weight
(
g);
calculated
from
observed
mean
body
length
(
this
table)
and
length/
weight
relationships
provided
by
Spratt
(
1981):
Tomales
Bay
W
=
0.2125(
x10­
4,
BL2.93'
6
bBL
San
Francisco
Bay
W
=
0.4278(
x10­
5,
BL3.2317.
=
mean
body
length;
Spratt
(
1981)
measured
"
body
length"
from
"
the
tip
of
the
snout
to
the
dL=
end
of
the
silvery
part
of
the
body."
mean
length;
Naumenko
(
1979)
did
not
mention
whether
"
length"
was
measured
as
standard,
fork,
total,
or
some
other
measure
of
fish
length.

been
initiated
coastwide
by
1973
(
Hardwick
1973;
Trumble
and
Humphreys
1985)
and
have
now
become
the
predominant
fishery
for
Pacific
herring.
Landings
in
California,
Oregon,
and
Washington
since
1979
are
shown
in
Table
3.

The
possibility
of
stable,
substantial
markets
and
high
prices
for
a
quality
product
in
roe
fisheries
led
to
the
development
of
fleets
that
by
the
1980'
s
had
many
times
the
needed
catching
capacity,
especially
since
effort
shifted
to
concentrate
on
the
dense
aggregations
which
typify
this
species'
prespawning
behavior.
To
prevent
overharvest
and
to
avoid
exceeding
processing
capacity,
most
State
or
Provincial
governments
now
manage
these
nearshore
stocks
by
using
limited
entry
permit
systems
for
very
brief
openings­­
sometimes
as
short
as
15
minutes
(
Hourston
1980;
Trumble
and
Humphreys
1985).
The
season
for
a
large
general
area
may
be
longer
(
sometimes
3
months).
The
culturing
of
prespawning
fish
in
impoundments
is
probably
not
a
viable
alternative
at
this
time
because
the
current
fishery
is
doing
well.
Should
impoundment
be
recommended
in
the
future,
experiments
on
impounded
fish
showed
that
mortality
was
low.
Age­,
length­,
and
weight­
specific
fecundity
were
in
the
range
of
fish
in
the
wild.
Density
and
cover
had
no
detectable
influence
on
mortality
(
Hay
and
Brett
1988;
Hay
et
al.
1988).

Herring
for
human
consumption,
other
than
as
sac­
roe,
still
command
a
market
share;

7
Table
3.
Pacific
herring
landings
in
short
tons
(
2,000
lb)
in
California,
Oregon,
and
Washington,
1977­
86.
Data
provided
by
State
management
agencies.

­
Landings
(
short
tons)

State
1979
1980
1981
1982
1983
1984
1985
1986
California
4,623
7,103
6,313
11,331
10,515
2,989
8,305
8,620
Oregon
88
70
74
72
73
89
82
=

Washingtonb
4,263
3,273
966
1,202
666
425
464
493
`
Data
not
available.
bTlle
sac­
roe
fishery
in
Washington
has
been
closed
since
1981
except
for
a
brief
opening
in
1982.
Data
for
198386
are
for
the
bait
fishery
only.

however,
this
and
other
fisheries
for
Pacific
herring
are
much
smaller
than
the
roe
fishery.
Pacific
herring
is
widely
used
for
bait
by
recreational
and
commercial
salmon
trollers,
halibut
longliners,
and
crabbers.
Small
fisheries
also
remain
that
harvest
fish
for
reduction
to
fish
meal
and
for
animal
food
in
zoos
and
aquaria
(
Trumble
and
Humphreys
1985).

Another
product
of
herring
fisheries,
and
more
recently
of
developing
aquacultural
interest,
is
spawn­
on­
kelp.
As
the
name
suggests,
both
the
eggs
and
the
algal
substrate
on
which
they
have
been
laid
are
harvested.
The
alga
is
often
a
kelp
species
but
may
be
any
of
a
number
of
other
algae
as
well­­
e.
g.,
GruciZutiu
(
Hardwick
1973).
The
harvest
of
algae
blanketed
with
naturally
spawned
eggs
had
been
practiced
in
California
since
1965
(
Hardwick
1973).
Again,
this
caviar­
like
product
is
exported
almost
entirely
to
Japan.
However,
the
expansion
of
Japanese
import
markets
in
the
early
1970'
s
encouraged
the
development,
led
by
British
Columbia
fishermen,
of
a
more
reliably
available
product
through
aquacultural
practices.
In
the
closed
pond
method,
schools
of
nearly
ripe
adults
are
encircled
by
nets
and
held
until
they
have
spawned
on
the
fronds
of
kelp
that
have
been
placed
in
the
enclosure.
In
1985,
the
spawn­
on­
kelp
fishery
in
northern
California
began
to
use
an
open­
pond
method
in
which
fronds
of
Macrocystis
pytiiferu
brought
in
from
southern
California
are
attached
to
rafts
in
the
vicinity
of
known
spawning
grounds
(
J.
Spratt,
pers.
comm.).
The
egg­
laden
fronds
provide
a
useful
technique
to
provide
spawning
substrate
at
particular
times
and
thereby
control
the
timing
of
egg
release.
Possible
conflicts
with
the
roe
fishery,
however,
have
caused
a
hesitancy
to
develop
spawn­
on­
kelp
fisheries
in
both
Puget
Sound
and
California.
An
experimental
spawn­
on­
kelp
operation
run
by
Native
Americans
may
be
developed
in
Port
Gamble
Bay,
Washington
(
Dwane
Day,
Washington
Department
of
Fisheries,
Olympia;
per­
s.
comm.).
Interest
has
also
been
expressed
in
developing
a
spawn­
on­
kelp
fishery
in
Coos
Bay,
Oregon
(
Jerry
Butler,
Oregon
Department
of
Fish
and
Wildlife,
Newport;
per­
s.
comm.).

stuckT
Haegele
and
Schweigert
(
1985)
indicated
that
the
variety
of
spawning
sites
and
times
made
it
difficult
to
identify
genetically
distinct
stocks
of
Pacific
herring.
They
remained
convinced,
however,
that
efforts
should
be
made
to
maintain
stock
diversity
because
if
the
time
of
spawning
is
genetically
influenced,
then
the
reestablishment
of
lost
stocks
may
be
impossible.
The
identification
of
separate
Pacific
herring
stocks
has
been
attempted
by
a
number
of
methods,
including
patterns
of
fecundity
(
Katz
1948),
parasitism
(
Arthur
and
Arai
1980),
and
8
growth
(
Gonyea
and
Trumble
1983).
A
complicating
factor
in
the
maintenance
of
stock
diversity
is
the
mixing,
particularly
in
stocks
exploited
by
offshore
food
or
reduction
fisheries,
that
may
occur
before
stocks
separate
for
spawning
(
Buchanan
1983;
Fried
and
Wespestad
1985).

Of
the
"
12
known
spawn
areas"
in
California
(
Trumble
and
Humphreys
1985),
only
San
Francisco
and
Tomales
Bays
support
major
herring
stocks.
Based
on
information
presented
by
Spratt
(
1981,
Table
3)
for
197380,
only
about
1%
of
the
total
California
catch
is
taken
in
the
Pacific
Northwest
region
(
Humboldt
Bay
and
Crescent
City).
Pacific
herring
stocks
in
Oregon
are
relatively
small
but
stable
(
Trumble
and
Humphreys
1985).
Yaquina
Bay
is
the
only
Oregon
bay
that
supports
a
commercial
roe
fishery,
but
Coos,
Umpqua,
and
Tillamook
Bays
support
smaller
bait
and
recreational
fisheries
(
Jerry
Butler,
pers.
comm.).
Most
of
the
Pacific
herring
fisheries
in
Washington
are
in
Puget
Sound
rather
than
along
the
open
coastline.
Gonyea
and
Trumble
(
1983)
suggested
the
existence
of
at
least
three
separate
stocks
(
Strait
of
Georgia,
Northern
Hood
Canal,
and
Case
Inlet)
in
the
vicinity
of
Puget
Sound.

Pophion
Dynamics
and
Management
Hourston
and
Haegele
(
1980)
estimated
that
of
all
stages
in
the
life
history
of
the
Pacific
herring,
larvae
experienced
the
greatest
mortality
(>
99%).
They
further
estimated
an
average
mortality
of
20%
at
the
egg
stage
and
a
total
annual
mortality
(
A)
of
50%
for
adults
(=
an
instantaneous
total
mortality,
2,
of
0.69).
Egg
mortality
may
sometimes
be
much
higher.
Hardwick
(
1973)
estimated
a
loss
to
predation
alone
of
56%~
99%
and
suggested
that
twothirds
of
this
mortality
occurred
within
the
first
3
days
of
spawning.
By
assuming
some
degree
of
compensatory
decrease
in
predation,
however,
he
suggested
that
a
harvest
by
spawn­
on­
kelp
ffihermen
of
"
10%
of
the
eggs
spawned
would
not
significantly
reduce
the
number
of
eggs
that
hatch."
High
egg
loss
may
only
occur
south
of
British
Columbia
on
the
U.
S.
coast,
where
total
egg
production
in
an
area
is
typically
small
and
there
may
be
no
swamping
of
predators
with
many
more
eggs
than
they
can
consume.
In
British
Columbia,
where
up
to
30,000
t
may
be
spawned
over
several
days
in
one
area,
egg
loss
rates
are
much
smaller.
Hourston
and
Haegele
(
1980)
cited
a
correlation
between
juvenile
abundance
and
abundance
at
recruitment
as
evidence
that
year­
class
strength
is
determined
by
the
time
Pacific
herring
have
reached
the
juvenile
stage.

In
a
review
of
Pacific
herring
management,
Trumble
and
Humphreys
(
1985)
wrote
that
"
most
estimates
of
instantaneous
natural
mortality
(
M)
[
for
adults]
fall
very
consistently
in
the
range
of
0.4­
0.5."
Similar
but
slightly
lower
estimates
were
given
by
Fried
and
Wespestad
(
1985,
M
=
0.39)
and
by
Schweigert
and
Hourston
(
1980,
M
=
0.36).
Stocker
et
al.
(
1985)
reported
that
the
instantaneous
natural
mortalities
in
the
Strait
of
Georgia
ranged
from
0.31
to
0.71.
A
review
by
Gunderson
and
Dygert
(
1988)
lists
M
at
0.56,
as
estimated
from
the
gonadosomatic
index
(
GSI).
They
found
a
positive
correlation
for
20
different
species
of
fish,
between
M
and
GSI,
indicating
that
the
GSI
of
those
species
can
predict
the
natural
mortality
rates
for
fishery
management
models.
Schweigert
and
Hourston
(
1980)
reported
a
mean
instantaneous
fishing
mortality
(
F)
for
1972­
79
of
0.59
for
a
heavily
exploited
Canadian
stocks.

By
combining
the
instantaneous
rates
of
natural
and
fishing
mortality
from
Schweigert
and
Hourston's
study
(
1980),
a
total
instantaneous
mortality
rate
(
M
+
F
=
Z)
of
0.95
can
be
calculated.
This
translates
to
an
annual
mortality
rate
(
A
=
1
­
e­`)
of
61%.
Trumble
and
Humphreys
(
1985)
reported
Z
values
of
0.50
(
A
=
39%)
for
a
relatively
unexploited
stock
and
0.62
(
A
=
46%)
for
an
exploited
stock
in
Puget
Sound.
Hourston
(
1980)
reported
that
"
apparent
annual
mortality
rates,
from
75%
to
over
90%,
did
not
depress
the
abundance
of
the
stocks"
in
the
1950'
s.
Annual
mortality
rates
of
75%
and
90%
translate
to
Z
values
of
1.39
and
2.30,
respectively.

Another
common
expression
of
the
influence
of
fishing
pressure
on
stocks
is
the
exploitation
rate
(
E)
which
expresses
instantaneous
fishing
mortality
as
a
percentage
of
instantaneous
total
mortality
(
i.
e.,
E
=
F/
Z).
Judging
from
the
9
Pacific
herring's
ability
to
compensate
for
exploitation
through
increased
growth
and
decreased
age
at
maturity,
Ware
(
1985)
estimated
that
E
should
not
exceed
0.2­
0.3
to
avoid
adversely
affecting
stock
resilience.
Similarly,
Fried
and
Wespestad
(
1985)
considered
the
balance
between
yield
and
maintenance
of
the
spawning
stock
to
produce
a
suggested
exploitation
rate
of
0.2.
Also
rather
conservative
in
his
estimate,
Spratt
(
1981)
suggested
a
quota
of
not
more
than
20%
of
the
previous
year's
spawning
biomass
for
California
roe
fisheries.

Smith
(
1985)
concluded
that
"
theoretical
population
approaches
and
correlative
environmental
indices
are
not
yet
sufficient
for
setting
catch
limits
.
.
.
direct
measurement
of
current
biomass
appears
to
be
necessary
for
setting
clupeoid
quotas."
On
the
basis
of
direct
estimation
techniques
such
as
hydroacoustics
and
spawning
ground
sutveys,
U.
S.
management
agencies
generally
set
quotas
as
a
percentage
(
usually
~
20%)
of
the
standing
stock.

ECOLOGICAL
ROLE
Feeding
Habits
If
other
environmental
conditions
are
sufficient
for
successful
hatching,
it
is
likely
that
larval
survival
is
dependent
on
timing
in
relation
to
predation
and
food
supply
(
Blaxter
and
Hunter
1982;
Alderdice
and
Hourston
1985).
Larval
Pacific
herring
begin
feeding
during
or
immediately
after
the
yolk­
sac
stage,
at
a
length
of
9.5­
11­
O
mm
(
Westernhagen
and
Rosenthal
1981;
Lasker
1985).
Earliest
food
consists
mainly
of
copepods,
invertebrate
eggs,
and
diatoms
(
Hart
1973).
As
larvae,
clupeoids
"
are
characterized
by
straight,
relatively
undifferentiated
guts"
(
Boehlert
and
Yoklavich
1984).

In
laboratory
aquaria,
Boehlert
and
Yoklavich
(
1984)
fed
14C­
labeled
rotifers
to
a
group
of
small
larvae
(
mean
notochord
length,
NL,
=
10.3
mm).
Assimilation
efficiency
ranged
from
44%
to
59%.
In
another
group
of
larger
larvae
(
mean
NL
=
13.8
mm),
which
were
fed
brine
shrimp
nauplii,
assimilation
efficiency
ranged
from
38%
to
68%.
In
both
groups,
assimilation
efficiency
was
inversely
related
to
ingestion
rate.
Even
with
the
decrease
in
assimilation
efficiency,
however,
overall
energy
uptake
was
greater
at
high
food
density.
This
pattern
of
maximizing
energy
by
maximizing
the
number
of
prey
taken
may
enable
larval
herring
to
better
exploit
patchy
food
sources
(
Boehlert
and
Yoklavich
1984).

If
larvae
are
unable
to
feed
soon
after
hatching,
they
may
"
give
up"
&
asker
1985)
and
die
of
starvation.
McGurk
(
1984)
noted
a
decrease
in
growth
and
an
increase
in
mortality
with
increasing
age
of
first
feeding
and
increasing
temperature.
The
time
from
exhaustion
of
the
yolk­
sac
to
the
age
of
irreversible
starvation
decreased
from
8.5
d
at
6
"
C
to
6
d
to
10
"
C.
McGurk
further
suggested
that
catastrophic
mortality
due
to
starvation
may
occur
in
18%­
36%
of
natural
populations
of
first­
feeding
Pacific
herring
larvae.
Westernhagen
and
Rosenthal
(
1981)
attributed
the
occurrence
of
emaciated
larvae
in
1976
in
the
Strait
of
Georgia
to
poor
food
supply
during
the
critical
period
(
sensu
Hjort
1914).
Contrary
to
theory,
however,
they
noted
that
this
same
1976
year­
class
yielded
an
extremely
good
recruitment.
This
finding
seems
to
cast
doubt
on
starvation
as
a
singular
cause
of
year­
class
failures
and
supports
the
conclusion
of
Cushing
(
1985)
that
we
should
perhaps
"
consider
predation
and
starvation
as
equivalent
factors,
not
exclusive
ones."

By
the
time
of
metamorphosis,
barnacle
and
mollusk
larvae,
bryozoans,
rotifers,
and
larval
fish
are
included
in
the
diet,
but
copepods
still
predominate
(
Hart
1973).
Levings
(
1983)
noted
that
the
diet
of
juveniles
45­
55
mm
long
depended
on
invertebrates
that
live
in
eelgrass
beds,
such
as
decapod
larvae,
harpacticoid
and
calanoid
copepods,
gammarid
amphipods,
and
barnacle
larvae.
During
summer,
while
the
fish
attain
lengths
of
70­
100
mm,
copepods
remain
an
important
diet
item
(
Hart
1973).
As
the
herring
mature,
copepods
may
be
superseded
by
euphausids.
However,
since
they
then
move
into
deeper
offshore
waters
where
sampling
is
difficult,
little
is
known
about
their
feeding
until
they
return
as
adults
for
spawning.

10
Pacific
herring
undertake
daily
vertical
migrations.
At
dusk,
they
move
up
in
the
water
column
and
begin
to
disperse
and
feed.
Ingestion
may
either
be
by
visually
mediated
particle
feeding
or,
when
particle
size
is
less
than
300400
pm,
by
filter­
feeding
(
Blaxter
1985).
The
herring
may
gather
near
the
bottom
during
the
day,
but
are
frequently
observed
in
midwater
schools
at
that
time.

During
their
spawning
migration
and
the
inshore
"
holding"
period,
Pacific
herring
may
reduce
their
intake
or
stop
feeding
altogether
(
Ware
1985).
Stacey
and
Hourston
(
1982)
examined
feeding
response
in
laboratory­
held
Pacific
herring
during
various
seasons.
Feeding
response
was
reduced
during
October
and
November
(
the
usual
period
of
spawning
migration)
and
was
at
its
lowest
during
February
and
March
(
the
period
immediately
before
spawning).
Herring
resume
feeding
heavily
after
spawning
and
continue
to
feed
through
the
summer
as
they
move
offshore
(
Stacey
and
Hourston
1982).
Over
summer
and
early
fall,
herring
increase
in
oil
content
and
condition
factor
(
Ware
1985).
These
stored
lipids
apparently
support
the
energy
requirements
of
the
herring
as
they
again
move
inshore
and
undergo
the
gonadal
development
that
precedes
spawning.

This
pattern
of
feeding
offshore
and
spawning
inshore
represents
a
movement
of
considerable
energy
from
offshore
to
inshore
waters.
Hay
and
Fulton
(
1983)
estimated
that
"
about
22%
of
the
total
herring
spawning
stock
biomass
is
released
as
milt
and
eggs."
They
noted
that
this
is
a
small
portion
of
the
carbon
budget
over
an
annual
cycle,
but
may
be
"
substantially
higher
than
maximum
estimates
of
primary
productivity"
over
the
period
from
spawning
until
larval
emergence.
Hay
and
Fulton
(
1983)
suggested
that
the
heavy
organic
input
of
eggs
and
milt
may
promote
a
burst
of
secondary
production,
especially
in
the
form
of
increased
microzooplankton
that
feed
on
the
organic
matter.
They
additionally
suggested
that
the
emergence
of
larvae
from
the
same
spawn
may
coincide
with
this
increased
zooplankton
availability.
In
other
words,
the
larval
herring
may
thus
be
the
beneficiary
of
this
burst
in
local
productivity.
­
of
iumlity
The
earliest
source
of
mortality
is
the
failure
of
eggs
to
hatch
because
of
unsuitable
environmental
conditions,
such
as
salinity
and
temperature
(
Alderdice
and
Hourston
1985).
Other
identified
sources
of
mortality
at
the
egg
(
or
zygote)
stage
include
physical
destruction
by
wave
action
during
storms
(
Hay
and
Miller
1982)
intertidal
exposure
and
desiccation
(
Haegele
and
Schweigert
1985b),
suffocation
due
to
high
egg
densities
or
silting
(
Haegele
and
Schweigert
1985b),
and
most
especially,
predation.
The
list
of
known
egg
predators
is
long,
but
birds
are
most
consistently
cited
as
the
major
predator
(
Hardwick
1973;
Hourston
and
Haegele
1980;
Alderdice
and
Hourston
1985).
Birds
that
feed
on
eggs
in
California
include
the
California
gull
(
Larus
califomicus),
mew
gull
(
L.
canus),
glaucous­
winged
gull
(
L.
glaucescens),
western
gull
(
L.
occidentalis),
coot
(
Fulicu
americana),
and
surf
scoter
(
Melunittu
perspicillatu)
(
Hardwick
1973).
Gulls
feed
directly
on
the
eggs.
Diving
ducks
may
cause
mortality
either
by
direct
consumption
or
by
dislodging
the
egg­
laden
algae
and
setting
them
adrift
to
wash
ashore.

Predation
on
larval
herring
may
be
extremely
high
(
Hourston
and
Haegele
1980).
Medusae
and
other
pelagic
invertebrates
may
be
the
major
predators.
Arai
and
Hay
(
1982)
demonstrated
in
laboratory
aquaria
that
several
species
of
medusae
common
to
coastal
waters
were
capable
of
feeding
on
Pacific
herring
larvae.
Field
surveys
led
them
to
believe
that
"
the
hydromedusae
Sursiu
tubulosu
and
Aequoreu
Victoria
may
be
the
most
abundant
during
the
time
of
peak
herring
larval
abundance."
Both
species
were
collected
with
herring
larvae
in
their
stomachs.
Ctenophores
and
chaetognaths
may
also
be
important
predators
of
larvae
(
Stevenson
1962).
Juveniles
of
a
common
pelagic
hyperiid
amphipod,
Hyperoche
medusurum,
may
occur
in
high
numbers
along
with
abundant
Pacific
herring
larvae
and
may
prey
on
the
larvae
(
Westerhagen
and
Rosenthal
1976).
Hourston
and
Haegele
(
1980)
suggested
that
juvenile
salmonids
on
their
seaward
migration
would
feed
on
larval
herring.
Other
fishes
and
invertebrates
that
have
been
observed
to
prey
11
on
herring
eggs
include
sturgeon
(
Acipenser
sp.),
smelt
(
family
Atherinidae),
surfperches
(
family
Embiotocidae),
and
crabs
(
probably
Cancer
sp.)
(
Hardwick
1973).
Hourston
and
Haegele
(
1980)
also
noted
that
even
juvenile
and
adult
Pacific
herring,
when
in
the
vicinity
of
the
spawning
grounds,
may
feed
"
voraciously"
on
the
eggs
and
newly
hatched
larvae
of
their
own
species.

Again,
little
is
known
of
predation
on
juvenile
herring.
Adults
are
susceptible
to
predation
while
holding
inshore
before
and
during
the
spawning
season.
Among
the
predators
that
feed
on
herring
at
these
times
are
salmon,
seals,
sea
lions,
killer
whales,
dogfish,
and
birds
(
Hourston
and
Haegele
1980).
Pacific
herring
were
the
most
important
prey
of
the
northern
fur
seal,
Callorhinus
ursinus,
in
the
inshore
waters
of
the
northern
coast
of
Washington
(
Perez
and
Bigg
1986).
Pacific
herring,
containing
2.17
kcal/
g,
were
among
the
prey
with
the
highest
energy
content.
When
herring
are
feeding
offshore,
important
predators
include
hake,
sablefish,
dogfish,
Pacific
cod,
and
salmon.

ENVIRONMENTAL
REQUIREMENTS
Salinity
Salinities
at
which
apparently
viable
Pacific
herring
eggs
have
been
found
range
from
3
to
35
ppt
(
Alderdice
and
Velsen
1971;
Alderdice
and
Hourston
1985;
Haegele
and
Schweigert
1985b).
Reported
optima
fall
in
a
somewhat
narrower
range
of
12­
26
ppt
(
Alderdice
and
Velsen
1971).
In
their
own
study,
Alderdice
and
Velsen
observed
maximum
egg
and
larval
survival
in
the
range
of
13­
19
ppt,
around
an
optimum
of
17
ppt.
Galkina
(
1957),
cited
by
Haegele
and
Schweigert
(
1985b),
noted
a
sharp
decrease
in
the
successful
fertilization
of
eggs
at
salinities
below
5
ppt.
Alderdice
and
Hourston
(
1985)
determined
an
"
incipient
lethal
limit"
(
3­
day
LCrO
with
3­
day­
old
larvae)
in
the
range
of
27.5­
31.7
ppt.
The
lower
72­
hour
median
tolerance
limit,
or
the
limit
at
which
50%
mortality
occurred
at
72
hours
of
exposure,
was
2.8­
5.2
ppt
for
larvae
O­
9
days
old
(
Alderdice
et
al.
1979).
The
upper
limit
was
33­
O­
35.8
ppt,
but
substantial
mortality
is
expected
above
20
ppt.
Tenrpemtune
The
range
of
temperatures
at
which
naturally
spawned
eggs
have
been
observed
is
fairly
broad,
ranging
from
below
0
to
14
"
C
(
Alderdice
and
Velsen
1971;
Haegele
and
Schweigert
1985b).
However,
most
natural
spawnings
occur
between
3
and
9
"
C
(
Alderdice
and
Velsen
1971).
Again,
in
their
own
studies,
Alderdice
and
Velsen
determined
that
optimum
egg
development
occurred
in
the
range
of
5.5
to
8.7
"
C
and
that
survival
of
eggs
and
larvae
was
highest
at
about
8.7
"
C.
They
also
noted
that
abnormalities
developed
in
the
lower
jaws
of
larvae
from
eggs
incubated
at
4.0
to
4.7
"
C.
Alderdice
and
Velsen
(
1971)
suggested
that
10
"
C
represented
an
approximate
upper
limit
to
natural
spawning
in
the
Pacific
Northwest.
However,
water
temperatures
of
10
to
12
"
C
are
about
average
for
natural
spawning
grounds
in
California
(
J.
Spratt,
pets.
comm.).

Optimal
temperatures
for
juvenile
and
adult
Pacific
herring
seem
to
be
a
few
degrees
higher
than
those
for
eggs
or
larvae.
Of
the
environmental
factors
analyzed
by
Haist
and
Stocker
(
1985),
sea
surface
temperature
best
fits
their
model
for
juvenile
growth.
The
suggested
optimum
temperature
was
12.2
"
C.
In
their
adult
surplus
energy
model
(
energy
for
somatic
and
gonadal
growth),
an
optimum
temperature
of
11.4
"
C
was
estimated.

Ttmperatwe
and
Salinity
lhteraction
Alderdice
and
Velsen
(
1971)
provided
an
extensive
review
of
the
literature
that
dealt
with
the
effects
of
temperature
and
salinity
on
Pacific
herring.
Most
studies
dealt
with
only
the
egg
and
larval
stages.
Alderdice
and
Velsen
(
1971)
concluded
that
"
Pacific
herring
eggs
are
considered
euryhaline
and
stenothermal."
More
specifically,
their
review
led
them
to
conclude
that
"
Pacific
herring
populations
on
the
North
American
coast
are
confined
to
regions
providing
protected
spawning
waters
of
reduced
salinity
(
8­
28
ppt
S)
at
temperatures
between
about
5.0­
5.5
"
C
and
8.8
or
9
"
C,"
and
that
the
size
of
these
populations
is
related
to
the
physical
extent
of
the
regions
that
provide
these
spawning
requirements.
Alderdice
and
Hourston
(
1985)
narrowed
these
ranges
when
they
concluded
that
the
Pacific
herring
"
appears
to
have
an
optimum
salinity­
temperature
maximum
for
physiological
performance.
during
its
early
life
history
in
the
region
of
12­
17
ppt
at
temperatures
near
6.5­
8.3
"
C."
They
also
determined
that
reproductive
success
occurs
over
a
wide
range
of
temperature
and
salinity
and
that
salinity
plays
a
major
role
below
7
"
C
but
not
above
that
temperature.

Substrate
Since
the
Pacific
herring
is
a
pelagic
rather
than
demersal
species
throughout
most
of
its
life
history,
information
on
substrate
use
is
entirely
related
to
spawning
and
egg
deposition.
References
to
the
specificity
of
spawning
substrate
selection
vary
considerably.
Hardwick
(
1973)
stated
that
once
the
herring
have
moved
into
shallow
waters,
"
they
spawn
on
whatever
substrate
is
available."
Haegele
and
Schweigert
(
1985b),
on
the
other
hand,
observed
that
"
eggs
are
laid
almost
exclusively
on
marine
vegetation,
algae
and
sea
grasses,
although
quite
frequently
eggs
adhere
to
the
rocky
substrate
to
which
the
algae
are
attached."
Within
the
vegetation
as
a
substrate
category,
however,
they
found
that
herring
"
do
not
appear
to
favor
one
type
.
.
.
over
another."
Some
of
the
plants
commonly
used
as
egg
deposition
sites
include
the
seagrass
Zostera
and
several
brown
or
red
algae
of
the
genera
Macrocystis,
Fucus,
and
Gracilaria.
Statements
of
preference
or
of
selectivity
in
onsite
substrate
use
that
are
based
on
nonmanipulative
field
studies
must
be
tempered
by
the
understanding
that
what
is
"
available"
may
have
been
strongly
biased
by
rather
specific
site
selection.

During
laboratory
observations
of
Pacilic
herring
spawning
behavior,
Stacey
and
Hourston
(
1982)
noted
that
"
rigidity
and
texture
appear
to
be
important
components
of
suitable
substrates."
In
general,
a
suitable
substrate
was
judged
to
be
one
that
was
"
rigid,
smooth,
and
free
of
sediment."
Haegele
and
Schweigert
(
1985b)
similarly
noted
that
spawn
was
deposited
on
substrates
"
free
from
silting."
Apparently,
the
presence
of
sediment
on
a
substrate
is
sufficient
to
inhibit
certain
behavioral
transitions
in
the
normal
spawning
sequence
(
Stacey
and
Hourston
1982).
The
use
of
vegetation
as
a
spawning
substrate
raises
a
very
practical
consideration
for
fishery
management.
Two
resources,
the
herring
and
the
algae
upon
which
they
spawn,
must
be
managed.
As
Hardwick
(
1973)
pointed
out
in
the
case
of
Tomales
Bay,
"
a
continued
supply
of
Gracilaria
is
essential
to
a
viable
herring
eggon
seaweed
fishery."
Unfortunately,
this
fishery
ended
in
1977
because
of
silt
contamination
of
the
product
(
J.
Sprat&
per­
s.
comm.).

I
found
no
reference
to
the
oxygen
requirements
of
any
Pacific
herring
life
history
stage
other
than
the
eggs.
Alderdice
and
Hourston
(
1985)
suggested
a
minimum
ambient
oxygen
concentration
of
2.5
mg/
ml
at
the
egg's
surface.
In
other
words,
to
achieve
this
ambient
oxygen
concentration
for
deeper
layers
within
an
egg
mass,
water
column
oxygen
concentration
must
be
much
higher.
The
amount
higher
depends,
of
course,
on
the
water's
ability
to
penetrate
the
egg
mass
and,
therefore,
upon
water
movement.
Haegele
and
Schweigert
(
1985)
suggested
that
eggs
elevated
from
the
bottom
on
vegetation
could
avoid
siltation
and
receive
better
circulation
for
waste
removal
and
oxygenation.

c0Muninants
Of
the
life
stages
of
the
Pacific
herring,
the
larvae
is
most
sensitive
to
the
water­
soluble
fraction
(
WSF)
of
crude
oil.
The
LC50
(
concentration
at
which
median
or
50%
mortality
occurred
was
0.37
ppm
WSF
and
the
LCso
for
indirect
exposure
(
exposure
of
prey
later
consumed)
was
6
ppm
WSF
(
Carls
1987).
Larval
growth
was
correlated
with
larval
feeding,
and
both
were
reduced
after
WSF
exposure.
Exposure
may
affect
growth
by
causing
biochemical
changes
or
by
reducing
feeding
rate.
In
clean
water,
larvae
rapidly
depurated
hydrocarbons
from
tissues,
and
survivors
resumed
growth
and
feeding
(
Carls
1987).

CONCERNS
Various
authors
whose
papers
I
reviewed
expressed
concerns
regarding
informational
needs
or
management.
Some
of
these
are
discussed
below.

One
rather
surprising
generalization
made
by
Cushing
(
1985)
was
that
"
management
of
herring
stocks
based
on
a
great
expenditure
of
research
has
not
been
very
successful."
In
view
of
such
management
difficulties,
the
very
basic
concerns
of
others
take
on
a
special
importance­­
especially
with
respect
to
coastal
development
policy.
I
refer
to
the
statement
by
Trumble
(
1983)
that
"
there
is
no
clear
instance
of
a
stock
successfully
moving
its
spawning
area
as
a
result
of
destruction
or
major
alteration
of
their
original
spawning
area."
A
study
of
the
transplantation
of
eggs
concluded
that
though
eggs
hatched,
a
new
spawning
population
did
not
become
established
(
Hay
and
Marliave
1988).
Clearly,
the
maintenance
of
relatively
undisturbed,
quiescent
areas
of
vegetation
is
a
valid
concern.

Also
pertinent
to
the
maintenance
of
functional
spawning
grounds
is
the
avoidance
of
activities
(
e.
g.,
dredging)
which
would
cause
silting
immediately
before,
during,
and
two
to
three
months
following
the
spawning
season.
The
inhibition
of
spawning
behavior,
suffocation
of
eggs,
and
destruction
of
product
quality
were
mentioned
earlier.
It
also
seems
likely
to
me
that
a
heavy
loading
of
suspended
sediment
would
be
ingested
or
feeding
inhibited
during
a
period
critical
to
the
nutrition
of
both
adult
and
newly
hatched
larvae.
Boehlert
and
Morgan
(
1985)
noted
that
sediment
at
"
low
suspension
levels"
roughly
equivalent
to
natural
conditions
actually
enhanced
larval
feeding
abilities.
Higher
loads,
as
might
be
expected
in
catastrophic
events,
inhibited
feeding.

Other
specific
concerns
relate
to
the
biology
of
the
species
and
the
efficiency
of
fishery
management.
Alderdice
and
Velsen
(
1971)
believed
that
there
was
a
need
for
the
systematic
collection
of
temperature
and
salinity
measurements
at
the
spawning
site
and
for
the
examination
of
temperature
and
salinity
tolerances
of
eggs
and
larvae
from
"
representative
stocks
over
the
range
of
the
species."
Such
data
could
be
used
to
build
a
data
base
for
correlating
recruitment
with
environmental
factors.
Hay
(
1985)
noted
that
the
time
and
place
at
which
recruits
join
the
spawning
population,
the
patterns
of
larval
dispersal
and
mortality,
and
the
distribution
and
ecology
of
Ogroup
juveniles
were
all
areas
that
are
poorly
understood.
Haegele
and
Schweigert
(
1985b)
were
concerned
with
the
maintenance
of
genetic
diversity
in
order
to
assure
no
further
reduction
in
the
production
capacity
of
herring
stocks.
Trumble
and
Humphreys
(
1985)
thought
that
there
was
a
need
to
better
separate
the
effects
of
natural
mortality
and
fishing
in
explaining
stock
declines.
These
same
authors,
as
well
as
Wilimovsky
(
1985)
expressed
concern
with
the
availability
and
precision
of
pre­
season
quota
estimates
to
better
prepare
the
herring
roe
industry.
Wilimovsky
additionally
asked
fishery
managers
to
more
formally
consider
the
risks
of
management
decisions
and
the
value
of
the
information
on
which
those
decisions
are
based.

14
Alderdice,
D.
F.,
and
F.
P.
J.
Velsen.
1971.
Some
effects
of
salinity
and
temperature
on
early
development
of
Pacific
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(
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pullasi).
J.
Fish.
Res.
Board
Can.
28:
1545­
1562.

Alderdice,
D.
F.,
and
AS.
Hourston.
1985.
Factors
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development
and
survival
of
Pacific
herring
(
Clupea
harergus
pallasi)
eggs
and
larvae
to
beginning
of
exogenous
feeding.
Can.
J.
Fish.
Aquat.
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42
(
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68.

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D­
F.,
T.
R.
Rao,
and
H.
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1979.
Osmotic
responses
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and
larvae
of
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Pacific
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Helgol.
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32:
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Arai,
M.
N.,
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D.
E.
Hay.
1982.
Predation
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medusae
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Pacific
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(
CZupea
harengus
paZZasi
larvae.
Can.
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Fish.
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Sci.
39:
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Arthur,
J.
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H.
P.
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1980.
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paZZusi
Valenciennes):
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indicators
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Can.
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Blaxter,
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H.
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Hardwick,
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Hay,
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A
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Hay,
D.
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Outram.
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16
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AS.
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Lasker,
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C.
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(
In
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A
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P.
E.
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C.
1955.
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1985.
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Ware,
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Life
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42
(
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Whitehead,
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J.
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Ring
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Can.
J.
Fish.
Aquat.
Sci.
42
(
Suppl.
1):
3­
20.

Wilimovsky,
N.
J.
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The
need
for
formalization
of
decision
algorithms
and
risk
levels
in
fshety
research
and
management.
Can.
J.
Fish.
Aquat.
Sci.
42
(
Suppl.
1):
258­
262.
c
L.

18
REPORT
E;;
MENTATION
11.
REWRT_
NO.

/
Bmloglcal
Report
82(
11.126)*
1.
Title
and
SubtItle
Species
Profiles:
Life
Histories
and
Environmental
Requirements
of
Coastal
Fishes
and
Invertebrates
(
Pacific
Northwest)­­
Pacific
Herring
­
I5.
Report
Date
)
December
1989
6.

7.
Author(
s)

Dennis
R.
Lassuy
9.
Performing
Organlration
Name
and
Address
Oregon
Cooperative
Fishery
Research
Unit
Oregon
State
University,
104
Nash
Hall
Corvallis,
OR
97331­
3803
_____­.­~.
It.
Sponsoring
Organization
Name
and
Address
U.
S.
Department
of
Interior
U.
S.
Army
Corps
of
Engineers
Fish
and
Wildlife
Service
Waterways
Experiment
Station
Research
and
Development
P.
O.
Box
631
Washington,
DC
20240
Vicksburg,
MS
39180
8.
Performlne
Organization
Rept.
NC
10.
PrplHt/
Task/
Wark
"
n
i
t
No.

11.
COntraCt
0,
Grant(
G)
No.

CC)

r
1%
Type
Of
RCpOrt
6
Penod
C
o
v
e
r
e
d
F
14.

15.
Supplementary
Notes
­

*
U.
S.
Army
Corps
of
Engineers
Report
No.
TR
EL­
82­
4.

16.
Abstract
(
Limit:
200
words)

Species
profiles
are
literature
summaries
of
the
taxonomy,
morphology,
distribution,
life
history,
ecological
role,
and
environmental
requirements
of
coastal
aquatic
species.
They
are
prepared
to
assist
coastal
managers,
engineers,
and
biologists
in
the
gathering
of
information
pertinent
to
coastal
development
activities.
The
Pacific
herring
has
a
long
history
of
exploitation
for
human
consumption,
animal
feed,
and
trade.
It
also
provides
food
for
a
wide
variety
of
pelagic,
intertidal,
and
avian
predators.
The
herring
roe
fishery
has
dominated
catches
since
Japan
opened
its
market
to
imports
in
the
early
1970'
s.
Pacific
herring
spawn
in
quiescent,
nearshore
areas,
primarily
on
marine
vegetation.
Spawning
peaks
in
the
Pacific
Northwest
region
during
February
and
March.
Larvae
remain
inshore,
transform
into
juveniles
after
2­
3
months,
then
move
offshore
in
the
fall.
Adults
move
inshore
on
their
spawning
migration
in
late
fall
and
early
winter.
Optimum
physiological
performance
during
the
early
life
history
is
achieved
at
about
12­
17
ppt
salinity
at
temperatures
near
6.5­
8.3"
C.
It
is
important
to
avoid
siltation
at
or
near
the
spawning
grounds
in
order
to
prevent
disruption
of
spawning
behavior
or
smothering
of
eggs.

17.
Document
Analysis
a.
Dcscr~
ptor.
Fisheries
Salinity
Growth
Oxygen
Feeding
habits
Life
cycles
Temperature
Sediments
b
.
Identihen/~
an.
Ended
Terms
Bays
Export
Inlets
Roe
Baitfish
Gonadosomatic
index
c.
COSATI
Field/
Group
Fecundity
Predators
Substrate
Pacific
herring
Clupea
harengus
pallasi
Life
history
Environmental
requirements
lg.
Availability
Statement
Unlimited
release
'
19.
Securuty
Class
(
This
Repat)

m::::
hi,
Page)

I
Unclassified
21.
NO
.
of
Pages
18
__­
22.
price
See
ANSI­
Z39.16)
OPTIONAL
FORM
272
M­
7:
(
Formerly
NTIS­
35)

Department
of
Commerce
As
the
Nation's
principal
conservation
agency,
the
Department
of
the
Interior
has
responsibility
for
most
of
our
nationally
owned
public
lands
and
natural
resources.
This
includes
fostering
the
wisest
use
of
our
land
and
water
resources,
protecting
our
fish
and
wildlife,
preserving
the
environmental
and
cultural
values
of
our
national
parks
and
historical
places,
and
providing
for
the
enjoyment
of
life
through
outdoor
recreation.
The
Department
assesses
our
energy
and
mineral
resources
and
works
to
assure
that
their
development
is
in
the
best
interests
of
all
our
people.
The
Department
also
has
a
major
responsibility
for
American
Indian
reservation
communities
and
for
people
who
live
in
island
territories
under
U.
S.
administration.

I
U.
S.
DEPARTMENT
OF
THE
INTERIOR
I
FISH
AND
WILDLIFE
SERVICE
I
TAKE
PRIDE
:

in
America
UNITED
STATES
DEPARTMENT
OF
THE
INTERIOR
FISH
AND
WILDLIFE
SERVICE
National
Wetlands
Research
Center
NASA­
Slide11
Computer
Complex
1010
Gause
Boulevard
Slidell,
IA
70458
