Document ID: EPA-HQ-OW-2002-0049-0064
Agency: epa
Document Type: Supporting & Related Material
Title: 
Posted Date: 2003-03-19T05:00Z

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A
B
C
D
E
F
G
H
I
J
K
L
M
N
O
FINAL
REPORT
BIOLOGICAL
CHARACTERISTICS
OF
NEARSHORE
FISHES
OF
CALIFORNIA:

A
REVIEW
OF
EXISTING
KNOWLEDGE
AND
PROPOSED
ADDITIONAL
STUDIES
FOR
THE
PACIFIC
OCEAN
INTERJURISDICTIONAL
FISHERIES
MANAGEMENT
PLAN
COORDINATION
AND
DEVELOPMENT
PROJECT
SUBMITTED
TO
AL
DIDIER
PACIFIC
STATES
MARINE
FISHERIES
COMMISSION
AUGUST
31,
2000
PRINCIPAL
INVESTIGATOR:

Dr.
Gregor
M.
Cailliet,
Professor
Moss
Landing
Marine
Laboratories
8272
Moss
Landing
Road
Moss
Landing,
California
95039
(
831)
632­
4432;
fax
(
831)
632­
4403
cailliet@
mlml.
calstate.
edu
MLML
RESEARCH
STAFF:

Erica
J.
Burton,
Project
Leader;
burton@
mlml.
calstate.
edu
Jason
M.
Cope,
Graduate
Research
Assistant;
jcope@
mlml.
calstate.
edu
Lisa
A.
Kerr,
Graduate
Research
Assistant;
lkerr@
mlml.
calstate.
edu
Moss
Landing
Marine
Laboratories
8272
Moss
Landing
Road,
Moss
Landing,
California
95039
(
831)
632­
4419;
fax
(
831)
632­
4403
CO­
INVESTIGATORS:

Dr.
Ralph
J.
Larson,
Professor
Dr.
Robert
N.
Lea
Department
of
Biology
California
Department
of
Fish
and
Game
San
Francisco
State
University
20
Lower
Ragsdale
Drive,
Suite
100
San
Francisco,
California
94132
Monterey,
California
93940
(
415)
338­
1027
(
831)
649­
2835;
fax
(
831)
649­
2894
rlars@
sfsu.
edu
rlea@
dfg.
ca.
gov
Mr.
David
VenTresca
Mr.
Eric
Knaggs
California
Department
of
Fish
and
Game
California
Department
of
Fish
and
Game
20
Lower
Ragsdale
Drive,
Suite
100
20
Lower
Ragsdale
Drive,
Suite
100
Monterey,
California
93940
Monterey,
California
93940
(
831)
649­
2881;
fax
(
831)
649­
2894
(
831)
649­
7193;
fax
(
831)
649­
2894
dventres@
dfg.
ca.
gov
ehknaggs@
dfg.
ca.
gov
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A
B
C
D
E
F
G
H
I
J
K
L
M
N
O
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49
A
B
C
D
E
F
G
H
I
J
K
L
M
PROJECT
SUMMARY
As
part
of
the
recent
Marine
Life
Management
Act,
the
California
Department
of
Fish
and
Game
is
required
to
produce
a
Fishery
Management
Plan
for
the
nearshore
region
as
described
in
the
Nearshore
Fisheries
Management
Act.
This
requires
the
compilation
of
a
species
list
that
includes
the
species
of
fishes
for
which
size
limits
are
prescribed
in
the
Act
and
other
vulnerable
species,
and
a
detailed
analysis
of
existing
knowledge
of
the
life
history
(
age,
growth,
age
at
maturity,
age
composition,
habitat
utilization,

reproduction,
recruitment,
trophic
relationships,
population
health,
and
stock
structure,
including
genetics)
of
nearshore
fish
species,

especially
those
subject
to
the
newly
developing
live
fish
(
premium)
fishery.
During
this
project,
we
compiled
a
list
of
species
currently
taken
in
the
nearshore
fishery
and
others
that
could
be
particularly
vulnerable
(
n=
124).
In
addition,
we
created
a
life
history
parameter
questionnaire
to
guide
our
literature
search
and
compilation
of
life
history
parameters,
surveyed
the
existing
literature,

constructed
a
life
history
database
of
the
nearshore
fish
species
list,
and
compiled
a
literature
cited
document.
Gaps
in
the
existing
knowledge
are
identified,
and
specific
research
projects
are
proposed
on
such
subjects
as
age
at
maturity,
age
validation,
reproduction,

trophic
interactions,
genetic
variance,
and
mobility
of
fish
stocks
subject
to
the
nearshore
fisheries
of
California.
It
is
hoped
that
this
project
will
result
in
additional
years
of
support
to
answer
questions
that
will
provide
management
agencies
with
sufficient
information
to
effectively
manage
this
fishery.

BACKGROUND
The
1998
Keeley
Bill
(
AB
1241),
which
is
now
law
as
the
Marine
Life
Management
Act
(
MLMA),
has
several
requirements,

including
size
limits
for
selected
species,
and
status
reports
and
Fishery
Management
Plans
(
FMPs)
to
be
prepared
by
the
California
Department
of
Fish
and
Game
(
CDFG)
for
important
commercial
and
sport
fishery
species
in
California.
Funding
has
been
made
available
from
the
Pacific
States
Marine
Fisheries
Commission
(
PSMFC)
to
support
a
research
institution
(
Moss
Landing
Marine
Labs,
MLML)
to
complement
the
work
CDFG
is
doing
on
this
subject.

Nearshore
fisheries
have
existed
in
California
for
decades
(
Lea
et
al.
1999),
but
a
recent
fishery
for
live
fish
that
are
used
in
the
restaurant
business
and
shipped
overseas,
started
in
southern
California
in
the
late
1980s,
spread
to
northern
California
in
the
early
1990s,
and
now
has
become
common
in
central
California
(
McKee­
Lewis
1996).
The
fishery
uses
gear
ranging
from
hook
and
line
to
pole
fishing
and
traps.
The
landings
of
live
finfish
in
California
have
exhibited
an
exponential
increase
from
~
20,000
to
1,200,000
pounds
in
the
past
7
years,
and
were
worth
from
~
$
20,000
to
$
1,300,000.
The
number
of
species
landed
in
this
fishery
range
from
~
5
to
>
50,
depending
upon
location,
season,
and
year.
Even
though
some
reports
of
landings
and
the
status
of
this
fishery
exist
(
CDFG
1998,
Starr
et
al.
1998)
much
information
is
still
needed.

The
role
of
life
histories
in
this
nearshore
fishery
has
not
yet
been
evaluated.
Could
the
live
fish
fishery
be
doomed
to
failure
because
it
targets
mainly
on
small,
immature
specimens
of
shallow,
nearshore
fishes,
some
of
which
are
not
targeted
in
any
other
way?

Certainly,
fisheries
for
juvenile
fishes,
especially
if
they
target
all
areas
where
these
juveniles
live,
cannot
last
long.
The
large
and
increasing
numbers
of
fishes
being
harvested
before
most
get
a
chance
to
reproduce
for
the
first
time
is
of
particular
concern
regarding
the
live
fish
fishery.
Since
rockfish
are
typically
long­
lived,
often
take
many
years
to
mature
(
Echeverria
1987,
Love
and
Johnson
1998,
Lea
et
al.
1999),
and
are
known
to
have
highly­
variable,
successful
recruitment
(
Karpov
et
al.
1995),
harvesting
most
individuals
before
they
mature
and
reproduce
can
be
disastrous
for
their
populations
and
ultimately
the
ecosystem.

The
live
fish
fishery
targets
on
sizes
and
species
of
fishes
that
were
not
heavily
fished
prior
to
now
(
Lea
et
al.
1999).
It
is
known
that
many
of
these
nearshore
fishes
are
very
site­
intensive.
Thus,
any
heavy
fishing
on
their
populations,
most
of
which
have
either
home
sites
or
territories
and
do
not
move
much
after
settling,
could
be
very
deleterious
to
the
local
population
densities.

In
June
1999,
the
University
of
California
Sea
Grant
Marine
Advisory
Program
and
CDFG
sponsored
a
Workshop
on
Assessing
and
Managing
Nearshore
Fisheries
in
Santa
Barbara.
Many
fishery
biologists
attended
this
useful
workshop
and
produced
numerous
recommendations,
including
the
need
to
obtain
the
best
possible
data
on
catch,
effort,
species
and
size
composition,
reproduction,

maturity,
location,
site
specificity,
larval
dispersal
and
recruitment,
and
socio­
economic
variables
to
enable
CDFG
to
evaluate
the
trends
in
this
increasing
fishery.
Page
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Summary
&
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A
B
C
D
E
F
G
H
I
J
K
L
1)
Species
List
The
first
objective
was
to
produce
a
list
of
the
species
now
taken
in
the
nearshore
fisheries
of
California.
This
list
includes
the
species
listed
in
the
MLMA,
but
also
includes
species
frequently
caught
that
are
not
on
this
size
limit
list
and
those
which
could
be
particularly
vulnerable.
Among
personnel
at
MLML,
and
Bob
Lea
from
CDFG,
we
decided
on
a
list
of
species
having
some
or
one
of
the
following
criteria:
1)
occurring
in
40
fathoms
or
less;
2)
listed
in
the
MLMA;
3)
listed
in
McKee­
Lewis
(
1996);
4)

common
in
the
sport
fishery;
5)
common
as
bait
fish;
6)
common
in
public
aquaria
or
the
aquaria
trade;
7)
common
as
bycatch
and
prone
to
overfishing;
or
8)
marketed
incorrectly
and
possibly
prone
to
overfishing.
These
criteria
could,
of
course,
include
almost
all
of
the
nearshore
fishes
off
the
coast
of
California.
Therefore,
our
species
list
was
subjectively
honed
down
from
>
239
to
124
species
(
42
families);
a
relatively
reasonable
number
of
species
to
survey
during
this
8­
month
project
(
Table
1).

2)
Literature
Survey
The
second
objective
was
to
survey
the
existing
published
literature
on
those
subjects
recently
included
in
the
Fisheries
Management
Plan
(
FMP)
and
Biological
Information
Catalog
system
being
developed
by
David
VenTresca
(
CDFG,
Monterey).

Life
history
subjects
in
our
survey
include,
but
are
not
limited
to,
age
and
growth
(
length/
weight/
age
relationships,
growth
parameters,
longevity,
age
composition);
spawning
and
reproductive
status
(
parturition
period,
size/
age
at
sexual
maturation,

fecundity);
recruitment
(
size
at
settlement,
recruitment
to
fisheries);
population
or
stock
genetics;
mortality
estimates
(
natural,

fishing,
total);
species­
habitat
associations;
trophic
interactions
(
feeding,
predation);
and
behavior
(
competition,
movements,

home
ranges).
We
constructed
a
LIFE
HISTORY
QUESTIONNAIRE
(
APPENDIX
A)
to
guide
our
literature
search,
and
used
this
questionnaire
to
compile
life
history
parameters
into
a
FLAT
DATABASE
(
MICROSOFT
 
EXCEL
WORKBOOK:

LIFE_
HISTORIES.
XLS).

3)
Database
Format
To
read,
understand,
and
utilize
the
database
and
its
format,
the
LIFE
HISTORY
QUESTIONNAIRE
(
APPENDIX
A)
should
be
used
while
viewing
the
database.
The
FLAT
DATABASE
is
composed
of
32
MICROSOFT
 
EXCEL
WORKSHEETS
within
a
SINGLE
WORKBOOK
(
LIFE_
HISTORIES.
XLS).
Worksheets
are
organized
by
family
name
or
species
group
(
elasmobranchs
and
flatfishes
only).
Within
a
spreadsheet,
species
are
listed
in
rows
within
column
B
(
Scientific
name).
Life
history
parameter
categories
are
listed
at
the
top
of
the
spreadsheet(
s)
from
left
to
right
(
following
the
order
of
the
LIFE
HISTORY
QUESTIONNAIRE,
APPENDIX
A),
columns
A
thru
DG
(
or
DH,
in
some
cases).
Life
history
information
is
then
contained
within
spreadsheet
cells.
Abbreviations
used
in
the
database
are
listed
at
the
end
of
APPENDIX
A.
The
literature
source
for
each
life
history
parameter
is
listed
at
the
bottom
of
each
column.
Therefore,
the
citations
are
column­
specific,
and
are
footnoted
accordingly
(
i.
e.
renumbered
for
each
column).
The
corresponding
literature
can
be
found
in
the
LITERATURE
CITED
document
(
APPENDIX
B),
where
citations
are
alphabetized.

In
some
cases,
life
history
categories
are
related;
therefore,
information
is
listed
across
a
single
row
(
encompassing
several
columns).
For
example,
the
"
von
Bertalanffy
Growth
Parameters
"
section
within
"
Age
and
Growth"
encompasses
several
columns
(
from
"
Geographic
Area"
to
"
Verification");
and
data
within
one
row
(
across
16
columns)
will
refer
to
one
study.
Other
similar
formats
are
found
in
the
"
Maturity,"
"
Fertilization
and
Spawning
Period,"
"
Habitat
Association
&
Trophic
Interactions,"

"
Genetic
Variance,"
"
Recruitment
Into
Fishery,"
and
"
Mortality"
sections.
Other
columns
stand
alone,
and
footnoted
citations
may
not
correspond
across
a
row,
from
column
to
column.
Page
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Database
Format
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55
A
B
C
D
Family
Name
Scientific
Name
Common
Name
Anarhichadidae
Anarrhichthys
ocellatus
eel,
wolf
Atherinopsidae
Leuresthes
tenuis
grunion,
California
Atherinopsis
californiensis
jacksmelt
Atherinops
affinis
topsmelt
Batrachoididae
Porichthys
notatus
midshipman,
plainfin
Bothidae
Paralichthys
californicus
halibut,
California
Citharichthys
sordidus
sanddab,
Pacific
Citharichthys
stigmaeus
sanddab,
speckled
Carangidae
Seriola
dorsalis
yellowtail
Chimaeridae
Hydrolagus
colliei
ratfish
Clinidae
Heterostichus
rostratus
kelpfish,
giant
Cottidae
Scorpaenichthys
marmoratus
cabezon
Hemilepidotus
hemilepidotus
Irish
lord,
red
Leptocottus
armatus
sculpin,
staghorn
Cynoglossidae
Symphurus
atricauda
tonguefish,
California
Dasyatididae
Urolophus
halleri
stingray,
round
Embiotocidae
Amphistichus
argenteus
surfperch,
barred
Embiotoca
jacksoni
surfperch,
black
Amphistichus
koelzi
surfperch,
calico
Damalichthys
vacca
surfperch,
pile
Amphistichus
rhodoterus
surfperch,
redtail
Rhacochilus
toxotes
surfperch,
rubberlip
Cymatogaster
aggregata
surfperch,
shiner
Embiotoca
lateralis
surfperch,
striped
Hyperprosopon
argenteum
surfperch,
walleye
Phanerodon
furcatus
surfperch,
white
Girellidae
Girella
nigricans
opaleye
Gobiidae
Lythrypnus
dalli
goby,
bluebanded
Lythrypnus
zebra
goby,
zebra
Gillichthys
mirabilis
mudsucker,
longjaw
Haemulidae
Anisotremus
davidsoni
sargo
Xenistius
californiensis
salema
Heterodontidae
Heterodontus
francisci
shark,
horn
Hexagrammidae
Hexagrammos
decagrammus
greenling,
kelp
Hexagrammos
superciliosus
greenling,
rock
Ophiodon
elongatus
lingcod
Table
1.
Nearshore
fishes
of
California
included
in
the
life
history
survey
(
124
species,
42
families).
List
is
alphabetized
first
by
Family
name,
then
by
common
name.
Page
5
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1167
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Species
List
(
Table
1)
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
A
B
C
D
Hexanchidae
Notorynchus
maculatus
shark,
sevengill
Hexanchus
griseus
shark,
sixgill
Kyphosidae
Hermosilla
azurea
zebra
perch
Labridae
Oxyjulis
californica
senorita
Semicossyphus
pulcher
sheephead,
California
Halichoeres
semicinctus
wrasse,
rock
Malacanthidae
Caulolatilus
princeps
whitefish,
ocean
Mugilidae
Mugil
cephalus
mullet,
striped
Muraenidae
Gymnothorax
mordax
eel,
California
moray
Myliobatididae
Myliobatis
californica
ray,
bat
Osmeridae
Thaleichthys
pacificus
eulachon
Spirinchus
thaleichthys
smelt,
longfin
Spirinchus
starksi
smelt,
night
Hypomesus
pretiosus
smelt,
surf
Platyrhinidae
Platyrhinoidis
triseriata
thornback
Pleuronectidae
Platichthys
stellatus
flounder,
starry
Parophrys
vetulus
sole,
English
Eopsetta
jordani
sole,
petrale
Lepidopsetta
bilineata
sole,
rock
Psettichthys
melanostictus
sole,
sand
Lyopsetta
exilis
sole,
slender
Pleuronichthys
coenosus
turbot,
C­
O
Pleuronichthys
decurrens
turbot,
curlfin
Hypsopsetta
guttulata
turbot,
diamond
Pleuronichthys
verticalis
turbot,
hornyhead
Pleuronichthys
ritteri
turbot,
spotted
Pomacentridae
Chromis
punctipinnis
blacksmith
Hypsypops
rubicundus
garibaldi
Rajidae
Raja
binoculata
skate,
big
Raja
inornata
skate,
California
Raja
rhina
skate,
longnose
Raja
kincaidii
skate,
sandpaper
Raja
stellulata
skate,
starry
Rhinobatidae
Rhinobatos
productus
guitarfish,
shovelnose
Sciaenidae
Menticirrhus
undulatus
corbina,
California
Genyonemus
lineatus
croaker,
white
Roncador
stearnsii
croaker,
spotfin
Umbrina
roncador
croaker,
yellowfin
Seriphus
politus
queenfish
Atractoscion
nobilis
white
sea
bass
Page
6
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1167
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­
Species
List
(
Table
1)
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
A
B
C
D
Scorpaenidae
Sebastes
melanops
rockfish,
black
Sebastes
chrysomelas
rockfish,
black­
and­
yellow
Sebastes
mystinus
rockfish,
blue
Sebastes
auriculatus
rockfish,
brown
Sebastes
dallii
rockfish,
calico
Sebastes
nebulosus
rockfish,
China
Sebastes
caurinus
rockfish,
copper
Sebastes
rubrivinctus
rockfish,
flag
Sebastes
carnatus
rockfish,
gopher
Sebastes
rastrelliger
rockfish,
grass
Sebastes
chlorostictus
rockfish,
greenspotted
Sebastes
elongatus
rockfish,
greenstriped
Sebastes
semicinctus
rockfish,
halfbanded
Sebastes
atrovirens
rockfish,
kelp
Sebastes
serranoides
rockfish,
olive
Sebastes
maliger
rockfish,
quillback
Sebastes
rosaceus
rockfish,
rosy
Sebastes
ovalis
rockfish,
speckled
Sebastes
hopkinsi
rockfish,
squarespot
Sebastes
constellatus
rockfish,
starry
Sebastes
saxicola
rockfish,
stripetail
Sebastes
nigrocinctus
rockfish,
tiger
Sebastes
serriceps
rockfish,
treefish
Sebastes
miniatus
rockfish,
vermilion
Sebastes
ruberrimus
rockfish,
yelloweye
Scorpaena
guttata
scorpionfish,
spotted
Sebastolobus
altivelis
thornyhead,
longspine
Sebastolobus
alascanus
thornyhead,
shortspine
Scorpididae
Medialuna
californiensis
halfmoon
Scyliorhinidae
Cephaloscyllium
ventriosum
shark,
swell
Serranidae
Paralabrax
nebulifer
bass,
barred
sand
Stereolepis
gigas
bass,
giant
sea
Paralabrax
clathratus
bass,
kelp
Paralabrax
maculatofasciatus
bass,
spotted
sand
Sphyraenidae
Sphyraena
argentea
barracuda,
California
Squalidae
Squalus
acanthias
shark,
spiny
dogfish
Squatinidae
Squatina
californica
shark,
Pacific
angel
Stichaeidae
Cebidichthys
violaceus
eel,
monkeyface
Xiphister
mucosus
prickleback,
rock
Stromateidae
Peprilus
simillimus
butterfish,
Pacific
Syngnathidae
Syngnathus
leptorhynchus
pipefish,
bay
Syngnathus
californiensis
pipefish,
kelp
Hippocampus
ingens
seahorse,
Pacific
Torpedinidae
Torpedo
californica
ray,
Pacific
electric
Page
7
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1167
5­
2220e
­
Species
List
(
Table
1)
166
167
168
169
A
B
C
D
Triakidae
Mustelus
henlei
shark,
brown
smoothhound
Mustelus
californicus
shark,
grey
smoothhound
Triakis
semifasciata
shark,
leopard
Galeorhinus
zyopterus
shark,
soupfin
Page
8
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1167
5­
2220e
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Species
List
(
Table
1)
123456789
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
A
B
C
D
E
F
G
H
I
ABBREVIATIONS
USED
IN
DATABASE
SE:
Standard
Error
SL:
Standard
Length
FL:
Fork
Length
TL:
Total
Length
BL:
Body
Length
DL:
Dressed
Length
NL:
Notocord
Length
?
L:
measurement
not
defined
m:
meters
cm:
centimeters
mm:
millimeters
If
information
is
not
provided,
the
following
abbreviations
should
be
used
to
explain
why
NA:
Not
Applicable
(
Refers
to
a
life
history
parameter
that
is
not
valid
for
a
species;
e.
g.

egg
stage
not
applicable
for
embiotocids).

NG:
Not
Given
(
Refers
to
a
study
where
similar
parameters
are
provided
but
not
for
the
specified
parameter;
e.
g.
a
maturity
study
with
age
at
first
maturity
but
not
age
at
50%
maturity;
an
age
and
growth
study
where
von
Bertalanffy
growth
parameters
are
provided
for
males
and
females,
but
not
for
the
sexes
combined.)

NF:
Not
Found
(
Refers
to
a
parameter
where
no
information
was
found;
either
it
does
not
exist,
or
we
could
not
find
the
information.)
Page
9
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ABBREVIATIONS
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Anarhichadidae1
Anarrhichthys
ocellatus
1
wolf
eel1
Sea
of
Japan
and
Aleutian
Is.
to
s.
CA
(
Imperial
Beach)
1
1subtidal
to
355
1­
Robins
et
al.
1991
1­
Eschmeyer
1998
1­
Robins
et
al.
1991
1­
Love
et
al
2000
1­
Love
et
al
2000
TAXONOMY
RANGE
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
F
G
H
I
JKLMNOPQ
Length
weight
equation
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)
to
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
Female
NF
NF
1
240
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Love
et
al
2000
AGE
&
GROWTH
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
R
S
T
U
V
W
X
Y
Z
AA
AB
AC
Age
composition
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
NF
NF
NF
NF
NF
NF
NF
NF
1
4
NF
Dioecious
1­
Matarese
et
al.
1989
1­
Parra
et
al.
2000
Promiscuous,
but
within
season
mate
fidelity1
r2
Page
12
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
AD
AE
AF
AG
AH
AI
AJ
AK
Mode
of
reproduction
Parental
Care
M
vs.
F
length
Secondary
Characteristics
Male
Female
Male
Female
Oviparous1,2,3
Yes1,2
1Males>
Females
NF
NF
NF
NF
1­
Watson
1996a
1­
Hart
1973
1­
Garrison
&
Miller
1982
1­
Garrison
&
Miller
1982
2­
Barsukov
1972
2­
Watson
1996a
2­
Cruys
&
Cantera
1980
2­
Friese
1966
3­
Garrison
&
Miller
1982
3­
Fitch
&
Lavenberg
1975
4­
Garrison
&
Miller
1982
5­
Cruys
&
Cantera
1980
6­
Love
1996
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
Breeding
Behavior
Sexual
dimorphism
1st
maturity
50%
maturity
nest­
guarded
by
both
sexes3,4,5,6
males
have
thicker
lips,
more
powerful
head1
females
show
bulging
abdominal
cavity
during
breeding
season2
Males
with
lumpy
head
and
oversized
jaw2
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
AL
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
Fertilized
females
present
Spawning
females
present
Both
reproductive
states
p
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.

NF
NF
NF
NF
California1,2
1­
Watson
1996a
2­
Love
1996
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
REPRODUCTION
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BC
BD
BE
BF
BG
Spawning
Frequency
Size
specific
fecundity
equation
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
NF
1NG
(
3,500
to
5,000)
1,4
10,000
(
NG)
NF
1,2
3
to
4
months
27,000
(
NG)
3
2
months
3
5,000
(
NG)

1­
Garrison
&
Miller
1982
1­
Matarese
et
al.
1989
1­
Watson
1996a
2­
Fitch
&
Lavenberg
1971
2­
Matarese
et
al.
1989
3­
Cruys
&
Cantera
1980
3­
Cruys
&
Cantera
1980
4­
Love
1996
Mean
(
range)
fecundity
in
eggs/
g
female
wt.
Fecundity
Fecundity
range
in
#
eggs
(@
cm,
grams,
or
age
of
female)
Page
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BH
BI
BJ
BK
BL
BM
BN
BO
Size
at
settlement
Size
Fish
Size
Larval
duration
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
(
mm)
Habitat
1
2
months
1150
to
610
NF
5.2
to
5.51
demersal,
adhesive1,2,3,4,5
118.6
to
47.6
pelagic1,2,3,4
2~
200
5.2
to
5.42
233.5
to
330
to
35
to
4
30.5
to
1­
Garrison
&
Miller
1982
1­
Watson
1996a
1­
Watson
1996a
1­
Watson
1996a
1­
Love
1996
1­
Watson
1996a
1­
Watson
1996a
2­
Marliave
1978
2­
Matarese
et
al.
1989
2­
Fitch
&
Lavenberg
1971
2­
Matarese
et
al
1989
2­
Barsukov
1972
3­
Fitch
&
Lavenberg
1975
3­
Friese
1966
3­
Garrison
&
Miller
1982
4­
Garrison
&
Miller
1982
4­
Cruys
&
Cantera
1980
4­
Cruys
&
Cantera
1980
5­
Cruys
&
Cantera
1980
rockfish;
kelp
greenlings1
Eggs
DISPERSAL
&
RECRUITMENT
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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BP
BQ
BR
BS
BT
BU
BV
BW
BX
Fish
Size
Food
habits
Foraging
guild
Foraging
method
When
/
Frequency
Predators
(
SL
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
carnivore
NF
NF
NF
147.6
to
maturity
NF
carnivore
NF
NF
(
fish,
such
as
walleye
pollock)
1
1­
Garrison
&
Miller
1982
1­
Watson
1996a
1­
Barraclough
1967b
(
zooplankton,
such
as
copepods;

larval
walleye
pollock)
1
Larvae
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Food
habits
Life
Stage
Juveniles
Page
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11
12
13
14
15
16
17
18
19
20
21
22
23
24
BY
BZ
CA
CB
CC
CD
CE
Adults
Food
habits
Predators
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
1lincod;
rockfish;
cabezon
carnivore
NF
nocturnal1,2
salmon1
NF
river
otters2
harbor
seals3
1­
Cruys
&
Cantera
1980
1­
Burge
&
Schultz
1973
1­
Hart
1973
1­
Hulberg
&
Graber
1980
1­
Fitch
&
Lavenberg
1971
2­
Barsukov
1972
2­
Hulberg
&
Graber
1980
2­
Barsukov
1972
2­
Stevens
et
al.
1984
3­
Walton
1979
3­
Barsukov
1972
3­
Olesiuk
et
al.
1990
4­
Dewees
&
Gotshall
1974
4­
Fitch
&
Lavenberg
1971
5­
Fitch
&
Lavenberg
1975
5­
Fitch
&
Lavenberg
1975
6­
Bodkin
1986
6­
Hallacher
&
Roberts
1985
7­
Friese
1966
7­
Friese
1966
8­
Cruys
&
Cantera
1980
8­
Hobson
1994
9­
Love
1996
9­
Cruys
&
Cantera
1980
10­
Love
1996
(
crustaceans;
echinoderms,
mostly
sea
urchins,

mollusks;
fish,
such
as
juvenile
rockfishes)

1,2,3,4,5,6,7,8,9,10
rocky
reefs
in
crevices;
stony
bottoms;

artificial
reefs;
kelp
forests
1,2,3,4,5,6,7,8,9
Page
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11
12
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14
15
16
17
18
19
20
21
22
23
24
CF
CG
CH
CI
CJ
CK
CL
CM
CN
CO
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
First
Recruitment
Full
Recruitment
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Total
length
(
age)
Total
length
(
age)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Nucleotide
diversity
(#)
Mean
heterozygosity
GENETICS
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16
17
18
19
20
21
22
23
24
CP
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
DA
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
Rate
of
Exploitation
Method
of
ge
range
use
Rate
of
Exploitation
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
alculate
mort
exploitation
(
u)
ratio
(
F/
Z)
Competition
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
DB
DC
DD
DE
DF
DG
Interactions
/
associations
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
NF
Residential1,2,3
Yes1,2
NF
Yes1,2,3,5
Yes,
even
when
juveniles4
1­
Parra
et
al.
2000
1­
Parra
et
al.
2000
1­
Miller
1982
1­
Parra
et
al.
2000
2­
Hulber
&
Graber
1980
2­
Quast
1968c
2­
Miller
1982
3­
Gascon
&
Miller
1981
3­
Hulber
&
Graber
1980
4­
Garrison
&
Miller
1982
5­
Cruys
&
Cantera
1980
symbiotic
relationships
with
rockfish,

lingcod,
and
sailfin
sculpin;
compete
with
Pacific
giant
octopus1
infested
by
parasites
from
the
family
Gyrodactylidae2
BEHAVIOR
Movement
Page
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Anarhichadidae
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11
12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
A
B
C
D
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Atherinidae1,2
Leuresthes
tenuis
1,2
California
grunion
Atherinopsidae
3
Atherinidae1,2
Atherinopsis
californiensis
1,2
jacksmelt
Atherinopsidae
3
Atherinidae1,2
Atherinops
affinis
1,2
topsmelt
Vancouver
Island,
nearshore,
also
in
estuaries
1
Atherinopsidae
3
TAXONOMY
RANGE
silversides,
California
smelt,
horse
smelt,
blue
smelt,

pescado
del
rey,
Peixe
rey,
Pesce
rey
1,2
smelt,
little
smelt,
least
smelt,
grunion,
silversides
1,2,5
San
Francisco,
northern
California
to
Bahia
Magdelena,

southern
Baja
California
1
Yaquina
Bay,
Oregon
to
at
least
Bahia
Magdelena,

southern
Baja
California
1
bay
smelt,
rainbow
smelt,
panzarotto,
little
smelt,
least
smelt,

silverside,
capron,
jack,
pescadillo,
panzarotti
1,2,3,4
Page
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Atherinopsidae
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
E
F
G
H
I
JK
Total
Length
(
cm)
Max.
reported
Geographic
area
Observed
depth
range
(
m)
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
Female
1
inshore
to
18
m
1Both:
W=
0.0089L3
(
immature
fall
fish
and
winter
fish)
(
L=
Body
lengthNF
119
cm
TL
NF
NF
NF
1Both:
W=
0.0096L3
(
adult
fall
fish)
(
L=
Body
length)

1nearshore
NF
1SL=­
1.167
x
TL(
0.832);
SE:
2.366
148.6
cm
TL
NF
NF
NF
1TL=
1.646
x
SL(
1.200);
SE:
2.841
1TL=
2.645
x
FL(
1.076);
SE:
5.453
1FL=­
1.509
x
TL(
0.925);
SE:
5.057
2Both:
W=
0.00000886L3.03574
(
L=
SL
(
mm))
1SL=­
6.217
x
TL(
0.863);
SE:
1.491
137.6
cm
TL
NF
NF
NF
3Both:
W=
0.0000992L2.59
(
L=
SL
(
mm))
(
specimens
over
135
mm)
1TL=
7.565
x
SL(
1.156);
SE:
1.725
1TL=­
0.228
x
FL(
1.092);
SE:
1.272
1FL=
0.399
x
TL(
0.914);
SE:
1.164
Linf
in
cm
(
SE
1
intertidal
and
nearshore,

also
in
estuaries
Page
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Atherinopsidae
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12
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14
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
L
M
N
O
P
Q
R
S
T
U
V
W
X
r2
Both
Male
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
AGE
and
GROWTH
Growth
Parameters
to
(
SE)

E)
k
(
SE)
Page
24
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1167
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Atherinopsidae
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
Y
Z
AA
AB
AC
AD
AE
Age
composition
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
1,2(
larvae),
5
2,3,54
yrs.
NF
dioecious
polygamous1
oviparous2,3
No
83
yrs
polyandrous1,2
4,6
3,5,6,1011
yrs.
NF
dioecious1
NF
oviparous1,2,3,4
No
79­
10
yrs.
(
30
cm)

911
yrs
(
40.6
cm)

3,7
16­
7
yrs.
NF
dioecious1
polyandrous3
oviparous1,2,3,4
No
5,6,97­
8
yrs.

108
yrs.
Breeding
Behavior
Page
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Atherinopsidae
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
AF
AG
AH
AI
AJ
AK
AL
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
females
larger2,3
NF
51
yr.
1
yr.
NG
NG
scales/
surface
reading
92
yr.
2
yr.
NG
NG
NG
NF
NF
32­
3
yrs
2­
3
yrs
NG
NG
NG
4>
14
cm
(
2­
3
yrs)
>
14
cm
(
2­
3
yrs)
NG
NG
NG
7>
20
cm
(
2
yrs)
>
20
cm
(
2
yrs)
NG
NG
NG
females
larger1,4,5
NF
1,2NG
1,22­
3
yrs.
NG
NG
NG
31­
2
yrs
2­
3
yrs
NG
NG
NG
610­
11
cm
(
1
yr.)
10­
11
cm
(
1
yr.)
NG
NG
NG
712
cm
(
2
yrs)
12
cm
(
2
yrs)
NG
NG
NG
82
yrs.
2
yrs.
NG
NG
NG
102
yr.
2yr.
NG
NG
scales
Sexual
dimorphism
1st
maturity
50%
maturity
Total
length
(
age)
at
REPRODUCTION
Page
26
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1167
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Atherinopsidae
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
Fertilized
eggs
present
Spawning
females
present
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.

No
between
scale
comparison
NF
southern
California4,5,7,8,9,10,13
NG
NG
CalCOFI
area2
*****
*****

southern
CA
to
northern
Baja18
NG
NG
NF
Tomales
Bay,
CA5,17
NG
NG
San
Pablo
Bay,
CA5,11,17
NG
NG
San
Francisco
Bay,
CA5,15,17
central
California6
Moss
Landing
Harbor,
CA15
*****
*****
*****
*****
*****

southern
California11,12,15,19,21
*****
*****

California18,20
northeast
Pacific1
CalCOFI
area2
*****
*****
*****

Pacific
coast10
NG
NG
NF
Coos
Bay,
Oregon3,22
NG
NG
San
Francisco,
CA6,11,17
*****
*****

NG
NG
Moss
Landing
Harbor,
CA15
*****
*****

NG
NG
central
California6
NG
NG
Anaheim
Bay,
CA22
No
No
San
Diego
Bay,
CA22
Newport
Bay,
CA5,14,16,17
*****
*****

CalCOFI
area2
*****
*****
*****
*****

California18
northeast
Pacific1
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
r
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Atherinopsidae
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15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
AZ
BA
BB
BC
BD
BE
BF
BG
Mean
Fecundity
Size
specific
Both
reproductive
states
present
eggs/
g
Number
of
eggs
fecundity
eq
Oct.
Nov.
Dec.
Spawning
Frequency
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
Multiple
spawners1,4,5,9
1,5up
to
3,000
eggs
every
two
weeks
NF
2,3,5,7,8,9,10,1410
days
2,4,91,000­
3,000
eggs
410.5
days(
23
C),
20.5days
(
18
C
31,500­
2,200
eggs
67.5
days
1314
days
62,200
(
avg.)

81,600
to
3,600
eggs
per
spawn
Multiple
spawners6,7,8,11,13
7over
2,000
eggs
per
female
NF
NF
37­
19
days
*****
*****

*****
*****
Multiple
spawners2,7,10,11
2200­
1,000
eggs
per
season
NF
19
days
411.3
days
(
21
C)

records
33,000
eggs
per
spawn/
spawning
every
two
weeks,
12,000­
24,000
per
11,475
eggs
(
13.7
cm
TL)­
2,528
eggs
(
17.1
cm
TL)
3,11,1235
days
(
55
F)
to
less
than
9
days
(
81
F)

several
hundred
(
11­
12
cm)

to1,000
(>
16
cm)
2,3
females
spawn
4­
8
times
a
season1,3,5
Fecundity
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
BH
BI
BJ
BK
BL
Eggs
Size
at
settlement
Size
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
140
days
NF
NF
2,4,51.5­
1.6
mm
230­
40
days
31.5­
2.2
6,71.75­
2.20
viable
limit
for
hatching
14.0­
28.5
C5,8
8<
2.0
mm
viable
limit
for
hatching
16.0­
26.5
C21
eurythermal27
3>
24
days
125
mm
NF
1,41.9­
2.5
mm
22.0­
2.5
mm
31.9­
2.5
mm
90.88­
2.4
mm
100.9­
2.2
mm
4>
18
days
NF
NF
1,4,101.5­
1.7
mm
21.62
mm
(
avg.)

31.4­
1.7
x
1.5­
1.8
mm
benthic
in
estuaries,
bays,
and
lagoons25
found
on
vegetation
in
shallow
water
estuaries
and
bays25
eggs
attached
to
algae,
eel
grass
hydroids,
and
submerged
aquatic
plants4,12,26
eggs
are
deposited
in
the
sand
at
night
just
below
the
high
spring
tide
line1,2,4,6,9,10,11,12,13,15,17,18,19,22
attached
to
algae,
grass,
and
submerged
aquatic
plants4,12,14,16,20,23,24,29
eggs
attached
to
spawning
substrate
and
to
one
another
via
adhesive
filaments1,12,20,30
eggs
spawned
in
estuaries
and
along
open
coast;
primarily
nighttime
spawners3
embryonic
development
occurs
from
12.8
to
26.8
C21
eggs
are
laid
primarily
on
eelgrass
and
adhere
to
macroalgae
on
tidal
flats25,26
eggs
attached
to
spawning
substrate
and
to
one
another
via
adhesive
filaments1,20,29,30
eggs
attached
to
estuarine
and
marine
plants
and
other
floating
objects3,20
eggs
develop
in
sand
and
hatch
when
uncovered
and
agitated
by
surf
of
next
high
tide15,17,18,22,28
duration
of
in­
beach­
sand
incubation
is
dictated
by
tidally
driven
wave
activity6,8,9
grunion
spawn
only
on
three
or
four
nights
following
each
full
or
new
moon6,7,17,26,28
DISPERSAL
&
RECRUITMENT
Page
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
BM
BN
BO
BP
BQ
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
26.5­
20.0
mm
SL
planktonic1
planktivores1
NF
(
larvae
retain
yolk
sac
for
4­
6
days)
1
reduced
salinity
tolerance
of
larvae
with
age8
isopods,
flies,
and
beetles6
NF
17.5­
19.5
mm
SL
planktonic1
planktivores
NF
26.0­
19.8
mm
SL
neuston
of
bays
and
shallow
coastal
waters2,

38.1
mm
TL
(
size
at
hatching)
found
in
mesohaline
salinities4,

57.5­
8.6
mm
(
at
hatching)­
25
mm
abundant
inshore,
common
in
surface
canopy13
shiner
surfperch
and
adult
topsmelt514.3­
15
mm
SL
planktonic1
planktivores
NF
24.3­
21
mm
SL
(
small
crustaceans)
4
35.4
mm
TL
(
size
at
hatching)

44.3­
18.5
mm
tolerates
euryhaline
conditions5
abundant
in
upper
few
inches
of
kelp13
eel
grass
beds
14
found
in
estuarine,
bay,
and
kelp
bed
habitats
and
actively
school
near
the
11
surface
waters
in
harbors,
along
sandy
beaches,
and
in
the
kelp
canopy6,12
(
Stenosomella
spp,
Labidocera
trispinosa
(
nauplii),
Euterpina
acutifrons
,
Euterpina
acutifrons
(
nauplii),
cirriped
nauplii,
bivalve
veligers,
Coscinodiscus
spp.,
Paracalanus
parvus
,
and
Oncea
spp.
)
2
54.3­
4.9
mm
(
at
hatching)
­
18.5
mm
found
over
soft,
unconsolidated
sediments
and
other
substrates11
planktonic,
school
near
the
surface,
found
in
embayments11
(
copepod
nauplii,
tintinnid
protozoans,
cyclopoid
copepods,
and
harpactacoid
copepods)
1,3
beetles,
possibly
annelids,

amphipods,
and
isopods4
larvae
showed
reduced
viability
below
and
above
25.2
C9
Larvae
Food
habits
(
Coscinodiscus
spp.,
Peridinium
spp.,

Stenosomella
spp.,
Euterpina
acutifrons,

E.
acutifrons
(
nauplii),
Acartia
tonsa
,
unidentified
copepods)
2
found
in
neuston
and
upper
water
column
in
coastal
waters2,3,7
sand
worms,
shore
birds,
isopods,

flies,
beetles,
and
ground
squirrels1,3
sand
worms,
shore
birds,
and
ground
squirrels2
found
in
neuston
and
upper
water
column
in
bays
and
coastal
waters2
collected
at
surface
during
the
day
with
a
shift
to
lower
depths
occurring
at
night10
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
BR
BS
BT
BU
BV
When
/
Frequency
Predators
Fish
Size
Habitat
Foraging
guild
fed
during
early
evening1
131.0­
38.8
mm
SL
found
in
shallow
waters
nearshore6
planktivores1
fed
during
early
evening1
kelp
bass
and
yellowtail2
125.8­
37.0
mm
SL
planktivores4
NF
NF
130.1­
39.3
mm
SL
planktivores4,6
218.5­
120.0
mm
(
algae
and
kelpfly
larvae)
2,5
(
algae
and
copepods)
3
abundant
in
surf
grass
beds5
(
insects,
amphipods,
and
copepods
(
51­
75
mm))

resident
populations
in
lagoons10
found
in
neritic,
estuarine,
and
bay
environments9
large
schools
of
juveniles
reside
in
estuaries
and
bays
in
summer;
in
fall,

they
migrate
to
coastal
waters3
surface
waters
in
harbors,
along
sandy
beaches,
and
in
the
kelp
canopy2,11
occur
along
sandy
beaches,
in
kelp
beds,
over
rocky
reefs,
and
around
9
halibut,
bass,
white
croakers,

and
other
large
predators1
Juveniles
preference
for
modal
temperature
of
25
C7
shallow
inshore
waters
in
bays
and
along
coast1
mostly
in
shallow
polyhaline
waters
of
bays
and
along
coast1
shallow
inshore
waters
used
as
nursery
grounds4
found
by
night
and
day
in
surface
waters
close
to
kelp
forests8
(
foraminiferans,
protozoans,
cladocerans,

ostracods,
copepods,
cirrepedian
larvae,

gammarid
amphipods,
caprellid
amphipods,

Callianassa
larvae,
decapod
zoea,
dipteran
adults,
dipteran
larvae,
algae,
and
detritus)
6
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
BW
BX
BY
BZ
CA
Foraging
method
When
/
Frequency
Predators
Habitat
Foraging
guild
NF
NF
planktivores1,2
omnivores8
found
in
midkelp
environment13
NF
kelp
bass
and
yellowtail4
omnivores8,17
(
plankton
and
small
fishes)
1
(
Engraulis
mordax
)
3,9
(
small
fish
and
crustaceans)
5
(
small
crustaceans)
7,19
(
green
algae
and
diatoms)
11
(
zooplankton
and
flora)
11
NF
kelp
bass3
omnivores8,17
(
plankton,
algae,
and
insect
larvae)
1,5
usually
found
near
surface2
(
Engraulis
mordax
)
3,9
abundant
in
surf
grass
beds8
(
planktonic
crustaceans)
4,5,7,14
found
in
euryhaline
conditions4,7,10,12
(
crustaceans,
amphipods,
and
copepods)
8
(
plant
material,
algae,
diatoms,
and
small
crustacea)
8,17
)
7
kelp
column
and
canopy11
(
zooplankton
and
flora)
11
(
detritus,
diatoms,
and
plant
material)
12
found
in
neritic,
estuarine,
and
bay
environments12
(
diatoms,
filamentous
algae,
detritus,
chironomic
midge
larvae,
and
amphipods)
6
found
in
kelp
beds,
harbors,
and
sandy
beaches3
opportunistic
feeder,
characterized
as
a
herbivore,

detrivore,
and
low
level
carnivore10
(
calanoid
copepods,
anchovies,
polychaete
worms,

fish
eggs,
zoea,
nauplei,
cypris
larvae,
and
13
(
Enteromorpha
spp
.,
and
other
filamentous
algae,

benthic
diatoms,
small
crustaceans,
and
detritus)
17
occurs
in
estuaries,
around
kelp
beds,

in
bays,
and
in
open
water2,6,7
often
found
in
lower
reaches
of
coastal
streams5
Food
habits
Fo
California
halibut,
sand
bass,
white
croakers,
and
other
large
predators,

including
man1,2
primarily
filter
feeders,

occasionally
more
predatory1
schools
nearshore,
usually
at
or
near
surface
along
open
coast
and
in
bays1
primarily
filter
feeders,

occasionally
more
predatory1
schools
nearshore
along
open
coast
and
in
bays1
schools
nearshore,
especially
in
bays
and
estuaries
and
near
kelp
beds1,6
feed
near
water
surface,
but
feed
on
bottom
when
in
shallow
water5
found
in
sand­
bottom
kelp
holdfast,

midkelp,
and
rocky­
bottom
kelp­
bed
habitat13
aggregates
by
day
in
surface
waters
close
to
kelp
forests,
but
at
night
larger
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22
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24
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
CB
CC
CD
CE
Foraging
method
When
/
Frequency
Predators
Substocks
NF
Diurnal
feeders1
least
terns
and
doublecrested
cormorants1
NF
larger
fish
and
humans12
Diurnal
feeders2
NF
larger
fish,
such
as
kelp
bass
and
yellowtail3
western
gull
and
Caspian
and
Fosters'
terns8
harbor
seal9,10
Caspian
terns11
sooty
shearwaters13
Diurnal
feeders2,3,4,5
several
subspecies1
larger
fish,
such
as
kelp
bass
and
yellowtail3
olive
rockfish4
birds,
such
as
sea
gulls5
California
gull
chicks7
western
gull
and
Caspian
and
Fosters'
terns8
harbor
seals9
Caspian
terns11
typically
feed
within
10­
15
feet
of
the
surface
over
shallow
rocky
areas
or
in
kelp
beds4
planktonic
feeders
(
ocean
dwellers),

bottom
feeder
(
estuarine
dwellers)
2,1
filter
feeders
and
opportunistic
particulate
predators3
feed
throughout
the
water
column,
but
especially
on
or
near
the
bottom3
Atherinops
affinis
affinis,
subspecies
in
salt
ponds
of
San
Francisco2
feed
throughout
the
water
column,
but
especially
on
or
near
the
bottom3
piscivorous
birds
and
fishes,
including
yellowtail6,14
Adults
ood
habits
marine
fishes
(
yellowtail),
marine
mammals
(
harbor
seal,

northern
fur
seals),
and
sea
birds
(
brown
pelicans,
western
gulls,

least
terns
Brandt's
and
doublecrested
cormorants)
1
yellowtail,
kelp
bass,
sharks,
other
piscivorous
fishes,

piscivorous
birds
(
brown
pelicans
and
gulls),
and
marine
mammals6
halibut,
bass,
white
croakers,
and
other
large
predators2
California
sea
lions,
harbor
seals,
least
terns,
and
Brandt's
and
double­
crested
cormorants1
feed
at
bottom,
swallowing
sand
and
shell
fragments
along
with
plant
and
animal
material
(
bay
dwellers)
1
filter
feeders
and
opportunistic
particulate
predators3
Atherinops
affinis
oregonia
(
ranges
from
Oregon
to
Humbolt
Bay),
A.
affinis
affinis
(
ranges
from
San
Francisco
and
surrounding
areas
to
Monterey),
A.
affinis
littoralis
(
integrades
with
A.
affinis
affinis
from
Monterey
to
Santa
Barbara
and
extends
south
to
San
Diego),
and
A.
affinis
insularum
(
Channel
Islands
form)
3,4
Page
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
CF
CG
CH
CI
CJ
CK
CL
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
GENETICS
Nucleotide
diversity
(#)
Mean
heterozygosity
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
CM
CN
CO
CP
CQ
CR
CS
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Effective
population
size
Fishery
/
Total
Length
(
age)
Fishery
/
Total
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
CT
CU
CV
CW
CX
CY
CZ
DA
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
DB
DC
DD
DE
DF
DG
Interactions
/
associations
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
NF
NF
Yes2,3,12
NF
ectoparasite:
Argulus
melanstictus
4
parasite:
trematode
(
Asymphylodora
atherinopsidis)
7
parasites:
15
families,
16
spp.
(
see
reference)
12
parasite:
trematode
(
Asymphylodora
atherinopsidis)
7
NF
NF
Yes2,3,4,6,7,12
NF
parasite:
nematode10
often
schools
with
topsmelt3,8,11
larvae
school
near
surface
waters7
cleaning
symbiosis
between
topsmelt,
Atherinops
affinis
,

cleaners
and
gray
whale,
Eschrichtius
robustus
1,2
NF
NF
Yes1,2,3,4,6,11
NF
schools
near
surface
waters5,7,9,10,14
larvae
school
near
surface
waters7,11
often
found
schooling
with
jacksmelt3,8
often
associated
with
schools
of
shiner
perch5
school
by
size10
this
species
is
commonly
impinged
on
power
plant
intake
screens10
moderate
to
dense
schools
above
sand
or
mud
bottoms13
resident;
limited
marking
experiments
idicate
the
grunion
are
nonmigratory3
move
inshore
and
into
bays
and
estuaries
to
spawn
during
late
winter
and
early
spring2
this
species
is
seldom
found
far
from
shore2
adult
Oxyjulus
californica
and
juvenile
Herosilla
azurea,

Girella
nigricans,
and
Medialuna
californiensis
have
been
observed
cleaning
adult
Atherinops
affinis8
highly
susceptible
to
adverse
effects
from
pollution
and
habitat
modification10,11
may
school
in
separate
sexes,

particularly
during
spawning
season8
may
school
in
separate
sexes,

particularly
during
spawning
season8
topsmelt
and
grunion
occur
together
in
larval
and
juvenile
stages9
unique
spawning
behavior
makes
them
vulnerable
to
any
reduction
or
change
in
spawning
habitat11
parasites:
copepods(
Bomolachus
pectinatus
,
Caligus
olsoni
,
and
Clauellopsis
),
brachiuran
(
Argulus
melanostictus
),
isopods(
Lironeca
californica
and
Nerocila
californica
),
trematodes
(
Asymphylodora
atherinosidis
and
Lepocredium
manteri
)
1
parasites:
Lepocreadium
manteri,
hyperparasite
(
Nosema
lepocreadii)
6
associated
with
Fundulus
parvipinnis,
Clevandia
ios,

Gillichthys
mirabilis,
and
Gambusia
affinis
3
school
in
murky
waters,
preferable
over
sand14
school
separately
from
jacksmelt,
but
in
the
same
area10
topsmelt
and
grunion
occur
together
in
larval
and
juvenile
stages9
many
move
into
estuaries
during
spring
and
summer
and
into
shallow,
open
coasts
in
fall
and
winter1,2
BEHAVIOR
Movement
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22
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
DH
DI
DJ
DK
DL
DM
DN
DO
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
A
B
C
D
1­
Eschmeyer
1998
1­
Robins
et
al
1991
1­
Eschmeyer
and
Herald
1983
1­
Love
et
al
2000
2­
Robins
et
al
1991
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
3­
Pers
comm
R.
N.
Lea,
CDFG,
Monterey
3­
Hart
1973
4­
Love
1996
5­
Thompson
1919
Page
39
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
E
F
G
H
I
JK
1­
Love
et
al
2000
1­
Clark
1925
1­
Karpov
and
Kwiecien
1988
1­
Love
et
al
2000
2­
Quast
1968a
3­
Schultz
1933
Page
40
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53
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
L
M
N
O
P
Q
R
S
T
U
V
W
X
Page
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59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
Y
Z
AA
AB
AC
AD
AE
1­
Fritzsche
et
al
1985
1­
Matarese
et
al
1980
1­
Emmett
et
al
1991
1­
Gregory
1992
1­
Matarese
et
al
1980
2­
Spratt
1986
2­
Breder
and
Rosen
1966
2­
Moser
1996a
3­
Love
1996
3­
Turner
1960
3­
Love
1996
3­
Fronk
1969
4­
Fitch
and
Lavenberg
1971
4­
Emmett
et
al
1991
4­
Clark
1929
5­
Gregory
1992
5­
Clark
1925
6­
DeLeon
1999a
6­
Emmett
et
al
1991
7­
Clark
1929
7­
Schultz
1933
8­
Clark
1925
9­
Ruagh
1976
10­
Emmett
et
al
1991
2­
Vasquez­
Yeomans
and
Sosa­
Cordero
1992
Page
42
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
AF
AG
AH
AI
AJ
AK
AL
1­
Hart
1973
1­
Fitch
and
Lavenberg
1975
2­
Fitch
and
Lavenberg
1971
2­
Gregory
1992
3­
Thompson
1919
3­
DeLeon
1999a
4­
Feder
et
al
1974
4­
Clark
1929
5­
Schultz
1933
5­
Clark
1925
6­
Carpelan
1955
7­
Emmett
et
al
1991
8­
Turner
1960
9­
Fritzsche
et
al
1985
10­
Schultz
1933
Page
43
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
1­
Matarese
et
al
1989
2­
Moser
1996a
3­
Hart
1973
4­
Spratt
1986
5­
Love
1996
6­
Wang
1981
7­
Walker
1952
8­
Fritzsche
et
al
1985
9­
Fitch
and
Lavenberg
1971
10­
Gregory
1992
11­
DeLeon
1999a
12­
Clark
1929
13­
Clark
1925
14­
Horn
and
Allen
1985
15­
Ruagh
1976
16­
Fronk
1969
17­
Emmett
et
al
1991
18­
C.
D.
F.
G.
1982
19­
Limbaugh
1955
20­
Turner
1960
21­
Feder
et
al
1974
22­
Scultz
1933
Page
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Atherinopsidae
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81
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85
86
87
88
89
90
91
92
93
94
95
96
97
AZ
BA
BB
BC
BD
BE
BF
BG
1­
Spratt
1986
1­
Spratt
1986
1­
Fritzsche
et
al
1985
1­
Matarese
et
al
1989
2­
Love
1996
2­
Love
1996
2­
Fronk
1969
2­
Spratt
1986
3­
Walker
1952
3­
Wang
1981
3­
Emmett
et
al
1991
3­
Love
1996
4­
Fritzsche
et
al
1985
4­
Walker
1952
4­
Hubbs
1965
5­
Fitch
and
Lavenberg
1971
5­
Fitch
and
Lavenberg
1971
5­
Ehrlich
and
Farris
1971
6­
Breder
and
Rosen
1966
6­
Thompson
1919
6­
David
1939
7­
DeLeon
1999a
7­
Emmett
et
al
1991
7­
Walker
1952
8­
Clark
1929
8­
Daugherty
1960a
8­
Fritzsche
et
al
1985
9­
Clark
1925
9­
Moffatt
and
Thomson
1978b
9­
Fitch
and
Lavenberg
1971
10­
Fronk
1969
10­
Gregory
1992
11­
Emmett
et
al
1991
11­
DeLeon
1999a
12­
Daugherty
1960a
12­
Emmett
et
al
1991
13­
Turner
1960
13­
Limbaugh
1955
14­
Daugherty
1960a
Page
45
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Atherinopsidae
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65
66
67
68
69
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71
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77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
BH
BI
BJ
BK
BL
1­
Vasquez­
Yeomans
and
Sosa­
Cordero
1992
1­
Emmett
et
al
1991
1­
Matarese
et
al
1989
1­
Moser
1996a
2­
Reynolds
et
al
1976
2­
Ahlstrom
et
al
1984
2­
Spratt
1986
3­
Clark
1929
3­
Moser
1996a
3­
Love
1996
4­
Ehrlich
et
al
1978
4­
Wang
1981
5­
Ehrlich
and
Farris
1971
5­
David
1939
6­
Moffatt
and
Thomson
1978b
6­
Moffatt
and
Thomson
1978b
7­
Walker
1952
7­
Fritzsche
et
al
1985
8­
Fritzsche
et
al
1985
8­
Breder
and
Rosen
1966
9­
Thompson
et
al
1936
9­
Clark
1929
10­
Fitch
and
Lavenberg
1971
10­
Emmett
et
al
1991
11­
Middaugh
et
al
1983
12­
Gregory
1992
13­
Breder
and
Rosen
1966
14­
DeLeon
1999a
15­
Moffatt
and
Thomson
1978a
16­
Horn1980
17­
Clark
1925
18­
Spratt
1981
19­
Thomson
and
Muench
1976
20­
Ruagh
1976
21­
Ehrlich
and
Muszynski
1981
22­
Thompson
1919
23­
Fitch
and
Lavenberg
1975
24­
Fronk
1969
25­
Emmett
et
al
1991
26­
C.
D.
F.
G.
1982
27­
Ehrlich
et
al
1979
28­
Daugherty
1960a
29­
Turner
1960
30­
Feder
et
al
1974
Page
46
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Atherinopsidae
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
BM
BN
BO
BP
BQ
1­
Fritzsche
et
al
1985
1­
Matarese
et
al
1980
1­
Moser
1996a
1­
Fritzsche
et
al
1985
2­
Fitch
and
Lavenberg
1971
2­
Moser
1996a
2­
Wang
1981
2­
Watson
and
Davis
1989
3­
Gregory
1992
3­
Middaugh
et
al
1990
3­
Fritzsche
et
al
1985
3­
Kauffman
et
al
1981
4­
Thompson
1919
4­
DeLeon
1999a
4­
Middaugh
et
al
1990
4­
Feder
et
al
1974
5­
Fronk
1969
5­
Emmett
et
al
1991
5­
Middaugh
and
Shenker
1988
6­
Daugherty
1960aa
6­
Greogory
1992
7­
Watson
and
Davis
1989
8­
Reynolds
et
al
1976
9­
Ehrlich
and
Muszynski
1981
10­
Kauffman
et
al
1981
11­
Emmett
et
al
1991
12­
Turner
1960
13­
Feder
et
al
1974
14­
Hoffman
1986
Page
47
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Atherinopsidae
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63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
BR
BS
BT
BU
BV
1­
Watson
and
Davis
1989
1­
Fitch
and
Lavenberg
1971
1­
Moser
1996a
1­
Wang
1981
1­
Fritzsche
et
al
1985
2­
Turner
1960
2­
Emmett
et
al
1991
2­
Gregory
1992
2­
Fitch
and
Lavenberg
1975
3­
DeLeon
1999a
3­
Horn
and
Allen
1985
4­
Horn
1980
4­
Ruagh
1976
5­
DeMartini
1981
5­
Quast
1968d
6­
Reynolds
et
al
1977
6­
Klingbeil
1972
7­
Reynolds
et
al
1977
7­
Mitchell
1953
8­
Hobson
et
al
1981
9­
Emmett
et
al
1991
10­
Allen
and
Horn
1975
11­
Turner
1960
Page
48
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64
65
66
67
68
69
70
71
72
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74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
BW
BX
BY
BZ
CA
microcarnivore15
sensitive
to
low
and
high
temperatures15
1­
Ruagh
1976
1­
Fritzsche
et
al
1985
1­
Moser
1996a
1­
Love
1996
2­
Fitch
and
Lavenberg
1971
2­
Love
1996
2­
Fritzsche
et
al
1985
3­
Love
and
Ebeling
1978
3­
Gregory
1992
3­
Coyer
and
Hall
1993
4­
Turner
1960
4­
Fronk
1969
4­
Fitch
and
Lavenberg
1975
5­
Moyle
1976
5­
Gregory
1992
6­
Carlisle
et
al
1964
6­
Moyle
1976
7­
DeLeon
1999a
7­
Carlisle
et
al
1964
8­
DeMartini
1981
8­
DeLeon
1999a
9­
Hobson
et
al
1981
9­
Coyer
and
Hall
1993
10­
Carpelan
1955
10­
Allen,
L.
G.
1982
11­
Quast
1968b
11­
Barry
et
al
1996
12­
Emmett
et
al
1991
12­
Horn
and
Allen
1985
13­
Limbaugh
1955
13­
Ruagh
1976
14­
Feder
et
al
1974
14­
Fitch
and
Lavenberg
1975
15­
Doudoroff
1945
15­
Quast
1968b
16­
Quast
1968d
17­
Emmett
et
al
1991
18­
Klingbeil
1972
19­
Turner
1960
occur
along
sandy
beaches,
in
kelp
beds,
over
rocky
reefs,
and
around
found
in
sand­
bottom
kelp
holdfast,

midkelp,
and
rocky­
bottom
kelp­
bed
habitat13
(
planktonic
crustaceans,
larval
bryozoans,
algae,
and
insect
larvae)
16
(
diatoms,
algae,
mysids,
jacksmelt
eggs,
and
calanoid
copepods)
13
(
diatoms,
copepods,
ostracods,
mysids,
algae,

amphipods,
and
zoea
larvae)
13
estuary/
bay
inhabitants
(
algae,
diatoms,
small
crustaceans,
benthic
invertebrates);
oceanic
inhabitants
(
gammarid
and
caprellid
amphipods,

mysids,
ostracods,
copepods,
and
crustacean
larvae)
17
(
detritus,
foraminiferans,
gastropods,
ostrocods,

copepods,
cumaceans,
tanaidaceans,
gammarid
amphiods,
caprellid
amphipods,
Callianassa
larva,

dipterans,
A.
affinis
eggs,
and
algae)
18
primarily
a
marine
fish
that
prefers
estuaries,
bays,
sloughs,
and
lagoons12
individuals
disperse
close
beneath
the
water
surface
over
deeper
water9
Page
49
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68
69
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79
80
81
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83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
CB
CC
CD
CE
4
subspecies
in
California:
San
Francisco
topsmelt,
Atherinops
affinis
affinis
;
bay
topsmelt,
A.
affinis
littoralis
;
kelp
topsmelt,
A.
affinis
cedroscensis
;
and
the
island
topsmelt,
A.
affinis
insularum
5
1­
Fronk
1969
1­
Fritzsche
et
al
1985
1­
Love
1996
1­
Fronk1969
2­
DeLeon
1999a
2­
Coyer
and
Hall
1993
2­
Fitch
and
Lavenberg
1971
2­
Carpelan
1955
3­
Ruagh
1976
3­
Hobson
et
al
1981
3­
Carlisle
et
al
1964
3­
Fronk
1969
4­
Quast
1968d
4­
Emmett
et
al
1991
4­
Love
and
Ebeling
1978
4­
Turner
1960
5­
Emmett
et
al
1991
5­
Hobson
et
al
1981
5­
Fronk
1969
5­
Feder
et
al
1974
6­
Emmett
et
al
1991
6­
Hubbs
1918
7­
Dierks
1990
8­
Harvey
1982
9­
Trumble
1995
10­
Torok
1994
11­
Parkin
1998
12­
Daugherty
1960a
13­
Krasnow
1978
14­
Feder
et
al
1974
6
subspecies:
Atherinops
affinis
littoralis,

A.
affinis
magdalenae,
A.
affinis
cedroscensis,
A.
affinis
guadalupae,
A.

affinis
affinis
,
and
A.
affinis
insularum
6
feed
near
water
surface,
but
feed
on
bottom
when
in
shallow
water5
Page
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Atherinopsidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
CF
CG
CH
CI
CJ
CK
CL
Page
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Atherinopsidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
CM
CN
CO
CP
CQ
CR
CS
Page
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Atherinopsidae
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58
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
CT
CU
CV
CW
CX
CY
CZ
DA
Page
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Atherinopsidae
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57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
DB
DC
DD
DE
DF
DG
due
to
spawning
habitat
any
pollution
or
alteration
of
bays
and
harbors
may
have
a
deterious
effect
on
the
species
11
parasites:
18
families,
19
spp.
(
see
reference)
12
parasite:
isopod
(
Lironeca
californica
)
13
1­
Fritzsche
et
al
1985
1­
Love
1996
1­
Matarese
et
al
1980
2­
Swartz
1981
2­
Emmett
et
al
1991
2­
Moser
1996a
3­
Allen,
L.
G.
1982
3­
C.
D.
F.
G.
1982
3­
Love
1996
4­
Benz
et
al
1995
4­
Coyer
and
Hall
1993
5­
Moyle
1976
5­
Fitch
and
Lavenberg
1975
6­
Canning
and
Olson
1980
6­
Gregory
1992
7­
Olson
1977
7­
DeLeon
1999a
8­
Sikkel
1986
8­
Ruagh
1976
9­
Ehrlich
et
al
1978
9­
Fitch
and
Lavenberg
1975
10­
Emmett
et
al
1991
10­
Fronk
1969
11­
C.
D.
F.
G.
1982
11­
Emmett
et
al
1991
12­
Love
and
Moser
1983
12­
Limbaugh
1955
13­
Bennett
1993
13­
Turner
1960
14­
Feder
et
al
1974
Page
54
of
1167
5­
2220e
­
Atherinopsidae
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
DH
DI
DJ
DK
DL
DM
DN
DO
Page
55
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1167
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Atherinopsidae
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99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
A
B
C
D
Page
56
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99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
E
F
G
H
I
JK
Page
57
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­
Atherinopsidae
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
L
M
N
O
P
Q
R
S
T
U
V
W
X
Page
58
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Atherinopsidae
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100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
Y
Z
AA
AB
AC
AD
AE
Page
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100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
AF
AG
AH
AI
AJ
AK
AL
Page
60
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1167
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Atherinopsidae
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
Page
61
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1167
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­
Atherinopsidae
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
AZ
BA
BB
BC
BD
BE
BF
BG
Page
62
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1167
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­
Atherinopsidae
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
BH
BI
BJ
BK
BL
Page
63
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1167
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­
Atherinopsidae
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
BM
BN
BO
BP
BQ
Page
64
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1167
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­
Atherinopsidae
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
BR
BS
BT
BU
BV
Page
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Atherinopsidae
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100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
BW
BX
BY
BZ
CA
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113
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118
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CB
CC
CD
CE
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112
113
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115
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CF
CG
CH
CI
CJ
CK
CL
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CM
CN
CO
CP
CQ
CR
CS
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112
113
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CT
CU
CV
CW
CX
CY
CZ
DA
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113
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128
DB
DC
DD
DE
DF
DG
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DH
DI
DJ
DK
DL
DM
DN
DO
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37
38
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40
41
42
43
44
45
46
47
48
49
50
51
52
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Batrachoiddidae1,2
Porichthys
notatus
1,2
plainfin
midshipman
1
intertidal
to
383
m
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Eschmeyer
and
Herald
1983
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
3­
Hart
1973
southeastern
Alaska
(
Sitka)?
British
Columbia
(
Smith
Sound)
to
Bahia
Magdelena,
southern
Baja
California
1
northern
midshipman,
bullhead,

singing
fish,
toad
fish,
grunter,

midshipman
1,2,3
TAXONOMY
RANGE
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
F
G
H
I
J
K
L
M
Total
Length
(
cm)
Max.
reported
Geographic
area
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
Female
Both
Male
1W=
0.0274
L0.605
(
sessile
larvae;
L=
SL)
1TL=
1.091
SL
+
4.8
138
cm
TL
NF
NF
NF
NF
NF
2W=
0.0087L3.04
1­
Crane
1981
1­
Demartini
1990
1­
Love
et
al
2000
2­
Phillips
1973
Linf
in
cm
(
SE)

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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
N
O
P
Q
R
S
T
U
V
W
X
r2
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
k
(
SE)
AGE
and
GROWTH
to
(
SE)
Growth
Parameters
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
Y
Z
AA
AB
AC
AD
AE
Age
composition
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
1,2,3
17
yrs.
NF
dioecious
polygamous1,3,5
oviparous1,2
nest
guarded
by
male1,2,3,5,6,7,8,9,11,12
polygynous2,3,4
1­
Campana
1984b
1­
Love
1996
1­
Hart
1973
1­
Moser
1996a
1­
Moser
1996a
2­
Phillips
1973
2­
Demartini
1988
2­
Garrison
and
Miller
1982
2­
Love
1996
3­
Ibara
1967
3­
Arora
1948
3­
Arora
1948
4­
Knapp
et
al
1999
4­
Knapp
et
al
1999
5­
Garrison
and
Miller
1982
5­
Breder
and
Rosen
1966
6­
Baxter
1999c
7­
Phillips
1973
8­
Garrison
and
Miller
1982
9­
Hubbs
1920a
10­
Turner
et
al
1969
11­
Allen,
M.
J.
1982
12­
Feder
et
al
1974
male
clears
area
for
egg
laying
and
guards
nest10
males
guard
nests
with
clutches
at
very
different
stages
of
development4,8
androgens
play
an
important
role
in
parental
care
behavior
of
males4
Breeding
Behavior
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22
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24
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AF
AG
AH
AI
AJ
AK
AL
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
males
larger1,2
13­
4
yrs
3­
4
yrs
NG
NG
NG
218­
19
cm
14­
15
cm
NG
NG
NG
large
sound
producing
muscles
in
males2
1­
Fitch
and
Lavenberg
1971
1­
Brantley
and
Bass
1994
1­
Phillips
1973
2­
Ibara
1967
2­
Ibara
1967
2­
Ibara
1967
1st
maturity
50%
maturity
Sexual
dimorphism
two
male
reproductive
morphs,

dimorphic
body
and
vocal
muscle
traits
1
Total
length
(
age)
at
REPRODUCTION
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22
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
Fertilized
eggs
present
Spawning
females
present
Both
reprod
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.

NG
NG
NF
CalCOFI
area1
NG
NG
San
Franscisco
Bay,
CA3
central
California2
*****
*****

Santa
Barbara,
CA2
California4
1­
Moser
1996a
2­
Love
1996
3­
Baxter
1999c
4­
Feder
et
al
1974
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BA
BB
BC
BD
BE
BF
BG
Mean
Fecundity
Size
specific
ductive
states
present
eggs/
g
Number
of
eggs
fecundity
eq
Nov.
Dec.
Spawning
Frequency
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
1,5200­
500
eggs
per
nest
NF
1F=
14+
5.13W,
r=
0.698
(
1978)
1,3,4,5,616­
20
days
2up
to
1,600
eggs
per
nest
1F=
40+
3.91W,
r=
0.652
(
1979)
215
days
3,6200
eggs(
avg.
female)
1F=
29+
4.92W,
r=
0.741
(
1984)

1F=
17+
5.95W,
r=
0.698
(
1985)

1F=
37+
4.28W,
r=
0.565
(
1986)

480­
200
eggs
per
season
720­
800
eggs
per
nest
1­
Fitch
and
Lavenberg
1971
1­
Hart
1973
1­
DeMartini
1990
1­
Hart
1973
2­
Baxter
1999c
2­
Crane
1986
2­
Love
1996
3­
Phillips
1973
3­
DeMartini
1988
3­
Crane
1981
4­
Love
1996
4­
Phillips
1973
5­
Arora
1948
5­
Garrison
and
Miller
1982
6­
Fitch
and
Lavenberg
1971
6­
Ibara
1967
7­
Garrison
and
Miller
1982
31,000­
1,200
eggs
per
nest
(
represents
production
of
5­
6
females)

single
spawner,
both
sexes
die
after
spawning1,2
females
usually
die
after
spawning,
males
may
spawn
more
than
once3
Fecundity
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30
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BH
BI
BJ
BK
BL
BM
Eggs
Size
at
settlement
Size
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
NF
NF
14.0­
6.0
mm
x
4.0­
8.0
mm
eggs
attached
to
nest
wall1
crabs
and
snails1
24.0­
7.5
mm
36.0
mm
44.0­
6.0
mm
1­
Crane
1981
1­
Moser
1996a
1­
Moser
1996a
1­
Crane
1986
2­
Wang
1981
2­
Hart
1973
3­
Breder
and
Rosen
1966
3­
Wang
1981
4­
Hubbs
1920a
4­
Crane
1981
2
5­
Arora
1948
6­
Fitch
and
Lavenberg
1971
7­
Garrison
and
Miller
1982
8­
Turner
et
al
1969
9­
Hubbs
and
Schultz
1939
10­
Limbaugh
1955
11­
Allen,
M.
J.
1982
12­
Feder
et
al
1974
demersal,
adhesive
eggs
spawned
in
shallow
intertidal
coastal
waters3,5
14­
5
weeks
larvae
break
away
from
disc
and
swim
free
of
the
eggs
laid
on
the
underside
of
intertidal
and
subtidal
rocks4,7,8,10,12
eggs
spawned
in
shallow
water
or
intertidal
zone
in
a
nest2,9,11
DISPERSAL
&
RECRUITMENT
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22
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24
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BN
BO
BP
BQ
BR
BS
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
17.0­
19
mm
larvae
attached
to
nest
wall1
planktivore3
NF
NF
27.0­
30
mm
TL
(
crustacean
larvae)
1,2
(
copepods)
4
(
crustacean
larvae
and
zooplankton)
5
1­
Moser
1996a
1­
Moser
1996a
1­
Hart
1973
1­
Arora
1948
2­
Wang
1981
2­
Wang
1981
2­
Crane
1986
3­
Arora
1948
3­
Love
1996
4­
Crane
1981
5­
Allen,
M.
J.
1982
attached
to
substrate
in
shallow
intertidal
coastal
waters2,3
Food
habits
attached
larvae
feed
on
yolk
reserves,
free
swimming
larvae
feed
on
small
crustacean
larvae
and
other
l
kt
5
Larvae
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30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BT
BU
BV
BW
BX
BY
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
NF
planktivore1
NF
NF
harbor
seals1
(
copepods)
1
young
gradually
migrate
to
deeper
waters2
(
copepods
and
amphipods)
2
(
small
crustaceans
and
zooplankton)
3,4
seek
cover
by
burrowing
in
the
mud
or
sand4
1­
Wang
1981
1­
Ibara
1968
1­
Torok
1994
2­
Phillips
1973
2­
Phillips
1973
3­
Garrison
and
Miller
1982
3­
Garrison
and
Miller
1982
4­
Limbaugh
1955
avoid
light
and
remain
buried
in
the
sand
during
the
day,
coming
out
at
night
to
feed
3
living
on
bottom
of
shallow
and
deeper
coastal
waters1
Juveniles
Food
habits
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BZ
CA
CB
CC
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
carnivore
detect
prey
by
motion
rather
than
sight1
nocturnal1,2,3,4,5,6
(
fishes
and
crustaceans)
1,6
feeds
in
midwater2
(
krill,
mysid
shrimps,
amphipods,
and
squids)
2
(
shrimplike
crustaceans
and
small
fish(
anchovies))
3
(
euphausiids
and
mysid
shrimp)
5
exhibit
diel
vertical
migration4,5
(
euphausiids,
mysids,
and
gammarids)
7
found
in
rocky­
bottom
kelp­
bed
habitat7
males
fast
while
guarding
eggs6
1­
Moser
1996a
1­
Hart
1973
1­
Ibara
1968
1­
Love
1996
2­
Love
1996
2­
Love
1996
2­
Phillips
1973
2­
Fitch
and
Lavenberg
1971
3­
Fitch
and
Lavenberg
1971
3­
Fitch
and
Lavenberg
1971
3­
Ibara
1967
3­
Phillips
1973
4­
Ibara
1968
4­
Phillips
1973
4­
Cailliet
et
al
1979
4­
Ibara
1967
5­
Phillips
1973
5­
Ibara
1967
5­
Allen,
M.
J.
1982
5­
Cailliet
et
al
1979
6­
Hubbs
and
Schultz1939
6­
Hubbs
1920a
6­
Garrison
and
Miller
1982
6­
Allen,
M.
J.
1982
7­
Limbaugh
1955
7­
Allen,
M.
J.
1982
8­
Allen,
M.
J.
1982
9­
Feder
et
al
1974
emerges
at
night,
swims
upward
and
snaps
at
the
plankton
as
it
drifts
and
swims
through
its
midst3
sand
and
mud
bottoms;
bury
themselves
in
substrate
during
the
2
3
8
9
vertical
migrator,
moving
off
the
bottom
in
search
of
prey
at
night4,5
occurs
in
deep
water
in
the
north
and
south
of
range6
ambush
prey
from
a
partially
buried
position
on
the
bottom
and
drift
in
the
water
column
and
snap
up
prey
that
comes
near5
demersal,
living
primarily
on
soft
bottom
of
the
continental
shelf;
near
rocky
areas
during
spawning1
Adults
Food
habits
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CD
CE
CF
CG
CH
CI
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
harbor
seals1,7,8
NF
NF
NF
NF
NF
rock
cod4
California
gull
chicks5
western
gull,
and
Caspian
and
Fosters'
terns6
California
sea
lion9
1­
Olesiuk
et
al
1990
2­
Love
1996
3­
Fitch
and
Lavenberg
1971
4­
Hubbs
1920a
5­
Dierks
1990
6­
Harvey
1982
7­
Trumble
1995
8­
Torok
1994
9­
Hawes
1983
kelp
bass,
northern
elephant
seals,
California
sea
lions,

northern
fur
seals,
Pacific
white­
sided
dolphins,
Steller
sea
lions,
pygmy
sperm
whales,
harbor
porpoises,

Pacific
striped
dolphins,
cormorants,
western
gulls,

common
murres,
arctic
loons
and
pink
footed
shearwaters2
rockcod,
giant
seabass,
lingcod,
sea
lions
and
numerous
other
large
predators3
GENETICS
Nucleotide
diversity
(#)

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40
41
42
43
44
45
46
47
48
49
50
51
52
CJ
CK
CL
CM
CN
CO
CP
CQ
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Length
(
age)
Fishery
/
Length
(
age)
Sampling
gear
eing
structure
/
Meth
NF
NF
NF
NF
NF
NF
NF
NF
Mean
heterozygosity
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CR
CS
CT
CU
CV
CW
CX
CY
Mortality
Estimates
Rate
of
Ex
Method
of
Age
range
used
Rate
of
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)

NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CZ
DA
DB
DC
DD
xploitation
Exploitation
Interactions
/
associations
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
NF
NF
NF
male
midshipman
humming
attracts
females
2,5
photophores
may
play
a
role
in
mating
behavior6,8
possesses
venom
apparatus,
not
toxic
to
humans
5
possesses
venom
apparatus,
may
have
toxic
action
7
parasites:
5
families,
10
spp.
(
see
reference)
10
parasites:
digenea
(
Tubluovesicula
lindbergi)
11
1­
Demartini
1991b
1­
Love
1996
2­
Crane
1986
2­
Garrison
and
Miller
1982
3­
Ibara
et
al
1983
3­
Ibara
1967
4­
Harper
1999
4­
Hubbs
1920a
5­
Phillips
1973
6­
Ibara
1967
7­
McAllister
1968
8­
Crane
1965
9­
Allen,
M.
J.
1982
10­
Love
and
Moser
1983
11­
Arai
1969
P.
notatus
migrates
into
intertidal
zone
for
spawning
in
late
spring
and
early
summer3,4
males
migrate
inshore
in
winter
for
spawning,
females
follow
soon
after,
both
sexes
return
to
deep
water
in
fall1,2
disruptive
counterillumination
used
by
some
adults
as
an
antipredatory
strategy4,5
invertebrates
compete
with
plainfin
midshipman
embryos
for
attachment
space1
possible
functions
of
bioluminescence:
mimicry,
warning,
courtship,

and
counterillumination9
BEHAVIOR
Moveme
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30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
DE
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Homing
ability
(
Home
range)
Schooling
Territoriality
NF
NF
Yes(
males
defending
nest)
1
1­
Arora
1948
ent
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
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89
90
91
92
93
94
95
96
97
98
99
100
101
A
B
C
D
E
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71
72
73
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81
82
83
84
85
86
87
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89
90
91
92
93
94
95
96
97
98
99
100
101
F
G
H
I
J
K
L
M
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71
72
73
74
75
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
N
O
P
Q
R
S
T
U
V
W
X
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71
72
73
74
75
76
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78
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80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
Y
Z
AA
AB
AC
AD
AE
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71
72
73
74
75
76
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79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
AF
AG
AH
AI
AJ
AK
AL
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73
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
BA
BB
BC
BD
BE
BF
BG
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67
68
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71
72
73
74
75
76
77
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79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
BH
BI
BJ
BK
BL
BM
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81
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83
84
85
86
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89
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92
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94
95
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97
98
99
100
101
BN
BO
BP
BQ
BR
BS
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68
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71
72
73
74
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80
81
82
83
84
85
86
87
88
89
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92
93
94
95
96
97
98
99
100
101
BT
BU
BV
BW
BX
BY
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70
71
72
73
74
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78
79
80
81
82
83
84
85
86
87
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89
90
91
92
93
94
95
96
97
98
99
100
101
BZ
CA
CB
CC
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72
73
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80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CD
CE
CF
CG
CH
CI
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72
73
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80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CJ
CK
CL
CM
CN
CO
CP
CQ
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CR
CS
CT
CU
CV
CW
CX
CY
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84
85
86
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88
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91
92
93
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95
96
97
98
99
100
101
CZ
DA
DB
DC
DD
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71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
DE
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
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22
23
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27
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29
A
B
C
D
E
F
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Carangidae1
Seriola
lalandi
3
n.
B.
C.
to
Chile1
1surface
to
69
1Male:
W=
0.00000005689FL2.89
S.
dorsalis
1
1Female:
W=
0.00000007747FL2.84
S.
lalandei
2
1Both:
W=
0.00000007439FL2.85
1­
Robins
et
al.
1991
1­
Love
et
al
2000
1­
Gates
&
Frey
1974
1­
Love
et
al
2000
1­
Love
et
al
2000
1­
Craig
1960b
2­
Crooke
1983
2­
Robins
et
al.
1991
3­
Eschmeyer
1998
TAXONOMY
RANGE
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
yellowtail;
California
yellowtail;
amberjack;
amberfish;

white
salmon;
cavasina;
jack1,2
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23
24
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29
G
H
I
J
K
L
M
N
O
P
Q
R
S
T
U
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)
to
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
Female
Both
Male
Female
Both
NF
1
193
s.
California
(
FL)
1
NF
NF
12.91
NF
NF
0.136
NF
NF
­
1.9
NF
NF
NF
1­
Love
et
al
2000
1­
Smith
&
Mekjian
1960
AGE
&
GROWTH
r2
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20
21
22
23
24
25
26
27
28
29
V
W
X
Y
Z
AA
AB
AC
Age
composition
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
scales
tag
&
recapture
No
NF
1,2,3
12
NF
Dioecious1
NF
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
2­
Baxter
1960b
3­
Love
1996
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12
13
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16
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18
19
20
21
22
23
24
25
26
27
28
29
AD
AE
AF
AG
AH
AI
AJ
AK
AL
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Oviparous1
NF
NF
NF
1NG
(
NG)
1
to
2
(
50.6)
NG
(
NG)
NG
(
NG)
NG
2,3,4
NG
(
NG)
55.9
(
2)
NG
(
NG)
71.1
(
3)
NG
1­
Strategic
Assessment
Branch
1990
1­
Smith
&
Paul
1960
2­
Crooke
1992b
3­
Fitch
&
Lavenberg
1971
4­
Love
1996
Sexual
dimorphism
1st
maturity
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
50%
maturity
Breeding
Behavior
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
state
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.

NG
NG
NF
WA
to
CA2
NG
NG
s.
California1
California3,4,6,7
*****
*****

Mexico2
1­
Smith
&
Paul
1960
2­
Strategic
Assessment
Branch
1990
3­
Watson
et
al.
1996
4­
Fitch
&
Lavenberg
1971
5­
Kramer
&
Smith
1973
6­
Baxter
1960b
7­
Love
1996
REPRODUCTION
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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19
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21
22
23
24
25
26
27
28
29
BB
BC
BD
BE
BF
BG
BH
es
present
Spawning
Frequency
Size
specific
fecundity
equation
Dec.
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
Larval
duration
NF
NF
NF
NF
Multiple4
2450,000
(
3
yrs.)

3,4
450,000
(
NG)
to
700,000
(
NG)

1­
Smith
&
Paul
1960
1­
Smith
&
Paul
1960
2­
Strategic
Assessment
Branch
1990
2­
Fitch
&
Lavenberg
1971
3­
Crooke
1992b
3­
Baxter
1960b
4­
Baxter
1960b
4­
Love
1996
1,4
729,000
(
72.8
FL)
to
3,914,000
(
105.3
FL)
eggs
Fecundity
range
in
#
eggs
(@

cm,
grams,
or
age)

Mean
(
range)
fecundity
in
eggs/
g
female
wt.

Once
(
prolonged
over
a
3
month
period);

older
capable
of
mulitple1,2,3
DISPERS
Fecundity
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
BI
BJ
BK
BL
BM
BN
BO
BP
BQ
Size
at
settlement
Size
Fish
Size
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
(
mm)
Habitat
Foraging
guild
Foraging
method
NF
NF
1
0.8
to
1.6
planktonic1,2,3
NF
1,33.2
to
24.5
pelagic1
NF
NF
2
1.4
23
to
4
to
3
1.27
4
1.69
1­
Smith
&
Paul
1960
1­
Feder
et
al
1974
1­
Strategic
Assessment
Branch
1990
1­
Watson
et
al.
1996
2­
Watson
et
al.
1996
2­
Watson
et
al.
1996
2­
Watson
et
al.
1996
3­
Brownell
1979
3­
Limbaugh
1955
3­
Sumida
et
al.
1985
4­
Baxter
1960b
Food
habits
Eggs
Larvae
SAL
&
RECRUITMENT
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111
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­
Carangidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
BR
BS
BT
BU
BV
BW
BX
BY
Fish
Size
Food
habits
When
/
Frequency
Predators
(
SL
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
NF
NF
124.5
to
maturity
drifitng
kelp1
carnivore
NF
NF
Least
terns1
(
pelagic
crustacans;
juvenile
fish)
1
1­
Sumida
et
al.
1985
1­
Mitchell
&
Hunter
1970
1­
Strategic
Assessment
Branch
1990
1­
Love
1996
Juveniles
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Page
112
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Carangidae
1234
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
BZ
CA
CB
CC
Adults
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
carnivore
NF
diurnal1,2,3,4,5
1­
Feder
et
al
1974
1­
Feder
et
al
1974
1­
Strategic
Assessment
Branch
1990
2­
Strategic
Assessment
Branch
1990
2­
Strategic
Assessment
Branch
1990
2­
Crooke
1992b
3­
Carlisle
et
al.
1964
3­
Crooke
1983
3­
Crooke
1983
4­
Turner
et
al.
1969
4­
Carlisle
et
al.
1964
4­
Fitch
&
Lavenberg
1971
5­
Mitchell
&
Hunter
1970
5­
Craig
1960a
5­
Schmitt
&
Strand
1982
6­
Limbaugh
1955
6­
Fitch
&
Lavenberg
1971
7­
CA
Dept
of
Fish
&
Game
1982
7­
Schmitt
&
Strand
1982
8­
Love
1996
8­
Limbaugh
1955
9­
Love
1996
pelagic/
neritic;
kelp
beds;
rocky
peaks;
drifting
kelp;
artificial
reefs
1,2,3,4,5,6,7,8
(
squid;
small
schooling
fishes,
such
as
sardine,
mackerel,
and
anchovy;
pelagic
crabs
and
other
crustaceans)
1,2,3,4,5,6,7,8,9
Page
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
CD
CE
CF
CG
CH
CI
CJ
CK
CL
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
NF
NF
NF
NF
NF
NF
NF
1­
Strategic
Assessment
Branch
1990
1­
Crooke
1992b
2­
Fitch
&
Lavenberg
1971
2­
Crooke
1983
3­
Brian
2000
None
from
s.
CA
to
Baja
CA3
Mean
heterozygosity
Baja
California
(
south
of
Cedros
Is.)
and
s.

California
(
north
of
Cedrose
Is.)
1,2
Nucleotide
diversity
(#)

pinnipeds;
large
fish,
such
as
giant
sea
bass
1,2
GENETICS
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
CM
CN
CO
CP
CQ
CR
CS
CT
CU
CV
CW
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
First
Recruitment
Full
Recruitment
Method
of
Effective
population
size
Total
length
(
age)
Total
length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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115
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Carangidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
CX
CY
CZ
DA
DB
DC
DD
DE
Rate
of
Exploitation
ge
range
use
Rate
of
Exploitation
Interactions
/
associations
alculate
mort
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

NF
NF
NF
NF
sometimes
schools
with
S.
argentea
1,6
1to
404
miles
some
residents,
but
mostly
transient
1
NF
cleaned
by
Oxyjulis
californica
2,3
may
clean
parasites
from
blue
sharks4
1­
Strategic
Assessment
Branch
1990
1­
Baxter
&
Collyer
1960
1­
Baxter
&
Collyer
1960
2­
Hobson
1971
2­
Collyer
1954
3­
Turner
et
al.
1969
4­
Fitch
&
Lavenberg
1971
5­
Schmitt
&
Strand
1982
6­
Limbaugh
1955
7­
Hose
et
al.
1983
8­
Love
&
Moser
1983
infected
by
parasites
from
the
families
Dasyrhynchidae,
Caligidae,
Pennellidae,

Acanthocolpidae,
Bucephalidae,

Hemiuridae,
and
Heteraxinidae8
displays
intraspecific
cooperative
foraging5
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants7
BEHAVIOR
Movement
2up
to
240
miles;
but
most
<
10
miles
Page
116
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10
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
DF
DG
Schooling
Territoriality
Yes1,2,3,4
NF
1­
Feder
et
al
1974
2­
Strategic
Assessment
Branch
1990
3­
CDF
&
G
1982
4­
Love
1996
Page
117
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Carangidae
1234
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Clinidae1,2
Heterostichus
rostratus
1,2
giant
kelpfish
1intertidal
to
40
m
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Eschmeyer
and
Herald
1983
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
British
Columbia
to
Cabo
San
Lucas,
southern
Baja
California,
including
Isla
Guadalupe1
kelpfish,
eel,
kelp
blenny,
butterfish
1,2
TAXONOMY
RANGE
Page
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12
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
F
G
H
I
J
K
L
M
N
Total
Length
(
cm)
Length
conversion
equations
Max.
reported
Geographic
area
to
weight
(
g)
equation
length
(
cm)
(
length
measurment)
Male
Female
Both
Male
Female
1Both:
W=
0.00310456L3.243
1SL=
0.598+
0.819(
TL)
(
larvae)
161
cm
TL
NF
NF
NF
NF
NF
NF
1SL=­
0.580+
0.906(
TL)
(
juveniles
and
adults)

1­
Stepien
1986
1­
Stepien
1986
1­
Love
et
al
2000
Linf
in
cm
(
SE)
k
(
SE)

Page
119
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Clinidae
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17
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
O
P
Q
R
S
T
U
V
W
X
r2
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Growth
Parameters
to
(
SE)

AGE
and
GROWTH
Page
120
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12
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15
16
17
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
Y
Z
AA
AB
AC
AD
AE
Age
composition
Other
refs.
Longevity
(
Length)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
1
15
yrs
(
females)
1
dioecious
seasonal
monogamy1
oviparous1,2,3,5
male
guards
nest1,2,3,4,5
13
yrs
(
males)

25
yrs.

37
yrs.

1­
Stepien
1986
1­
Matarese
et
al
1980
1­
Stepien
1986
1­
Coyer
1982
1­
Matarese
et
al
1980
1­
Matarese
et
al
1980
2­
Stepien
1986
2­
Moser
1996a
2­
Moser
1996a
3­
Tenera
Environmental
Services
2000
3­
Love
1996
3­
Coyer
1982
4­
Tenera
Environmental
Services
2000
4­
Breder
and
Rosen
1966
5­
Coyer
1982
5­
Limbaugh
1955
6­
Holder
1907
Breeding
Behavior
Page
121
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1234
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
AF
AG
AH
AI
AJ
AK
AL
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
females
larger1
118.6
cm
(
1­
1.5
yrs.)
18.6
cm
(
1­
1.5
yrs.)
NG
NG
otolith/
whole
22
yrs.
2
yrs.
NG
NG
NG
1­
Stepien
1986
1­
Stepien
1986
1­
Stepien
1986
2­
Stepien
1986
2­
Tenera
Environmental
Services
2000
adult
males
and
females
exhibit
different
melanin
patterns1
1st
maturity
Total
length
in
cm
(
age
in
yrs)
at
REPRODU
50%
maturity
Sexual
dimorphism
adult
males
are
brown
(
infrequently
olive
green)
and
adult
females
exhibit
three
color
morphs;
red,
brown,
and
green1
Page
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10
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
Fertilized
eggs
present
Spawning
females
present
Both
reprod
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.

No
interreader
comparison
California5
NG
NG
northeast
Pacific1
CalCOFI2
*****
*****
*****

throughout
range
3
*****
*****
*****

Santa
Catalina
Island,
CA4
*****
*****
*****
*****
*****

1­
Matarese
et
al
1980
2­
Moser
1996a
3­
Stepien
1986
4­
Love
1996
UCTION
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
Page
123
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­
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BA
BB
BC
BD
BE
BF
BG
Fecundity
Mean
Fecundity
Size
specific
ductive
states
present
eggs/
g
Number
of
eggs
fecundity
eq
Nov.
Dec.
Spawning
Frequency
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
1,2,3400­
1,200
eggs
13,550
eggs
(
155
mm
TL
female)
NF
1,2,312­
17
days
2700
eggs
(
avg.)
214
days
(
avg.)

1­
Matarese
et
al
1980
1­
Matarese
et
al
1980
1­
Wang
1981
1­
Matarese
et
al
1980
2­
Stepien
1986
2­
Stepien
1986
2­
Stepien
1986
3­
Tenera
Environmental
Services
2000
3­
Love
1996
3­
Love
1996
may
spawn
several
times
a
year1,2,3
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BH
BI
BJ
BK
BL
BM
Size
at
settlement
Size
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
1approx.
8
week
NF
11.35
mm
(
avg.)
adhesive
eggs1,5,11
22
weeks
1,31.4
mm
(
avg.)

21.2­
1.4
mm
eggs
spawned
in
coastal
intertidal
to
30
m6
1­
Stepien
1986
1­
Stepien
1986
1­
Matarese
et
al
1980
1­
Matarese
et
al
1980
1­
Coyer
1982
2­
Love
1996
2­
Moser
1996a
2­
Moser
1996a
3­
Stepien
1986
3­
Stepien
1986
4­
Coyer
1982
5­
Love
1996
6­
Wang
1981
7­
Hubbs
1920b
8­
Breder
and
Rosen
1966
9­
Orton
1955
10­
Limbaugh
1955
11­
Tenera
Environmental
Services
2000
DISPERSAL
&
RECRUITMENT
Eggs
30­
50
mm
TL
(
fronds
of
giant
kelp,
brown
algae,
and
green
1
Gibbonsia
elegans,
Heterostichus
rostratus,
Semicossyphus
pulcher,
and
Oxyjulis
californica
1
eggs
are
found
on
sargassum,
ribbon
kelp,
giant
kelp,
and
surfgrass10
two
color
morphs
of
eggs;
purple
yolk
with
orange
oil
globules
and
yellow
yolk
and
yellow
oil
globules9
eggs
attached
in
algal
nest
and
brooded
by
male2,3,4,5,7,8
Page
125
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52
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54
BN
BO
BP
BQ
BR
BS
BT
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Fish
Size
15.5­
50.0
mm
SL
pelagic1,2
planktivore
NF
NF
NF
130­
50
mm
to
maturity
25.1­
45.0
mm
SL
rocky
reef
species3
near
kelp
beds
in
coastal
waters2
1­
Matarese
et
al
1980
1­
Matarese
et
al
1980
1­
Stepien
1986
1­
Stepien
1986
2­
Moser
1996a
2­
Moser
1996a
2­
Wang
1981
3­
Pondella
and
Stephens
1994
HA
(
single­
celled
algae,
rotifers,

mollusk
larvae,
and
barnacle
and
copepod
larvae)
1
Food
habits
Larvae
newly
hatched
larvae
have
large
yolk
sacs1
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17
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BU
BV
BW
BX
BY
BZ
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Habitat
NF
NF
NF
1tunas
and
lancetfish
among
kelp
beds
and
algae
of
coastal
waters3
inhabits
surf
grass
beds4
rocky
reef
species4
1­
Stepien
1986
1­
Moser
et
al
1986
1­
Moser
1996a
2­
Stepein
1987
2­
Stepein
1987
3­
Wang
1981
3­
Hubbs
1920b
4­
Pondella
and
Stephens
1994
4­
DeMartini
1981
5­
Hobson
et
al
1981
6­
Stepien
1986
7­
Quast
1968b
8­
Hubbs
1952
9­
Limbaugh
1955
10­
Holder
1907
BITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Food
habits
kelp
forests
and
rocky
bottom
with
macrophyte
cover1
fronds
of
giant
kelp,
the
brown
alga,

Sargassum
muticum,
and
green
surfgrass1
majority
of
juvenile
color
morphs
were
collected
in
plant
habitats
that
closely
matched
the
color
of
the
fish2
found
in
sand­
bottom
kelp
holdfast,

midkelp,
and
rocky­
bottom
kelp­
bed
habitat9
lies
almost
motionless
along
the
strands
of
kelp,
usually
at
about
a
45
°
angle,
with
the
head
up
or
down8
common
in
kelp
column
and
canopy,

also
about
laminarian
algae
and
eel
grass
on
the
bottom7
Juveniles
hovers
among
columns
of
giant
kelp
druing
both
day
and
night5
majority
of
adult
color
morphs
were
collected
in
plant
habitats
that
closely
matched
the
color
of
the
fish2,3,6,8,10
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38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CA
CB
CC
CD
CE
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
carnivores
NF
diurnal
feeders1,2
Brandt's
cormorant
and
least
terns1
1Santa
Catalina
Island
San
Diego
Punta
Clara
San
Benito
Islands
(
mysids,
isopods,
amphipods,
shrimp,
and
fish)
2
Guadalupe
Islands
(
small
crustaceans,
small
fish,
and
mollusks)
3,4
(
isopods
and
decapods
(
113­
283
mm)
5
1­
Love
1996
1­
Love
1996
1­
Love
1996
1­
Stepien
and
Rosenblatt
1991
2­
Quast
1968d
2­
Hobson
et
al
1981
2­
Hubbs
1952
3­
Limbaugh
1955
4­
Feder
et
al
1974
5­
Mitchell
1953
Food
habits
Adults
2subspecies
Heterosticus
rostratus
guadalupensis
(
from
Guadalupe
Island)

(
small
crustaceans;
such
as
mysids,
shrimps,
and
amphipods,
small
fish;
such
as
clingfish,
kelp
perch,
and
senorita)
1
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CF
CG
CH
CI
CJ
CK
CL
CM
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
NG
NG
1.25
+
0.07
25.00%
NG
0.07
+
0.02
allozymes
NG
1.30
+
0.09
25.00%
0.07
+
0.02
1.20
+
0.06
20.00%
0.07
+
0.03
1.23
+
0.07
22.50%
0.05
+
0.02
1.04
+
0.04
3.57%
0.01
+
0.01
Nucleotide
diversity
(#)
Mean
heterozygosity
GENETICS
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23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CN
CO
CP
CQ
CR
CS
CT
CU
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Fishery
/
Length
(
age)
Fishery
/
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)

NF
NF
NF
NF
NF
NF
NF
NF
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22
23
24
25
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CV
CW
CX
CY
CZ
DA
DB
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Interactions
/
associations
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
NF
NF
NF
NF
NF
NF
illicit
cleaning
behavior
from
senoritas
1
larvae
tend
to
school
with
mysid
shrimp
2
parasites:
5
families,
5
spp.
(
see
reference)
3
1­
Coyer
1982
2­
Love
1996
3­
Love
and
Moser
1983
MORTALITY
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
DC
DD
DE
DF
DG
DH
DI
DJ
DK
DL
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
NF
Residents1
NF
Yes
(
larvae)
1,2,3
Yes1,2,3,4,5
(
newly
recruited
juveniles)
2,3
adults
solitary3
1­
Tenera
Environmental
Services
2000
1­
Stepien
1986
1­
Stepien
1986
2­
Love
1996
2­
Limbaugh
1955
3­
Feder
et
al
1974
3­
Feder
et
al
1974
4­
Coyer
1982
5­
Holder
1907
BEHAVIOR
Movement
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17
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
DM
DN
DO
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
A
B
C
D
E
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67
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69
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71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
F
G
H
I
J
K
L
M
N
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
O
P
Q
R
S
T
U
V
W
X
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
Y
Z
AA
AB
AC
AD
AE
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
AF
AG
AH
AI
AJ
AK
AL
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BA
BB
BC
BD
BE
BF
BG
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BH
BI
BJ
BK
BL
BM
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BN
BO
BP
BQ
BR
BS
BT
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BU
BV
BW
BX
BY
BZ
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
CA
CB
CC
CD
CE
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
CF
CG
CH
CI
CJ
CK
CL
CM
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145
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
CN
CO
CP
CQ
CR
CS
CT
CU
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
CV
CW
CX
CY
CZ
DA
DB
Page
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
DC
DD
DE
DF
DG
DH
DI
DJ
DK
DL
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
DM
DN
DO
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30
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Cottidae
Hemilepidotus
hemilepidotus
2,3,4
red
Irish
lord
3,4,5
Sea
of
Okhotsk
to
Monterey
Bay,
California
1
Intertidal
to
51
m
(
156
ft)
1
[
spotted
Irish
lord,
bullhead,
red
sculpin]
3
Kamchatka
(
Russia)
to
Monterey
Bay,
CA
(
common
AK,
rare
CA)
2
San
Miquel
Is,
southern
CA
3
intertidal
to
275
m
(
908
ft)
4
Eastern
Kamchatka,
Russia
and
Commander
Is,
Bering
Sea
to
St
Paul
I,
Aleutian
Is4
Diablo
Cove,
central
CA
5
Cottidae
Leptocottus
armatus
2,3,4
Pacific
staghorn
sculpin
3,5
Chignik,
Alaska
to
San
Quintin
Bay,
Baja
California
1
Intertidal
to
98
m
(
300
ft)
1
[
bullhead,
staghorn
sculpin
4,
cabezon,

buffalo
sculpin,
smooth
cabezon]
3
S
Bering
Sea
to
northern­
central
Baja2
max
275
m
(
906
ft)
3
TAXONOMY
RANGE
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15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
F
G
H
I
J
KLM
AGE
and
GROWTH
Length
(
mm)
to
Max.
recorded
Geographic
Area
Linf
in
mm
(
SE)

weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
3
Male:
W
=
4.6759
x
10­
4TL
2.4063
Female:
W
=
7.3659
x
10­
6TL
3.1510
NF
1
51
cm
TL
(
20
in)
NF
NF
NF
NF
NF
4
W
=
0.000341
SL
2.896
2
FL
=
1.166(
SL)
+
1.549
SL
=
0.856(
FL)
­
1.116
(
range
56
­
178
mm
SL;
n=
10)
1
30.5
cm
TL
(
12
in)
3
San
Juan
Island,
WA
(
TL)
NG
NG
NG
NG
5
W
=
5.14
x
10­
3
TL
3.31
(
TL
=
cm)
2
TL
=
1.013(
FL)
­
0.083
FL
=
0.987(
TL)
+
0.093
(
range
67
­
213
mm
FL;
n=
10)
2
46
cm
TL
(
18
in)
4
San
Francisco
Bay,
CA
(
TL)
NG
NG
NG
NG
6
log
W
=
­
5.243
+
3.134
(
log
TL)

(<
190
mm
TL)
2
SL
=
0.846(
TL)
­
1.056
TL
=
1.181(
SL)
+
1.462
(
range
68
­
215
mm
TL;
n=
10)
5
Anaheim
Bay,
CA
(
SL)
NG
NG
NG
NG
8
W
=
3.9­
6
x
TL
3.2037
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19
20
21
22
23
24
25
26
27
28
29
30
N
O
P
Q
R
S
T
U
V
W
X
Y
Z
k
(
SE)
to
(
SE)

Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
Length)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
6
>
6
yr
NG
NG
NG
NG
NG
NG
NG
NG
3
otoliths/
whole
immersed
in
glycerin
(
n=
262)
No
No
NF
3
10
yr
(
228
mm
TL)

NG
NG
NG
NG
NG
NG
NG
NG
4
Length­
frequency
<
2yr;
Otoliths/
Whole
&

cleared
>
2yr
(
n=
146)
No
No
4
Male:
3
yr
(
193.5
mm
TL);

Female:
3
yr
(
197.3
mm
TL)

NG
NG
NG
NG
NG
NG
NG
NG
5
length­
frequency;
otoliths/
whole
(<
1yr)
No
No
5
NG
r2
Growth
Parameters
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12
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14
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17
18
19
20
21
22
23
24
25
26
27
28
29
30
AA
AB
AC
AD
AE
AF
Age
composition
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
NF
Dioecious
Male:
Promiscuous
and/
or
Polygynous
2
Oviparous
1
Both
sexes:
guard
nest3,4
Female:
larger
3
NF
Dioecious
NF
Oviparous
1
NF
NF
external
fertilization
Breeding
Behavior
Sexual
d
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
AG
AH
AI
AJ
AK
AL
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Male:
black
&
white
pelvic
fins
Female:
white
4
4
22.0
­
31.3
cm
TL
(
4­
6
yr)
4
28.7
­
38.5
cm
TL
(
4­
6
yr)
NG
NG
NG
NF
3
NG
(
SF
Bay,
CA)
3
12.8
­
15.3
cm
TL
(
1
yr;
SF
Bay,
CA)
NG
NG
Otoliths
/
whole
3
12.0
­
12.2
cm
TL
(
1
yr;
Walker
Creek,
CA)
3
12.0
­
12.2
cm
TL
(
1
yr;
Walker
Creek,
CA)
NG
NG
Otoliths
/
whole
50%
maturity
dimorphism
1st
maturity
Total
length
in
cm
(
age
in
yrs)
at
REPROD
Page
154
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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
Fertilized
eggs
present
Spawning
females
present
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.

NG
NG
NF
Puget
Sound
and
nearby
areas,
WA
7
No
No
NF
Walker
Creek
estuary
(
SF
Bay,
CA)
8
*****
*****

No
No
San
Pablo
Bay
(
SF
Bay,
CA)
9
*****
*****

San
Francisco
Bay
10
Anaheim
Bay
(
S
CA)
11
*****
*****

Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
s
DUCTION
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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
AY
AZ
BA
BB
BC
BD
BE
BF
Mean
Fecundity
Size
specific
Both
reproductive
states
present
eggs/
g
Number
of
eggs
fecundity
eq
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)

once
annually
4
NF
NF
3
Log
(
ova)
=
2.9145
log(
TL)
­
2.4311
(
range:
326
­
382
mm
TL)

once
annually
5
3
5000
eggs/
female
NF
4
#
maturing
eggs
=
0.355
TL
1.849
(
range:
~
135
­
235
mm
TL)

group­
synchronous
(
2­
4
clutches)
6
no
evidence
of
multiple
spawning6
pawning
records
Fecundity
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10
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
BG
BH
BI
BJ
BK
Size
at
settlement
Size
Egg
duration
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)

NF
NF
NF
NF
3
1.5
­
1.6
mm
5
1.5
­
1.7
mm
11
NG
8
NG
9
NG
12
NG
2,3
9­
14
days
at
15oC
NF
NF
NF
3
1.4
­
1.5
mm
(
avg
1.43
mm)

6
1.36
­
1.50
mm
DISPERSAL
&
RECRUITMENT
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
BL
BM
BN
BO
Habitat
Predators
Fish
Size
Habitat
demersal,
adhesive,
guarded
3,11
NF
2
~
5
­
20
mm
NL
pelagic
2
­
1
to
+
1
m
5
3
5
­
6
to
>
19
­
23
mm
SL
pelagic
3
egg
masses
adhered
to
substrate
(
rocks,

inverts)
5,11
7
22
mm
FL
(
n=
1)
pelagic
7
shallow
water
or
intertidal
zone
8
8
6.5
­
10.5
mm
pelagic
8
tough,
pink
conspicuous
masses,
often
above
low­
tide
mark
9
11
16.5
­
22
mm
FL
pelagic,
surface
11
shallow
water,
among
sessile
inverts
in
areas
of
high
currents
12
demersal
3,6,7,10
NF
2
~
4­
11
mm
TL
pelagic
2
adhesive
3,6,10
3
3.8­
5.0
mm
SL
(
hatching);
15­
20
mm
SL
(
transformation)
pelagic
2
4
4.5
mm
SL
(
hatching);
15­
20
mm
SL
(
metamorphosis)
NG
4
5
3­
11
mm
SL
tidal
embayment
&
seasonal
estuary
(
Elkhorn
Slough,

central
CA)
5
6
3.8­
5.0
mm
TL
(
at
hatching)
planktonic
in
polyhaline
to
mesohaline
areas
(
Sacramento­
San
Joaquin
estuary,
Moss
Landing
Harbor
and
Elkhorn
Slough,
CA)
6
9
~
3.9
­
20
mm
SL
shallow
water,
riverine,
estuarine,
marine
9
Eggs
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16
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19
20
21
22
23
24
25
26
27
28
29
30
BP
BQ
BR
BS
BT
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Fish
Size
NG
2
NF
NF
NF
NF
NG
3
copepods
7
NG
8
copepods,
amphipods
11
NG
2
NF
NF
NF
2
<
70
mm
SL
NG
3
2
>
70
mm
SL
NG
4
3
45­
79
mm
TL
NG
5
3
80­
119
mm
TL
NG
6
4
NG
planktivorous
9
5
66­
166
mm
TL
(
juv
&
adult)

6
3.3­
24.3
cm
TL
7
54­
140
mm
SL?
(
juv
&
adult)

7
59­
220
mm
SL?
(
juv
&
adult)

7
105­
290
mm
SL?
(
juv
&
adult)

8
107­
160
mm
TL
(
juv
&
adult)

9
99­
205
mm
SL
(
juv
&
adult)

Food
habits
Larvae
Page
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18
19
20
21
22
23
24
25
26
27
28
29
30
BU
BV
BW
BX
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
NF
NF
NF
NF
estuary
(
Anaheim
Bay,
southern
CA)
2
carnivore:
decapods,
amphipods,
fishes
2
NF
estuary
(
Anaheim
Bay,
southern
CA)
2
carnivore:
(
decapods,
fishes)
&
opportunistic
benthic
feeder
2
bay
(
Padilla
Bay,
WA)
3
carnivore:
amphipods,
isopods
3
bay
(
Padilla
Bay,
WA)
3
carnivore:
amphipods,
isopods,
crabs
3
estuary
(
Elkhorn
Slough,
central
CA)
4
epifaunal
crustaceans
(
amphipods,
copepods,
polychaetes)
4
marsh
(
China
Poot,
Alaska)
5
generalist:
amphipods,
diptera
larvae,
plants,
fish,
detritus,
insects,
crustacean,
oligochaetes
5
estuary
(
Yaquina
Bay,
OR)
6
NG
6
lagoon
(
Big
Lagoon,
Humboldt
Co,
CA)
7
amphipods
7
river
(
Mad
River,
Humboldt
Co,
CA)
7
algae,
shrimp,
amphipod,
Dungeness
crab
legs,
fish
7
bay
(
Humboldt
Bay,
Humboldt
Co,
CA)
7
Dungeness
crab,
fish,
amphipods,
polychaete,
isopods,
shrimp
7
intertidal
8
Dungeness
crab
8
bay/
estuary
(
San
Francisco
Bay,
CA)
9
shrimp,
crabs,
fish,
amphipods,
Callianassids,
bivalve
mollusks,

polychaetes,
isopods,
other
9
Food
habits
Life
Stage
Juveniles
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Page
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15
16
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18
19
20
21
22
23
24
25
26
27
28
29
30
BY
BZ
CA
Predators
Fish
Size
Habitat
NF
2
287­
385
mm
TL
(
breeding
females)
guarding
nests
2
2
220­
313
mm
TL
(
breeding
males)
guarding
nests
2
19
NG
Nanaimo,
B.
C.
19
gulls
and
cormorants
1
3
NG
estuary
(
SF
Bay,
northern
CA)
3
Great
Blue
Heron
(
Ardea
herodias
)
2
3
NG
estuary
(
Walker
estuary,
northern
CA)
3
Caspian
tern
(
Hydroprogne
caspia
)
3
4
 
120
mm
TL
bay
(
Padilla
Bay,
WA)
4
waterfowl,
ducks
4
5
NG
Mugu
Lagoon,
southern
CA
5
western
grebe
(
Aechmophorus
occidentalis
)
5
9
NG
estuary
(
Elkhorn
Slough,
central
CA)
9
mammals
(
harbor
seal,
CA
sea
lion,
N.
sea
lion)
6
6
66­
166
mm
TL
(
juv
&
adult)
marsh
(
China
Poot,
Alaska)
6
size
NG
(
juv
or
adult):
harbor
seal
(
Phoca
vitulina
richardsi
)
9
6
3.3­
24.3
cm
TL
estuary
(
Yaquina
Bay,
OR)
6
size
NG
(
juv
or
adult):
California
gull
(
Larus
californicus
)
10
7
54­
140
mm
SL?
(
juv
&
adult)
lagoon
(
Big
Lagoon,
Humboldt
Co,
CA)
7
Caspian
tern
(
Sterna
caspia
)
8,11
7
59­
220
mm
SL?
(
juv
&
adult)
river
(
Mad
River,
Humboldt
Co,
CA)
7
western
gull
(
Larus
occidentalis
)
11
7
105­
290
mm
SL?
(
juv
&
adult)
bay
(
Humboldt
Bay,
Humboldt
Co,
CA)
7
harbor
seal
(
Phoca
vitulina
richardsi
)
12
8
107­
160
mm
TL
(
juv
&
adult)
intertidal
8
10
>
70
mm
SL
tideflat
(
Coos
Bay,
OR)
10
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20
21
22
23
24
25
26
27
28
29
30
CB
CC
CD
CE
CF
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
Female
(
breeding):
carnivore
(
crabs,
shrimp,
fishes)
2
NF
Both
sexes:
suggested
nocturnal
1
NF
NF
Male
(
breeding):
carnivore
(
crabs,
shrimp,
fishes,

conspecific
eggs)
2
carnivore:
crabs
(
Hemigrapsus,
Cancer
productus
),

shrimps,
barnacles,
mussels
19
carnivore:
shrimps,
fishes
3
NF
NF
gulls
and
cormorants
1
NF
carnivore:
amphipods,
nereid
worm,
dipteran
larvae,

shrimp
3
Great
Blue
Heron
(
Ardea
herodias
)
2
carnivore:
isopods,
crabs,
fish
4
Caspian
tern
(
Hydroprogne
caspia
)
3
carnivore:
bivalve
siphons
5
waterfowl,
ducks
4
epifaunal
crustaceans
(
crabs,
amphipods,
macroalgae)

9
western
grebe
(
Aechmophorus
occidentalis
)
5
generalist:
amphipods,
diptera
larvae,
plants,
fish,
detritus,
insects,
crustacean,
oligocha
etes
6
mammals
(
harbor
seal,
CA
sea
lion,
N.
sea
lion)
6
NG
6
harbour
seal
7
amphipods
7
size
NG
(
juv
or
adult):
harbor
seal
(
Phoca
vitulina
richardsi
)
9
algae,
shrimp,
amphipod,
Dungeness
crab
legs,
fish
7
size
NG
(
juv
or
adult):
California
gull
(
Larus
californicus
)
10
Dungeness
crab,
fish,
amphipods,
polychaete,

isopods,
shrimp
7
caspian
tern
(
Sterna
caspia
),
western
gull
(
Larus
occidentalis
)
11
Dungeness
crab
8
harbor
seal
(
Phoca
vitulina
richardsi
)
12
burrowing
ghost
shrimp
(
Callianassa
californiensis
)
10
Food
habits
Adults
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24
25
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28
29
30
CG
CH
CI
CJ
CK
CL
CM
CN
CO
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
First
Recruitment
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Length
(
age)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Nucleotide
diversity
(#)
Mean
heterozygosity
GENETICS
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Cottidae
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
CP
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
DA
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
Rate
of
Exploitation
Full
Recruitment
Method
of
Age
range
used
Rate
of
Exploitation
Fishery
/
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
Page
164
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Cottidae
1234
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
DB
DC
DD
DE
DF
Interactions
/
associations
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

NF
Parasite:
mature
Trematode
(
Derogenes
varicus
);
Bering
Sea2
2
<
5
m2
to
500­
900
m2
NF
NF
Parasites:
9
Families;
~
13
Species;
See
Refs
11
Parasites:
Digenea
(
7
sp),
Nematoda
(
Ascarophis
sebastodis
),

Acanthocephala
(
Echinorhynchus
gadi
)
12
NF
Parasites:
nematode
Spirocamallanus
pererai
in
gastrointestinal
track
3
3
0­
800
m
(
Anaheim
Bay,
CA)
Resident:
Elkhorn
Slough,
central
CA
(
tidal
embayment
&
seasonal
estuary)
1
NF
Parasite:
nematode
larvae
Contracaecum
in
coelum
3
Resident:
Elkhorn
Slough,
central
CA
(
tidal
embayment
&
seasonal
estuary)
2
Parasite:
trematode
(
Gyrodactylus
sp)
on
gills
3
Resident:
Elkhorn
Slough,
central
CA
(
tidal
embayment
&
seasonal
estuary)
3
Parasites:
trematodes
(
Tubulovesicula
lindbergi
&
Stephenostomum
sp)

in
small
intestine
3
Parasite:
cestode
larvae
(
Acanthobothrium)
in
small
intestine
3
Parasite:
glossiphonid
leech
at
isthmus
3
Parasite:
cyclopoid
copepod
(
Holobomolochus
prolixus)
at
operculum
3
Parasite:
nematode
(
Contracaecum
magnus)
4
Parasite:
Trematode
(
Ascocotyle
diminuta
)
on
gills;
Pt
Mugu,
CA
5
Parasite:
Haemogregarina
leptocotti
in
blood;
Humboldt
Bay,
CA
6
Parasite:
acanthocephalan
(
Echinorhynchus
gadi
)
in
intestine;
Coos
Bay,

OR
8
Parasite:
acanthocephalans
(
Podocotyle
olssoni,
P.
reflexa
)
in
intestine;

Coos
Bay,
OR
8
Movement
BEHAVIOR
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Cottidae
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
DG
DH
Schooling
Territoriality
NF
NF
NF
NF
Page
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Cottidae
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
A
B
C
D
E
Cottidae
Scorpaenichthys
marmoratus
3,4
cabezon3,4,5;
giant
marbled
sculpin1,2,3
Sitka,
Alaska
to
Pt.
Abreojos,
central
Baja
California
1,2,3
Intertidal
to
82
m
(
250
ft)
1,2
(
Hemitripterus
marmoratus)
1,
2
[
bullhead,
blue
cod,
sculpin,
giant
sculpin,

marble
sculpin,
bull
cod,
salpa,
scorpion,

biggyhead,
scaleless
sculpin]
3
Max
@
110
m
(
335
ft)
3
[
cab,
caby,
cabezony,
bullhead]
6
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
F
G
H
I
J
KLM
1
W
=
1.85
x
10­
5
TL
2.9836
1
SL
=
0.83
TL
1
99
cm
TL
(
39
in)
1
Puget
Sound,
WA
(
TL)
690.25
(
NG)
740.87
(
NG)
NG
0.241
(
NG)

2
W
=
6.84
x
10­
6
TL
3.16
2
central
Cal,
S
Cal,
&
Baja
(
TL)
604
(
NG)
760
(
NG)
NG
0.83
(
NG)

7
Male:
W
=
2.546
x
10­
6
TL
3.307
Female:
W
=
8.780
x
10­
6
TL
3.113
Both:
W
=
5.498
x
10­
6
TL
3.185
1
O'Connell's
data
(
central
Cal,
S
Cal,
Baja)
recalculated
536
(
NG)
679
(
NG)
NG
0.457
(
NG)

Page
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Cottidae
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38
39
40
41
42
43
44
45
46
47
48
49
50
N
O
P
Q
R
S
T
U
V
W
X
Y
Z
0.354
(
NG)
NG
­
1.23
(
NG)
0.84
(
NG)
NG
NG
NG
NG
1
Otoliths/
whole;
fin
rays/
sectioned
(
n=
87)
No
between
structures
NF
1
Male:
17
yr
(
648
mm
TL)

Female:
16
yr
(
725
mm
TL)

0.91
(
NG)
NG
NG
NG
NG
NG
NG
NG
2
otoliths/
whole
(
n=
105)
No
intrareader
2
Male:
9+
yr
(
531
mm
TL)

Female:
13+
yr
(
720
mm
TL)

0.230
(
NG)
NG
NG
NG
NG
NG
NG
NG
1
otoliths/
whole
(
n=
105)
No
NG
Page
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38
39
40
41
42
43
44
45
46
47
48
49
50
AA
AB
AC
AD
AE
AF
NF
Dioecious
Polygynous
1
Oviparous
1
Male:
guard
nest1,2
Female:
larger,
older
1
tend
eggs
laid
on
Iridaea
splendens
5
Female:
larger
2
Page
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Cottidae
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34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AG
AH
AI
AJ
AK
AL
Juveniles:
silver­
red
1
1
34.3
cm
TL
(
2
yr)
1
44.5
cm
TL
(
3
yr)
NG
NG
NG
Male:
(
adult):
red
Female:
(
adult):
green
1,2
2
~
33.9
cm
TL
(
2+
yr)
2
~
44.5
cm
TL
(
3+
yr)
NG
NG
Otoliths
/
whole
Both
sexes:
(
Non­
breeding
adults):

white1,2
Male:
(
near
spawning
females):

brown,
red,
black
Male:

(
nesting):
white
Female
(
adult):
turquoise­
green
3
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Cottidae
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34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
NG
NG
NF
British
Columbia
1
No
between
structures
British
Columbia
2
southern
Strait
of
Georgia,
B.
C.
3
Puget
Sound,
WA
4,5
*****
*****

central
CA
4,5
central
CA
6
*****

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Cottidae
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37
38
39
40
41
42
43
44
45
46
47
48
49
50
AY
AZ
BA
BB
BC
BD
BE
BF
batch
spawner1,3
1
26.4
­
14.5
per
batch
(
2.5
­
10.5
kg
TW)
NF
1
#
eggs
=
0.29
TL
­
87.54
(
range:
500
­
775
mm
TL)

at
least
twice
during
spawning
season2
2
34.8
­
21.0
per
batch
(
1.4
­
4.6
kg
TW)
2
Y
=
27.3
+
1.53X
(
range:
1.4
­
4.6
kg
TW;
Y
=

1000'
s
of
eggs;
X
=
100'
s
of
g
TW)

*****
Page
173
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Cottidae
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BG
BH
BI
BJ
BK
1
2­
3
weeks
Nov
­
Mar
1
1
40
mm
TL
(~
3­
4
months),
tidepools
NF
1
1.4
­
1.7
mm
3­
4
months
3
2
<
0.40
­
1.5
mm
3
1.4
­
1.9
mm
4
1.5
mm
5
1.5
­
1.6
mm
7
NG
13
nests:
46
cm
diam
(
5
­
10
cm
thick)

14
NG
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36
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38
39
40
41
42
43
44
45
46
47
48
49
50
BL
BM
BN
BO
NG
1
eggs
toxic
to
man
1
1
10­
40
mm
TL
planktonic
1
NG
2
eggs
toxic
to
lab
animals
1,3,5,6,7
2
4­
35
mm
SL
(
hatch
at
4­
6
mm)
pelagic
(
Oregon
<
65
km
offshore)
2
demersal,
adhesive,
attached
eggs
3
eggs
avoided
by
natural
predators
2
3
NG
pelagic
3
hard
surfaces
(
rock,
iron,
steel,
wood)
15
Artedius
harringtoni,
Embiotoca
lateralis,

Damalichthys
vacca,
other
male
cabezon
4
6
hatch:
3.1­
4.8
mm
TL
NG
6
5
NG
10
8
­
17
mm
FL
pelagic,
surface
10
demersal,
adhesive,
clusters
7
11
14
­
22
mm
FL
pelagic,
surface
11
intertidal,
under
rocks,
deep
crevices,
subtidal
13
12
3
­
6
mm
SL
(
hatching);
14
mm
SL
(
transformation)
pelagic
until
~
35
mm
12
algae
(
Iridaea
splendens
)
14
13
6
mm
TL
(
hatching)
NG
13
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Cottidae
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36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BP
BQ
BR
BS
BT
10,11
60­
118
mm
TL
(
0+)

12
~
26­
31.4
mm
TL
(
juv)

13
45­
245
mm
TL
(
juv
&
adult)

13
45­
79
mm
TL
(
juv)

13
80­
119
mm
TL
(
juv)

14
NG
(
juvenile
or
adult)

15
20­
220
mm
TL
(
juv
&
adult)

16
<
100
mm
TL
18
33­
290
mm
TL
(
juv
&
adult)

copepods
(
crustaceans)
1
NF
NF
NF
1
41­
200
mm
TL
(
arbitrary:
juvs)

NG
2
1
201­
400
mm
TL
(
arbitrary:
pre­
adults)

NG
3
17
200­
499
mm
SL
(
juv
&
adult
)

NG
6
19
NG
barnacle
larvae
(
Cirripedia:
Balanus
cypris
stage),

copepod
(
nauplii,
Copepodites,
Pseudocalanus
minutus
),
amphipod
(
Parathemisto
pacifica
),

decapod
(
Brachyura
zoea),
Euphausiacea
(
furcilia),

fish
(
Atheresthes
stomias
)
10
copepods,
fish
(
Stichaeidae)
11
NG
12
NG
13
Page
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Cottidae
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40
41
42
43
44
45
46
47
48
49
50
BU
BV
BW
BX
estuary
(
Grays
Harbor,
WA)
10,11
carnivore:
nereid
polychaetes,
ghost
shrimp,
sand
shrimps,
juv
crabs,
bivalve
siphon
tips,
unid
crustaceans
10,11
bay
(
Skagit
Bay,
Puget
Sound,
WA)
12
carnivore:
crustaceans,
polychaetes,
insects
12
bay
(
Padilla
Bay,
WA)
13
generalist:
amphipod,
isopods,
crabs,
polychaetes,
moluscs,

fishe,
algae,
wood,
rock,
shrimp
13
bay
(
Padilla
Bay,
WA)
13
carnivore:
amphipods,
isopods,
polychaetes,
molluscs,
other
13
bay
(
Padilla
Bay,
WA)
13
carnivore:
amphipods,
isopods,
crabs,
polychaetes,
other
13
estuary
(
west
of
Vancouver
Island,
B.
C.,
Canada)
14
amphipods,
insects
(
Chironamid
larvae
&
pupae),
Oligochaetes,

isopods
(
Gnorimosphaeroma
bregonensis
),
eggs,
calanoid
copepods
14
bay,
cove,
strait
(
Strait
of
Juan
de
Fuca)
15
generalist:
decapods
(
Natantia,
Reptantia),
mysids,
amphipods,

fish
(
incl
Pleuronectidae,
Cottidae),
polychaetes,
isopods
15
estuaries
(
Vancouver
Island,
B.
C.
Canada)
16
generalist:
amphipods
(
Eogammarus
confervicolus
,
Corophium
sp
),
polychaetes
(
Nereis
sp
),
isopod
(
Gnorimosphaeroma
oregonensis
),
chum
salmon
fry
(
Oncorhynchus
keta
)
16
bay
(
Everett
Bay,
WA)
18
carnivore:
amphipods
(
Corophium
,
gammarids),
fish
(
juv
staghorn
sculpin,
sandlance,
starry
flounder,
threespine
stickleback),
ghost
shrimp
(
Callianassa
)
18
demersal;
hard
bottoms;
tidepools
1
carnivore:
Crustacea,
fish
1
NF
NF
demersal;
hard
bottoms;
tidepools
1
carnivore:
Crustacea,
Mollusca,
fish
1
S
Cal
rocky
inshore
and
kep
beds
17
cephalopods
(
octopus,
squid),
crabs
(
cancroid,
spider),

crustaceans,
shrimp,
fish,
algae
(
red
&
green),
gastropods,

lamellibranchs
17
Diablo
Cove,
CA
19
fish
(
incl
juv
rockfish),
crustaceans,
mollusks,
Octopus
sp,
see
ref
for
more
ID
19
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­
Cottidae
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32
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34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BY
BZ
CA
11
99­
205
mm
SL
(
juv
&
adult)
bay/
estuary
(
San
Francisco
Bay,
CA)
11
12,13
 
118
mm
TL
(
1+
and
older)
estuary
(
Grays
Harbor,
WA)
12,13
14
45­
245
mm
TL
(
juv
&
adult)
bay
(
Padilla
Bay,
WA)
14
14
>
120
mm
TL
(
adult)
bay
(
Padilla
Bay,
WA)
14
15
NG
(
juvenile
or
adult)
estuary
(
west
of
Vancouver
Island,
B.
C.,
Canada)
15
16
20­
220
mm
TL
(
juv
&
adult)
bay,
cove,
strait
(
Strait
of
Juan
de
Fuca)
16
18
33­
290
mm
TL
(
juv
&
adult)
bay
(
Everett
Bay,
WA)
18
sea
otters,
pigeon
guillemots,
least
terns,
Brandt's
cormorants
7
1
401­
720
mm
TL
(
arbitrary
)
demersal;
hard
bottoms1
1
 
488
mm
tidepools
1
19
NG
Nanaimo,
B.
C.
19
20
NG
(
Adult)
Kelp
bed
bottom
20
17
200­
499
mm
SL
(
juv
&
adult
)
S
Cal
rocky
inshore
and
kep
beds
17
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36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CB
CC
CD
CE
CF
shrimp,
crabs,
fish,
amphipods,
Callianassids,
bivalve
mollusks,
polychaetes,
isopods,
other
11
carnivore:
ghost
shrimp,
nereid
polychaete,
mud
shrimp,
juv
crabs,
gammarid
amphipod,
sand
shrimp,

fish
12,13
generalist:
amphipod,
isopods,
crabs,
polychaetes,

moluscs,
fishe,
algae,
wood,
rock,
shrimp
14
generalist:
crabs,
isopods,
fishes,
algae,
wood,
rock
14
amphipods,
insects
(
Chironamid
larvae
&
pupae),

Oligochaetes,
isopods
(
Gnorimosphaeroma
bregonensis
),
eggs,
calanoid
copepods
15
generalist:
decapods
(
Natantia,
Reptantia),
mysids,

amphipods,
fish
(
incl
Pleuronectidae,
Cottidae),

polychaetes,
isopods
16
carnivore:
amphipods
(
Corophium
,
gammarids),
fish
(
juv
staghorn
sculpin,
sandlance,
starry
flounder,

threespine
stickleback),
ghost
shrimp
(
Callianassa
)
18
carnivore:
Crustacea,
Mollusca,
fish,
eggs
1
NF
NF
sea
otters,
pigeon
guillemots,
least
terns,
Brandt's
cormorants
8
NF
NG
1
carnivore:
fish
(
Hemilepidotus
hemilepidotus,
fish
eggs,
small
perch),
crabs
(
Epialtus
,
Cancer
,

Hemigrapsus
),
shrimp
19
mesocarnivore
20
cephalopods
(
octopus,
squid),
crabs
(
cancroid,

spider),
crustaceans,
shrimp,
fish,
algae
(
red
&

green),
gastropods,
lamellibranchs
17
Page
179
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1167
5­
2220e
­
Cottidae
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CG
CH
CI
CJ
CK
CL
CM
CN
CO
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
180
of
1167
5­
2220e
­
Cottidae
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CP
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
DA
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
181
of
1167
5­
2220e
­
Cottidae
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
DB
DC
DD
DE
DF
Parasite:
trematode
(
Tubulovesicula
sp
)
in
stomach,
intestine,
gill
region;

Coos
Bay,
OR
8
Parasite:
nematodes
(
Contracaecum
sp,
Anisakis
sp
)
in
intestine
and
body
musculature;
Coos
Bay
OR
8
Parasite:
trematode
(
Derogenes
varicus,
Genolinea
manteri
)
in
stomach
9
Parasites:
21
Families;
~
43
Species;
See
Refs
11
Parasites:
Digenea
(
5
sp),
Nematoda
(
2
sp),
Acanthocephala
(
2
sp)
12
NF
Parasite:
Nematode
(
Capillaria
margolisi
)
in
pyloric
caeca1
1
0
naut
mi
NF
NF
Parasite:
Protozoa,
Sarcomastigophora
(
Trypanoplasma
beckeri
)
in
blood
and
body
fluids
7
Parasite
and
vector
for
T.
beckeri
:
Leech,
Hirudinea,
Piscicolidae
(
Malmiana
diminuta
)
7
Parasite:
trematode
(
Genolinea
robusta
)
in
stomach
9
Parasite:
copepod
(
Chondracanthus
gracilis
)
in
gill
cavity
10
Parasite:
copepod
(
Chondracanthus
pinguis
)
10
Parasites:
11
Families;
~
18
Species;
See
Refs
11
Page
182
of
1167
5­
2220e
­
Cottidae
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
DG
DH
NF
NF
Page
183
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
A
B
C
D
E
1
­
Lea
and
Eschmeyer
1988
1
­
Clemens
and
Wilby
1961
1
­
Miller
and
Lea
1972
1
­
Miller
and
Lea
1972
2
­
Eschmeyer
1998
2
­
Pillsbury
1957
2
­
Eschmeyer
and
Herald
1983
2
­
Eschmeyer
and
Herald
1983
3
­
Miller
and
Lea
1972
3
­
Gates
and
Frey
1974
3
­
Love
et
al.
2000
3
­
Love
et
al.
2000
4
­
Robins
et
al.
1991
4
­
Miller
and
Lea
1972
4
­
Allen
and
Smith
1988
4
­
Allen
and
Smith
1988
5
­
Robins
et
al.
1991
5
­
Burge
and
Schultz
1973
6
­
Reilly
et
al.
1994
Page
184
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
F
G
H
I
J
KLM
1
­
O'Connell
1953
1
­
O'Connell
1953
1
­
Miller
and
Lea
1972
1
­
Lauth
1987
2
­
Lauth
1987
2
­
Karpov
and
Kwiecien
1988
2
­
Hart
1973
2
­
O'Connell
1953
3
­
DeMartini
and
Patten
1979
3
­
Weiss
1969
4
­
Tasto
1975
4
­
Jones
1962
5
­
Bayer
1985
5
­
Tasto
1975
6
­
Weiss
1969
7
­
Lea
et
al.
1999
8
­
Armstrong
1991
Page
185
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
N
O
P
Q
R
S
T
U
V
W
X
Y
Z
1
­
O'Connell
1953
1
­
Lauth
1987
2
­
Lauth
1987
2
­
O'Connell
1953
3
­
Weiss
1969
3
­
Weiss
1969
4
­
Jones
1962
4
­
Jones
1962
5
­
Tasto
1975
5
­
Tasto
1975
6
­
Love
1996
Page
186
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
AA
AB
AC
AD
AE
AF
1
­
Lauth
1987
1
­
Matarese
et
al.
1989
1
­
Feder
et
al.
1974
1
­
O'Connell
1953
2
­
DeMartini
and
Patten
1979
2
­
Lauth
1987
2
­
Lauth
1987
3
­
DeMartini
and
Patten
1979
3
­
DeMartini
and
Patten
1979
4
­
Garrison
and
Miller
1982
5
­
Burge
and
Schultz
1973
Page
187
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
AG
AH
AI
AJ
AK
AL
1
­
O'Connell
1953
1
­
Fitch
and
Lavenberg
1971
1
­
Fitch
and
Lavenberg
1971
1
­
Fitch
and
Lavenberg
1971
2
­
Orton
1955
2
­
O'Connell
1953
2
­
O'Connell
1953
2
­
O'Connell
1953
3
­
Lauth
1987
3
­
Jones
1962
3
­
Jones
1962
3
­
Jones
1962
4
­
DeMartini
and
Patten
1979
4
­
Garrison
and
Miller
1982
4
­
Garrison
and
Miller
1982
Page
188
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
1
­
Fitch
and
Lavenberg
1971
1
­
Fitch
and
Lavenberg
1971
1
­
Hart
1973
2
­
O'Connell
1953
2
­
O'Connell
1953
2
­
Fraser
1923
3
­
Jones
1962
3
­
Jones
1962
3
­
Pillsbury
1957
4
­
Lauth
1987
5
­
Lauth
1988
6
­
O'Connell
1953
7
­
DeMartini
and
Patten
1979
8
­
Jones
1962
9
­
DeVlaming
et
al.
1984
10
­
Boothe
1967
11
­
Tasto
1975
Page
189
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
AY
AZ
BA
BB
BC
BD
BE
BF
1
­
O'Connell
1953
1
­
Lauth
1988
(
TW
=
total
weight)
1
­
Lauth
1988
2
­
Lauth
1987
2
­
O'Connell
1953
(
TW
=
total
weight)
2
­
O'Connell
1953
(
TW
=
total
weight)

3
­
Lauth
1988
3
­
Jones
1962
3
­
DeMartini
and
Patten
1979
4
­
DeMartini
and
Patten
1979
4
­
Jones
1962
5
­
Jones
1962
6
­
DeVlaming
et
al.
1984
Page
190
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
BG
BH
BI
BJ
BK
16
NG
1
­
O'Connell
1953
1
­
O'Connell
1953
1
­
O'Connell
1953
1
­
O'Connell
1953
2
­
Jones
1962
2
­
Forrester
1969a
2
­
Lauth
1988
3
­
Matarese
et
al.
1989
3
­
Tenera
Environmental
Services
2000
3
­
Matarese
et
al.
1989
4
­
Barnhart
1932
5
­
Demartini
and
Patten
1979
6
­
Jones
1962
7
­
Yoklavich
et
al.
1992
8
­
Hart
1973
9
­
Clemens
and
Wilby
1961
10
­
Wang
1981
11
­
Gorbunova
1966
12
­
Garrison
and
Miller
1982
13
­
Feder
et
al.
1974
14
­
Burge
and
Schultz
1973
15
­
Lauth
1987
16
­
Marliave
1975a
Page
191
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
BL
BM
BN
BO
cluster
of
egg
masses
on
moderately
exposed
bedrock,
4
m
depth
16
14
6­
9
mm
FL
surface
waters,
Saanich
Inlet,
Vancouver
Isl.
14
1
­
O'Connell
1953
1
­
Hubbs
and
Wick
1951
1
­
O'Connell
1953
1
­
O'Connell
1953
2
­
Lauth
1988
2
­
Pillsbury
1957
2
­
Richardson
and
Washington
1980
2
­
Richardson
and
Washington
1980
3
­
Matarese
et
al.
1989
3
­
Burns
1988
3
­
Matarese
et
al.
1989
3
­
Matarese
et
al.
1989
4
­
Barnhart
1932
4
­
Lauth
1987
4
­
Jones
1962
4
­
Jones
1962
5
­
Demartini
and
Patten
1979
5
­
Fuhrman
et
al.
1969
5
­
Yoklavich
et
al.
1992
5
­
Yoklavich
et
al.
1992
6
­
Jones
1962
6
­
Fuhrman
et
al.
1970
6
­
Wang
1981
6
­
Wang
1981
7
­
Yoklavich
et
al.
1992
7
­
McAllister
1968
7
­
Barraclough
1967a
7
­
Barraclough
1967a
8
­
Hart
1973
8
­
Gorbunova
1966
8
­
Gorbunova
1966
9
­
Clemens
and
Wilby
1961
9
­
Emmett
et
al.
1991
9
­
Emmett
et
al.
1991
10
­
Wang
1981
10
­
Barraclough
and
Fulton
1968
10
­
Barraclough
and
Fulton
1968
11
­
Gorbunova
1966
11
­
Barraclough
1967b
11
­
Barraclough
1967b
12
­
Garrison
and
Miller
1982
12
­
Ambrose
1996
12
­
Ambrose
1996
13
­
Feder
et
al.
1974
13
­
Marliave
1975a
13
­
Marliave
1975a
14
­
Burge
and
Schultz
1973
14
­
Barraclough
et
al.
1968
14
­
Barraclough
et
al.
1968
15
­
Lauth
1987
16
­
Marliave
1975a
Page
192
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
BP
BQ
BR
BS
BT
copepod
nauplii
14
1
­
O'Connell
1953
1
­
Cass
et
al.
1990
1
­
O'Connell
1953
2
­
Richardson
and
Washington
1980
2
­
Tasto
1975
3
­
Matarese
et
al.
1989
3
­
Dinnel
et
al.
1990
4
­
Jones
1962
4
­
Barry
et
al.
1996
5
­
Yoklavich
et
al.
1992
5
­
Wolf
et
al.
1983
6
­
Wang
1981
6
­
Bayer
1985
7
­
Barraclough
1967a
7
­
Porter
1964
8
­
Gorbunova
1966
8
­
Fernandez
et
al.
1993
9
­
Emmett
et
al.
1991
9
­
Boothe
1967
10
­
Barraclough
and
Fulton
1968
10
­
Armstrong
1991
11
­
Barraclough
1967b
11
­
Armstrong
et
al.
1995
12
­
Ambrose
1996
12
­
Smith
1980
13
­
Marliave
1975a
13
­
Dinnel
et
al.
1990
14
­
Barraclough
et
al.
1968
14
­
Birtwell
et
al.
1984
15
­
Thornburgh
1978
16
­
Mace
1983
17
­
Quast
1968d
18
­
Conley
1977
Page
193
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
BU
BV
BW
BX
1
­
O'Connell
1953
1
­
O'Connell
1953
2
­
Tasto
1975
2
­
Tasto
1975
3
­
Dinnel
et
al.
1990
3
­
Dinnel
et
al.
1990
4
­
Barry
et
al.
1996
4
­
Barry
et
al.
1996
5
­
Wolf
et
al.
1983
5
­
Wolf
et
al.
1983
6
­
Bayer
1985
6
­
Bayer
1985
7
­
Porter
1964
7
­
Porter
1964
8
­
Fernandez
et
al.
1993
8
­
Fernandez
et
al.
1993
9
­
Boothe
1967
9
­
Boothe
1967
10
­
Armstrong
1991
10
­
Armstrong
1991
11
­
Armstrong
et
al.
1995
11
­
Armstrong
et
al.
1995
12
­
Smith
1980
12
­
Smith
1980
13
­
Dinnel
et
al.
1990
13
­
Dinnel
et
al.
1990
14
­
Birtwell
et
al.
1984
14
­
Birtwell
et
al.
1984
15
­
Thornburgh
1978
15
­
Thornburgh
1978
16
­
Mace
1983
16
­
Mace
1983
17
­
Quast
1968d
17
­
Quast
1968d
18
­
Conley
1977
18
­
Conley
1977
Page
194
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
BY
BZ
CA
1
­
Girard
1858
1
­
O'Connell
1953
1
­
O'Connell
1953
2
­
MacGinitie
1935
2
­
DeMartini
and
Patten
1979
2
­
DeMartini
and
Patten
1979
3
­
Miller
1943
3
­
Jones
1962
3
­
Jones
1962
4
­
Clemens
and
Wilby
1961
4
­
Dinnel
et
al.
1990
4
­
Dinnel
et
al.
1990
5
­
Palmer
1962
5
­
Peterson
and
Quammen
1982
5
­
Peterson
and
Quammen
1982
6
­
Treacy
1985
6
­
Wolf
et
al.
1983
6
­
Wolf
et
al.
1983
7
­
Love
1996
7
­
Porter
1964
7
­
Porter
1964
8
­
Parkin
1998
8
­
Fernandez
et
al.
1993
8
­
Fernandez
et
al.
1993
9
­
Trumble
1995
9
­
Barry
et
al.
1996
9
­
Barry
et
al.
1996
10
­
Dierks
1990
10
­
Posey
1986
10
­
Posey
1986
11
­
Harvey
1982
11
­
Boothe
1967
11
­
Boothe
1967
12
­
Torok
1994
12
­
Armstrong
1991
12
­
Armstrong
1991
13
­
Armstrong
et
al.
1995
13
­
Armstrong
et
al.
1995
14
­
Dinnel
et
al.
1990
14
­
Dinnel
et
al.
1990
15
­
Birtwell
et
al.
1984
15
­
Birtwell
et
al.
1984
16
­
Thornburgh
1978
16
­
Thornburgh
1978
17
­
Quast
1968d
17
­
Quast
1968d
18
­
Conley
1977
18
­
Conley
1977
19
­
Fraser
1923
19
­
Fraser
1923
20
­
Quast
1968b
20
­
Quast
1968b
Page
195
of
1167
5­
2220e
­
Cottidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
CB
CC
CD
CE
CF
1
­
O'Connell
1953
1
­
DeMartini
and
Patten
1979
1
­
Girard
1858
2
­
DeMartini
and
Patten
1979
2
­
MacGinitie
1935
3
­
Jones
1962
3
­
Miller
1943
4
­
Dinnel
et
al.
1990
4
­
Clemens
and
Wilby
1961
5
­
Peterson
and
Quammen
1982
5
­
Palmer
1962
6
­
Wolf
et
al.
1983
6
­
Treacy
1985
7
­
Porter
1964
7
­
Olesiuk
et
al.
1990
8
­
Fernandez
et
al.
1993
8
­
Love
1996
9
­
Barry
et
al.
1996
9
­
Trumble
1995
10
­
Posey
1986
10
­
Dierks
1990
11
­
Boothe
1967
11
­
Harvey
1982
12
­
Armstrong
1991
12
­
Torok
1994
13
­
Armstrong
et
al.
1995
14
­
Dinnel
et
al.
1990
15
­
Birtwell
et
al.
1984
16
­
Thornburgh
1978
17
­
Quast
1968d
18
­
Conley
1977
19
­
Fraser
1923
20
­
Quast
1968b
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196
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Cottidae
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52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
CG
CH
CI
CJ
CK
CL
CM
CN
CO
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Cottidae
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52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
CP
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
DA
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52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
DB
DC
DD
DE
DF
1
­
Moravec
and
McDonald
1981
1
­
Lea
et
al.
1999
1
­
Yoklavich
et
al.
1991
2
­
Kruse
1977
2
­
DeMartini
and
Patten
1979
2
­
Yoklavich
et
al.
1992
3
­
Tasto
1975
3
­
Tasto
1975
3
­
Barry
et
al.
1996
4
­
Smedley
1934
5
­
Armitage
1997
6
­
Hill
and
Hendrickson
1991
7
­
Burreson
1979
8
­
Dunlap
1951
9
­
Lloyd
1938
10
­
Fraser
1919
11
­
Love
and
Moser
1983
12
­
Arai
1969
Page
199
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­
Cottidae
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52
53
54
55
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
DG
DH
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9
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11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Triakidae
2
Triakis
semifasciata
1,2
leopard
shark
Oregon
to
Baja
and
northern
Gulf
of
California
1,2,3,4
1,2
3.7­
91
m
catshark,
tiger
shark
1,2,3
Triakidae
2
Mustelus
henlei
1,2
brown
smoothhound
Oregon
(
Coos
Bay)
to
Gulf
of
California,
Ecuador
and
2
inshore
to
335
m
mud
shark,
dogfish,
paloma,
Peru
1,3,4
sand
shark,
Henle's
shark
1,3
Triakidae
2
Mustelus
califoriensis
1,2
grey
smoothhound
northern
California
(
Cape
Mendocino)
to
Mexico
2
inshore­
67
m
mud
shark,
dogfish,
paloma,
(
Mazatlan),
including
Gulf
of
California
1,2,3,4
sand
shark,
gray
shark
1,3
Triakidae
2
Galeorhinus
galeus
2
soupfin
shark
temperate
waters
nearly
worldwide;
2
nearshore
to
471m
school
shark,
oil
shark,
soupfin,
northern
British
Columbia
to
Gulf
of
California;

tope
shark
1,3,4,5,9
Peru
and
Chile
1,3,4,5
Squalidae
1,2
Squalus
acanthias
1,2
spiny
dogfish
circumglobal;
Sea
of
Japan
to
Bering
Sea
and
to
2
intertidal­
1,236
m
spur
dog,
rock
salmon,
dogshark
central
Baja
California
and
Gulf
of
California
1,3,4,5
picked
dogfish,
pinback
shark,
tiburon
de
espina,
kaad,
aburatsunozame1,3,4,5,10
TAXONOMY
RANGE
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22
23
24
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
F
G
H
I
J
Total
Length
(
cm)
Max.
reported
Geographic
Area
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
1
Males:
W=
0.000009772L2.9806
(
W=
lbs.)
NF
1,2,3
2.1
m
central
California(
TL)
1
149.9
Females:
W=
0.000005754L3.1044
(
W=
lbs.)

NF
NF
1,2,3,4
95cm
central
California
2,13
97.6(
195)

(
95%
CI)

2
Males:
log
W
=
5.20
­
10
+
2.88
logL
NF
1,2,3
1.6
m
central
California
2,13
142.4(
185)

Females:
log
W
=
4.02
­
10
+
3.19
logL
(
L=
mm)
(
95%
CI)

Males:
W=
0.000002673L3.26954
NF
1,2,3,5
2
m
NF
NF
Females:
W=
0.000000032L4.15605
3
Females:
loge
(
weight
+
1)=
0.0119TL
­
0.2676;
TL<
50
cm(
r2=.
91)
NF
2,3,4,5
160
cm
mid­
Atlantic
and
s.
New
England
3
82.49
loge
(
weight
+
1)=
0.0293TL
­
1.1714;
TL>
50
cm
(
r2=.
96)

3Males:
loge
(
weight
+
1)=
0.00114TL
­
0.2532;
TL<
50
cm(
r2=.
92)
Strait
of
Georgia
4
97.3
loge
(
weight
+
1)=
0.0235TL
­
0.8535;
TL>
50
cm
(
r2=.
91)
Strait
of
Georgia
5
NG
north
and
west
Scottish
coast
6
79.7
4
Full
term
females:
W=
0.00000017
L3.47
(
mm)
British
Columbia14
99.8
4Barren
Females:
W=
0.00000305L3.03
(
mm)

4Males:
W=
0.00000189L3.09
(
mm)

11Both:
W=
0.00267L3.783
(
W=
kg.)

11Males:
W=
0.00292L3.0641
(
W=
kg.)

11Females:
0.00185L3.7012
(
W=
kg.)
L
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24
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
K
L
M
N
O
P
Q
R
S
T
U
r2
Female
Both
Male
Female
Both
Male
Female
Both
Male
Female
Both
160.2
153.6
0.089
0.073
0.082
­
2.03
­
2.74
­
2.31
NG
NG
NG
86.1(
68)
97.7(
64)
0.225(
0.196)
0.285(
0.059)
0.224(
0.05)
­
1.375(
0.925)
­
1.086(
0.688)
­
1.296(
0.49)
NG
NG
NG
101.8(
165)
154.4(
245)
0.218(
0.177)
0.35(
0.063)
0.168(
0.058)
­
1.032(
0.42)
­
1.002(
0.296)
0.339
NG
NG
NG
­
1.271
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
100.5
NG
0.1481
0.1057
NG
­
2.67
­
2.9
NG
NG
NG
NG
128.5
NG
0.07
0.036
NG
­
4.5
­
6.9
NG
NG
NG
NG
114.94
NG
NG
0.0437
NG
NG
­
3.557
NG
NG
NG
NG
101.4
NG
0.21
0.11
NG
­
2
­
3.6
NG
NG
NG
NG
125.3
NG
0.07
0.048
NG
­
4.7
­
4.88
NG
NG
NG
NG
AGE
and
GROWTH
to
(
SE)
Growth
Parameters
Linf
in
cm
(
SE)
k
(
SE)
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
V
W
X
Y
Z
AA
AB
Age
composition
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
Length)
information
Sexuality
vertebral
centra/
silver
nitrate
tetracycline
mark­
recapture
betw.
reader
comparison
1,10,11
1
9­
10
(
120
cm)
NF
dioecious
1
stain,
thin
sectioning
technique
edge
analysis
1
females
may
live
to13­
16
yrs.

2,3
24
yrs.
(
130
cm)

1230
yrs.
(
200
cm
TL)

vertebral
centra;
sectioned
and
edge
analysis
betw.
reader
comparison
NF
3
13
yrs.
NF
dioecious
x­
rayed
7
13
yrs.
(
100
cm)

vertebral
centra;
sectioned
and
edge
analysis
betw.
reader
comparison
NF
3
9
yrs.
NF
dioecious
x­
rayed
7
9
yrs(
125
cm)

NF
NF
NF
NF
NF
NF
dioecious
dorsal
fin
spines;
surface
reading
No
interreader
comparison
2,3,4,5,13,14,15(
juveniles),
16
4
25­
30
yrs.
NF
dioecious
5
40
yrs.

x­
ray
spectrometry
No
compared
to
dorsal
spines
1160
+
yrs.

dorsal
fin
spine
No
No
dorsal
fin
spine;
surface
reading
No
interreader
comparison
dorsal
fin
spine;
surface
reading
No
interreader
comparison
12females:
13
yrs
(
100
cm
TL),

males:
7
yrs
(
90
cm
TL)

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30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AC
AD
AE
AF
AG
AH
AI
AJ
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
promiscuous
aplacental
viviparity
1,2,3
No
females
grow
larger6
males
have
claspers
1
NG
118
cm
NG
2
NG
104
cm
NG
3
100
cm
(
7
yrs)
105
cm
(
10
yrs,)
NG
4
NG
110­
129
cm
(
12­
14
yrs)
NG
2970­
119
cm(
7
yrs)
100­
129
cm(
10
yrs)
NG
32NG
100­
129
cm(
11­
21
yrs)
NG
promiscuous
viviparous
1,2,4,5
No
females
grow
larger
1,6,7
males
have
claspers
2
NG
67
cm
NG
yolksac
placenta
15
5,23,32
52­
66
cm.(
3
yrs)
51­
63
cm.(
2­
3
yrs)
NG
6
61
cm
61
cm
NG
promiscuous
viviparous
1,2,4,5
No
females
grow
larger
7
males
have
claspers
2
NG
87.5
cm
NG
6,23
58­
64
cm
(
1­
2
yrs)
70
cm
(
2­
3
yrs)
NG
promiscuous
aplacental
viviparity1,2,4,5,6,7,18
No
NF
males
have
claspers
7
135
cm
150
cm
approx.
140
cm
6,24
122­
171
cm
(
8
yrs)
131­
186
cm(
11yrs)
NG
8
140
cm
155
cm
NG
31approx.
152
cm
>
152
cm
NG
promiscuous
aplacental
viviparity
1,2,4,5,8,9,16
No
females
grow
larger1,8
males
have
claspers
NG9
76
cm
NG
10
72
cm(
15
yrs)
76
cm
(
18
yrs)
78.5cm
(
19
yrs)

11NG
80
cm
(
24
yrs)
NG
8NG
NG
72
cm
28NG(
14
yrs)
93.5
cm
(
23
yrs)
NG
30NG
NG
72.3
cm
759
cm
72.8
NG
REPRODUCTION
Breeding
Behavior
Sexual
dimorphism
1st
maturity
50%
m
Total
length
in
cm
(
age
in
yrs)
at
Page
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16
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AK
AL
AM
AN
AO
AP
AQ
AR
Fertilization
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.

NG
NG
NG
NG
NF
Elkhorn
Slough,
CA
1
NG
NG
NG
NG
San
Diego,
CA
3
NG
vertebral
centra/
silver
nitrate
tetracycline
mark­
recapture
betw.
reader
comparison
west
coast
4
stain,
thin
sectioning
technique
and
edge
analysis
Pacific
coast
2
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NF
Elkhorn
Slough,
CA
1
NG
vertebral
centra;
sectioned
and
edge
analysis
betw.
reader
comparison
and
central
California
12
x­
rayed
Pacfic
coast
2
NG
NG
NG
NG
NG
NG
NG
NG
1
central
California
12
NG
vertebral
centra;
sectioned
and
edge
analysis
betw.
reader
comparison
and
Anaheim
Bay,
CA
6
x­
rayed
approx.
155cm
NG
NG
NG
2
Santa
Catalina
Island,
CA
14
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
93.5
cm
NG
NG
NG
1,2,3,5
Strait
of
Georgia,
B.
C.
8
93.5
cm
(
29)
used
age
and
growth
estimates
No
No
San
Diego,
CA
3
of
Jones
and
Geen
1977
Pacific
coast
2
93.4
cm
(
35.5)
dorsal
spines/
NA
No
No
Unkown
location17
93.5
cm
NG
NG
NG
NG
dorsal
spines/
surface
reading
No
interreader
comparison
94.2
cm
NG
NG
NG
NG
NG
NG
NG
Fertilization
and
Spawning
maturity
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18
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
Mean
Fecundity
Both
Parturition
eggs/
g
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
(
weight
range)

annual
breeding
cycle1,5
1,2
7­
36
offspring
3
6­
24
offspring
*****
*****
4
12
offspring
(
average)

^^^^^
^^^^^
5,6,284­
29
offspring
7
4­
33
offspring
313­
11
young
single
spawner
3,7
1­
8
embryos
22
1­
10
embryos
5,28
3­
5
young
per
litter
311­
8
young
single
spawner
3,8
3­
16
embryos
22
3­
15
embryos
7
2­
5
offspring
single
spawner
23
16­
54
fertilized
eggs
2,7,9,30
6­
52
offspring
2,9,10
average
of
35
young
*****
^^^^^
females:
biannual
spawners
11
4­
11
eggs
males:
annual
spawners
1
12
2­
16
embryos
10
3­
14
offspring
13
4­
32
offspring
5,7,9
1­
20
offspring
8
20­
30
pups
262­
20
pups,
8
pups
(
avg.)

291­
25
embryos
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
27North
Pacific:
2­
17
pups,
6­
7
(
avg.)
,
Sea
of
Japan:
3­
25
pups,
12
pups(
avg.),

northwest
Atlantic:
1­
9
pups,
4
pups(
avg.),

northeast
Atlantic:
1­
10
pups,
3­
5
pups(
avg.)

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37
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39
40
41
42
43
44
45
46
47
48
49
50
51
BE
BF
BG
BH
Fecundity
Size
specific
Number
of
eggs
fecundity
eq
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Gestation
period/
egg
duration
Larval
duration
1
6
embryos
(
121
cm)
NF
1,26approx.
12
month
gestation
NA
2
approx.
15
embryos
(
123­
125.9
cm)
2
10­
12
month
gestation
approx.
16
embryos(
126­
128.9
cm)

approx.
19
embryos(
129­
134.9
cm)

approx.
22
embryos(
135­
143
cm)

approx.
30
embryos(
144­
146.9
cm)

approx.
32
embryos(
147­
149.9
cm)

3
6
embryos
(
64.6
cm)
NF
NF
NA
3
10
embryos
(
86
cm)

NF
NF
3,4
10­
month
gestation
NA
5
10­
11
month
gestation
NF
NF
5,6,7,20,2512
month
gestation
NA
NF
1
Eggs:
loge
F=
3.04
loge
TL­
11.81;
r2=
0.40
3,8,9
approx.
2
yr.
gestation
NA
Candled
embryos:
loge
F=
3.33
loge
TL­
13.22;
r2=
0.36
10
18­
24
month
gestation
Large
embryos:
loge
F=
3.80
loge
TL­
15.49;
r2=
0.40
9
22­
24
month
gestation
2
y=
0.27x­
18.25;
y=
brood
size,
x=
length
11
12
month
gestation
2220
months
2421
months
DISPERSAL
&
R
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BI
BJ
BK
BL
BM
BN
Eggs
Size
at
settlement
Size
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
Fish
Size
NA
NF
NA
NA
NA
NA
NA
NF
NA
NA
NA
NA
NA
NF
NA
NA
NA
NA
NA
NF
NA
NA
NA
NA
NA
NF
NA
NA
NA
NA
RECRUITMENT
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16
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BO
BP
BQ
BR
BS
BT
BU
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Fish
Size
Habitat
NA
NA
NA
NA
NA
1,2,3,4
18­
20
cm
to
maturity
pupped
in
estuaries3
NA
NA
NA
NA
NA
2,3
19
cm
to
maturity
common
in
enclosed,
shallow,
muddy
bays
9
20
cm
to
maturity
NA
NA
NA
NA
NA
9
20
cm
to
maturity
bottom
dwelling
in
shallow
muddy
bays
2
NA
NA
NA
NA
NA
3
36
cm
to
maturity
176
found
in
shallow
waters
4
NA
NA
NA
NA
NA
5
approx.
24­
76
cm
shallow
bays
and
estuaries
2
1120
cm.­
maturity
(
approx.
60
cm.)

Food
habits
Larvae
sandy
or
muddy
flats,
cobble
bottoms,

rocky
reef,
and
kelp
beds1,2,3
remain
in
bays
and
estuaries
up
to
2
yrs.

then
move
offshore2
sloughs
and
estuaries
serve
as
a
nursery
ground12
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18
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BV
BW
BX
BY
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Foraging
guild
Foraging
method
When
/
Frequency
Predators
carnivores
opportunistic
feeder
on
bottom
dwellers
1,2
NF
larger
sharks,
ie.
sixgill
and
sevengill
1
(
crabs)
1,2,3,4,5,6,7
may
disturb
mud
to
capture
food
3
may
use
suction
to
capture
prey
4
carnivore
(
crabs)
3,8
feeder
on
bottom
dwellers,
may
disturb
NF
NF
mud
to
capture
food
3
pursuit
and
capture
over
mudflats
5
carnivore
9
may
disturb
mud
or
suck
organisms
to
no
diel
pattern
1
NF
(
crabs)
8
capture
food
5
(
crabs,
clams,
fish,
polychaetes,
echinorinid)
10
pursuit
and
capture
over
mudflats
5
(
small
decapods)
11
carnivore
(
fish)
12
NF
NF
NF
(
invertebrates
and
fish)
7
carnivore
(
bony
fish,
cartilageonous
fish,
invertebrates)
3
NF
NF
pups
eaten
by
lincod,
sablefish,
white
shark
planktivores
13
(
euphasiids,
herring,
ctenophore,
and
other
fish)
11
(
planktonic
invertebrates,
such
as
euphausiids
and
ctenophores)
20
(
invertebrates,
such
as
euphausiids,
ctenophores,
and
shrimp)
21
(
herring,
young
salmon,
sticklebacks,
and
sand
lance)
22
Food
habits
Juveniles
continuous,
or
intermittent
feeding6
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BZ
CA
CB
Habitat
Foraging
guild
Foraging
method
sandy
or
muddy
flats,
cobble
bottoms,
rocky
reef,
and
kelp
beds
1,2,3
benthic
carnivore1,2,3,43
opportunistic
feeder
on
bottom
dwellers
1
found
in
polyhaline
to
euryhaline
waters3,13
(
fish,
crabs,
fish
eggs,
innkeeper
worms)
1,4
may
disturb
mud
to
capture
food
2
kelp
bed
and
adjacent
kelpless
cobble
area
4
may
use
suction
to
capture
prey
3
(
crabs,
isopods,
shrimp,
worms,
fish,
clams,
and
fish
eggs)
8
found
in
shallow
bays
and
coves,
often
close
over
cobble
bottom
16
(
mobile
grapsid
crabs)
45
(
crabs,
shrimp,
clam
siphons,
fish,
aand
small
sharks
and
rays)
54
common
in
enclosed,
shallow,
muddy
bays
2
feeder
on
bottom
dwellers,

found
in
polyhaline
to
euryhaline
waters13
may
disturb
mud
to
capture
food
2
(
crabs,
isopods,
shrimp,
worms,
fish,
clams,
and
fish
eggs)
8
pursuit
and
capture
over
mudflats
4
(
small
crabs
and
shrimp)
40,54
(
small
crustaceans
and
fish)
43,54
bottom
dwelling
in
shallow
muddy
bays
2
carnivore
(
crabs,
ghost
shrimp,
innkeeper
worm,
and
sm.
fish)
4,5
may
disturb
mud
or
suck
(
primarily
crabs)
10,48
organisms
to
capture
food
4
(
clams,
large
decapods)
11
pursuit
and
capture
over
mudflats
4
coastal­
pelagic
species
of
temperate
waters,
found
in
opportunistic
carnivore
(
pelagic
and
bottom
fish,
squid)
12,13,14,53
feeds
on
bottom
and
in
water
column
18
shallow
bays
and
in
submarine
canyons
2
(
fish,
chondrichthyans,
invertebrates)
15
feeds
in
midwater19
males
and
females
segregate
by
sex,
males
in
deeper
waters
mainly
piscivores
(
fishes,
squid,
cephalopods)
44
and
females
inshore
5
boreal
to
warm­
temperate,
inshore
and
offshore,
and
opportunistic
carnivore
(
bony
fish
and
invertebrates)
5,16
primarily
bottom
feeder,
but
from
surface
to
bottom
2
(
crustaceans
and
small
fish)
14,17
also
rise
to
the
surface
to
feed
5
(
fish,
shrimp,
crabs)
8
forages
throughout
the
water
column19
(
fish,
cephalopods,
crustaceans)
18
(
ratfish,
herring,
krill)
37,41
(
shrimp,
herring,
flatfish,
sand
lance,
squid,
and
octopus)
37
(
euphasiid,
herring,
sandlance,
ctenophore,
and
other
fish)
39
(
herring,
euphausiids,
jellyfish,
and
octopus)
46
Food
habits
Adults
(
Clupea
sp
.,
Scomber
scomber
,
Gadus
sp
.,
Pleuronectes
sp
.,

Trigla
sp
.,
Callionymus
lyra
,
and
Raia
sp
.)
47
(
invertebrates,
crabs,
clam
siphons,
polychaetes,
innkeeper
worms,
octupi,
and
fish)
3,5,6,7
mainly
piscivores
(
fish,
such
as
herring,
northern
anchovy,
lincod,

and
Pacific
hake,
crustaceans,
and
eupausiids)
44
carnivore
(
crabs,
shrimp,
isopods,
squid,
polychaete
worms,

tunicates,
and
sm.
fish)
4,5,6,9,10
(
krill
crab,
shrimp,
octopus,
squid,
clam,
snail,
worm,
jellyfish,

anemone,
and
plants)
42
found
in
sand­
bottom
kelp
holdfast,
midkelp,
and
rocky­
bottom
kelpbed
habitat16
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18
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CC
CD
CE
CF
CG
CH
Genetic
varia
Haplotype
diversity
Mean
number
of
When
/
Frequency
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)

NF
no
major
predators
except
man
1
NF
NF
NF
NF
possibly
other
shark
species
2
NF
sevengill
and
leopard
sharks
3
NF
NF
NF
NF
no
diel
pattern
1
NF
NF
NF
NF
NF
NF
spotted
sevengill
shark,
the
NF
NF
NF
NF
great
white
shark,
marine
mammals,
and
man
2
NF
larger
sharks,
bluefin
tuna,
and
large
bony
fishNF
NF
NF
NF
six­
gill
shark
5
larger
sharks,
such
as
the
porbeagle
shark,

blue
shark,
and
tiger
shark
6
GENETICS
Nucleotide
diversity
(#)

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14
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CI
CJ
CK
CL
CM
CN
CO
CP
RECRUITMENT
INTO
FISHERY
nce
%
Polymorphic
First
Recruitment
Full
Recruitment
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Length
(
age)
Fishery
/
Length
(
age)
Sampling
gear
NF
NF
NF
NF
1
4­
13
yrs.
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
S
Mean
heterozygosity
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CQ
CR
CS
CT
CU
CV
CW
CX
Y
Mortality
Estimates
Method
of
Age
range
used
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
NG
NG
NG
1
M=
0.28
(
first
yr.
of
life)
0.038­
0.163
NG
Hoenig's
equation
NG
1
M=
0.14
(
after
first
yr.)

2
M=
0.15
NG
NG
Hoenig's
equation
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
3
M=
0.094(
instantaneous)
NG
NG
age­
structure
model
NG
4
M=
0.10
202
NG
NG
NG
NG
MORTALITY
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CY
CZ
DA
DB
DC
Rate
of
Exploitation
Rate
of
Exploitation
Interactions
/
associations
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
NG
NG
NF
Observed
hunting
anchovies
with
piked
dogfish
5
parasites:
13
families,
20
spp.
(
see
reference)
10
NG
NG
12
Tracking:
overall
movement
rate:
8.1+
0.5
m
min­
1
12
movements
correlated
with
tides
NF
NF
NF
parasites:
8
families,
15
spp.
(
see
reference)
10
2,15160
km
in
3
months
NF
NF
NF
parasites:
8
families,
9
spp.
(
see
reference)
10
NF
NF
NF
NF
parasites:
7
families,
9
spp.
(
see
reference)
10
NG
NG
sablefish,
Pacific
cod,
soupfin
5
move
offshore
in
winter
shark,
and
sea
lions
1
and
inshore
in
summer
NG
NG
parasites:
39
families,
75
spp.
(
see
reference)
10
6
>
7,000
km
2
long
range
movements
>
1,600
km
1
tag
recoveries
indicate
that
most
leopard
sharks
are
residents
of
San
Francisco
Bay,
although
some
move
out
of
bay
in
fall
and
winter
BEHAVIOR
3,4
approx.
1,000
miles
northward
(
at
large
2yrs.
1
3/
4
months)

14majority
of
tag
recoveries
made
in
the
vicinity
of
release;
movement
from
20­
1,200
nautical
miles
(
1­
7
yrs)

found
in
association
with
hake,
pollock,
salmon,
herring,
Leuroglossus
,

myctophid,
and
several
other
species
of
midwater
fish
7
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16
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
DD
DE
DF
DG
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
resident,
limited
local
movement
1
sensitive
to
electrical
fields
in
Yes
1,2
NF
water,
may
be
used
in
orientation
or
locating
food
2
NF
NF
NF
NF
northern
movement
in
summer
2
NF
Yes
5,6
NF
Transient
1,2,3
NF
Yes
2,4,5,6,7,8
NF
seasonal
migration
4
school
segregated
by
sex9,10
forms
schools
w/
grey
and
brown
smoothhounds3,4,11
tend
to
move
out
of
bays
and
estuaries
in
winter15
Movement
inhabits
inshore
estuarine
waters
during
spring
and
moves
offshore
during
winter
months1,15
reported
to
migrate
from
southern
to
central
California
in
the
summer1
seasonal
migration
assoc.
with
reproductive
cycles5
Yes
,
schools
often
composed
of
one
sex4
Yes,
forms
schools
with
leopard
sharks2,4
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Elasmobranchs
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
A
B
C
D
E
Squatinidae
1,2
Squatina
californica
1,2
pacific
angel
shark
Unverified
report
from
southeastern
Alaska.
Probably
1
3­
46
m
California
angel
shark,
angel
at
least
Puget
Sound,
Washington
to
Bahia
Magdelena,

shark,
monkfish,
northern
angel
southern
Baja
California
and
Gulf
of
California;
Panama
shark,
squat,
squato
1,3
to
Chile
1,2,3,4
Heterodontidae
1,2
Heterodontus
francisi
1,2
horn
shark
central
California
to
Gulf
of
California,
probably
1,2
intertidal
to
150
m
Port
Jackson
shark,
bullhead
shark
Ecuador
and
Peru
1,2,3,4
horned
shark,
California
horn
shark
1,3
2,
5
Scyliorhinidae
1,2
Cephaloscyllium
ventriosum
1,2
swell
shark
central
California
(
Monterey
Bay)
to
Aculpulco
2
5­
457
m
California
swell
shark,
balloon
shark,
Mexico,
also
Chile
1,2,3,4,5
puffer
shark
1,3,5
Hexanchidae
1,2
Hexanchus
griseus
1,2
sixgill
shark
circumglobal
in
temperate
seas;
southern
Japan,
1,2
surface
to
2,000
m,

cowshark,
shovelnosed
shark,
Aleutian
Is.,
Alaska
to
the
southern
tip
of
Baja
1,3,4
adults
usually
below
91
m
mud
shark,
sixgill
cowshark,
young
in
deeper
parts
of
griset,
caffa
bota,
bluntnose
sixgill,
bays,
adults
in
deep
requin
griset
1,3,4,5
water
Hexanchidae
1,2
Notorynchus
cepideanus
1,2
sevengill
shark
circumglobal
in
temperate
waters;
southern
Japan,
2
1­
46
m
cowshark,
mud
shark,
spotted
northern
British
Columbia
to
southern
Baja
California
1,3
shallow
bays
cowshark,
sevengill
cowshark,
and
in
the
Gulf
of
California
1,3,4
Pacific
sevengill
cowshark,
broadnose
sevengill
shark,
pratnez,
canabota,

gata
1,3,4,5
Rhinobatidae
1,2
Rhinobatos
productus
1,2
shovelnose
guitarfish
San
Francisco
to
Gulf
of
California,
1
nearshore­
15
m
shovelnose
shark,
pointed
nose
rare
north
of
Monterey
1,3
guitarfish,
guitarfish
1,3
Dasyatididae
Urolophus
halleri
1,2
round
stingray
n.
California
(
Eureka)
to
Panama,
including
Gulf
of
1,2
1­
21
m
Urolophidae
Urobatis
halleri
2
stingaree,
stingray
1,3
California
1,3,4,5
Page
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Elasmobranchs
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63
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
F
G
H
I
J
5
Males:
y=
1.33
­
0.00831x
+
0.0000152x2
(
r2=
0.96)
NF
1,2,3,4
1.52
m
Pacific
Coast
7
NG
Females:
y=
2.095
­
0.0115x
+
0.0000182x2
(
r2=
0.98)
Pacific
Coast
7
NG
NF
NF
3,4
122
cm
NF
NF
NF
NF
2,3,4
1
m
NF
NF
6
Females:
y=­
37.5
+
(
6.64x10­
2)(
x1)+(­
3.11x10­
5)(
x1
2)+(
1X10­
8)(
x1
3)
NF
3
5+
m
NF
NF
(
polynomial
regression)
1
4.7
m
6
4.82
m
6
Males:
y=­
3.39
+
(
1.23X10­
2)(
x1)+(­
1.58x10­
5)(
x1
2)+(
1.10X10­
8)(
x1
3)
NF
1
2.7
m
NF
NF
Females:
y=­
59
+
(
0.14)(
x1)+(­
1.1x10­
4)(
x1
2)+(
3.08X10­
8)(
x1
3)
7
2.96
m
(
polynomial
regression)

NF
NF
2
1.7
m
Long
Beach,
California(
TL)
8
142
NF
ratio
of
DW
to
TL
is
1:
1.75
1,2,3
56
cm
NF
NF
Page
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Elasmobranchs
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
K
L
M
N
O
P
Q
R
S
T
U
NG
133.8
NG
NG
0.151
NG
NG
­
1.309
NG
NG
NG
NG
120.7
NG
NG
0.523
NG
NG
­
0.424
NG
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
594
228
0.095
0.016
0.047
­
3.942
­
3.8
­
4.03
0.7
0.94
0.81
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
220
of
1167
5­
2220e
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Elasmobranchs
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53
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
V
W
X
Y
Z
AA
AB
tag
and
recapture
No
No
1,6,7
NF
NF
dioecious
tag
and
recapture
No
No
NF
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
NF
dioecious
vertebral
centra;
whole
terramycin
marked,
held
in
intrareader
comparison
and
NF
3
10
yrs.
NF
dioecious
captivity
and
edge
11
yrs.
(
125
cm)

analysis
NF
NF
NF
12
8
yrs.
NF
dioecious
Page
221
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1167
5­
2220e
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Elasmobranchs
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53
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
AC
AD
AE
AF
AG
AH
AI
AJ
promiscuous
aplacental
viviparity
4,9
No
NF
males
have
claspers
12
90­
100
cm
TL
90­
100
cm
TL
NG
5
75­
80
cm
86­>
108
cm
NG
13
102­
107
cm
100­
109
cm
NG
promiscuous
oviparous
1,2,4,5
No
NF
males
have
claspers
5
58­
71
cm
>
58
cm
NG
6
61
cm
61
cm
NG
promiscuous
oviparous
2,4,5,9
No
NF
males
have
claspers
NF
NF
NF
promiscuous
aplacental
viviparity
2,5,9,10
No
NF
males
have
claspers
14
309­
331
cm
NG
NG
15,16
421
cm
NG
NG
6
335
cm
427
cm
NG
promiscuous
aplacental
viviparity
2,9,11
No
NF
males
have
claspers
15,17
132
cm
NG
NG
18
140
cm
218
cm
NG
promiscuous
aplacental
viviparity
4
No
NF
males
have
claspers
NG
2
110
cm
TL
NG
19
90­
100
cm(
8
yrs)
99
cm(
7
yrs)
NG
promiscuous
aplacental
viviparity
4
No
NF
male
dentition
1
27
14.6
cm
DW(
2.6
yrs)
14.5
cm
DW
(
2.6
yrs)
NG
males
have
claspers
and
siphon
sac
5
Page
222
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2220e
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Elasmobranchs
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
AK
AL
AM
AN
AO
AP
AQ
AR
NG
NG
NG
NG
3
Pacific
coast
2,15
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NF
San
Diego,
CA
3
NG
NG
NG
NG
Pacific
coast
2
Unspecified
location
9
NF
NF
NF
NF
NF
NF
NG
NG
NG
NG
1
British
Columbia
2
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
1,3,4
central
California
10
NG
NG
NG
NG
NG
NG
NG
NG
3
Anaheim
Bay,
CA
5
NG
vertebral
centra;
whole
terramycin
marked,
held
in
intrareader
comparison
Pacific
coast
2
captivity
NG
rearing
in
captivity,
tag
and
recaptureNG
NG
3
southern
California7
width
frequency,
and
length
frequency
Pacific
coast
2
Page
223
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1167
5­
2220e
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Elasmobranchs
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53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
annual
spawner
2
5,14
10­
11
embryos
7
8­
13
offspring
24
average:
6
offspring
single
spawner
3
single
spawner
7
2
eggs
per
clutch
single
spawner
15
51
embryos
5,7,8,9,16
22­
108
offspring
single
spawner
7,17,18,
29
82­
95
offspring
single
spawner
7
<
28
offspring
annual
breeding
cycle
4
20
1­
6
embryos
7
3
offspring
(
average)

Page
224
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1167
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2220e
­
Elasmobranchs
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BE
BF
BG
BH
NF
21
10
months
NA
NF
12
young
hatch
in
8­
9
months
NA
2,10,13
young
hatch
in
7­
9
months
NF
NF
2,10,12
young
hatch
in
7.5­
10
months
NA
13
7.5
months
4
51
embryos
(
420
cm)
NF
15
12
month
gestation
NA
NF
NF
2,14,15
12
month
gestation
NA
NF
NA
NF
NF
16
3
month
gestation
NA
Page
225
of
1167
5­
2220e
­
Elasmobranchs
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BI
BJ
BK
BL
BM
BN
NA
NF
NA
NA
NA
NA
NA
NF
NF
lodged
under
reefs
or
northern
elephant
seals
1
NA
crevices
4,5
NA
NF
NF
NF
northern
elephant
seals
1
NA
NA
NF
NA
NA
NA
NA
NA
NF
5
8­
77
mm
(
internal
eggs)
NA
NA
NA
NA
NF
NA
NA
NA
NA
NA
NF
NA
NA
NA
NA
Page
226
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1167
5­
2220e
­
Elasmobranchs
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BO
BP
BQ
BR
BS
BT
BU
NA
NA
NA
NA
NA
2
21
cm
to
maturity
sand
and
mud
bottoms
5,6
6
24
cm
to
maturity
202
NA
NA
NA
NA
NA
2,3
15
cm
to
maturity
Rocky
to
sandy
areas
6
NA
NA
NA
NA
NA
2
13
cm
to
maturity
rocky
areas,
often
in
crevices
6
NA
NA
NA
NA
NA
<
120
cm
often
found
inshore
2
2
65
cm
to
maturity
3
61
cm
to
maturity
NA
NA
NA
NA
NA
7
43­
186.5
cm
TL
found
in
shallow
open
coast
and
bays
2,4,7
2,3
45
cm
to
maturity
NA
NA
NA
NA
NA
3
15
cm
to
maturity
shallow
sandy
areas
4,6
NA
NA
NA
NA
NA
3
approx.
8­
25
cm
Sand
or
mud
bottom
off
beaches
and
in
bays
and
sloughs
1
juveniles
found
more
inshore
than
adults
4,8
Page
227
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1167
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Elasmobranchs
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BV
BW
BX
BY
carnivore
benthic
ambusher
6,7
NF
NF
carnivore
(
worms,
anemones)
14
scours
seafloor
in
search
of
prey
8
nocturnal
and/
or
NF
crepuscular
feeder
2,3
NF
NF
nocturnal
3
northern
elephant
seals
3
carnivore
(
cephalopods,
teleosts,
chondrichthyans)
15
NF
NF
NF
carnivore
NF
NF
NF
carnivore
(
prim.
crabs)
8
bottom
feeder
9
continuously
throughout
day
1
NF
carnivore
bottom
feeders
9
continuously
throughout
day
1,4,5NF
(
polychaetes,
clams)
10
browsers,
crush
shells
10
(
worms,
shrimp,
crabs,
amphipods)
14
Page
228
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1167
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­
Elasmobranchs
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BZ
CA
CB
(
anchovies,
other
fish,
shrimp,
and
euphausiids)
50
(
fish,
mollusks,
crustaceans,
annelids,
and
coelenterates)
52
sand
and
mud
bottoms
6,7
benthic
ambusher
1,5,6,7
often
buried
in
mud
or
sand
2,8,16
found
in
sand­
bottom
kelp
holdfast
and
rocky­
bottom
kelp­
bed
habitat
16
(
fish,
squids,
sea
cucumbers)
4
(
bottom
and
epipelagic
fishes
and
squids)
5
rocky
to
sandy
areas
7
carnivore
(
crustaceans
and
small
fish)
19
scours
seafloor
in
search
of
prey
8
rocky
bottom,
kelp
beds,
sandy
draws
between
rocks
and
on
sand
flats
(
gastropods,
crustaceans,
pelecypods,
sipunculoids,

rocky­
bottom
kelp­
bed
habitat16
cephalopods,
small
fish)
20
(
benthic
invetebrates
and
small
fish)
5
rocky
areas,
often
in
crevices
7,14
carnivore
(
fish)
13
suction
or
"
yawn";
in
which
the
rocky,
algal­
covered
areas
of
kelp
beds,
also
in
algal­
covered
bottom
(
bony
fish,
dead
and
alive,
and
crustaceans)
5
mouth
is
held
open
until
prey
without
kelp
2
wanders
in
or
is
carried
in
by
current
5
rocky­
bottom
kelp­
bed
habitat16
locates
prey
by
detecting
electrical
fields
5
marine
and
benthic
or
pelagic,
on
the
continental
and
insular
carnivores
NF
shelves
and
upper
slopes
2
(
primarily
bony
and
cartilagenous
fish)
21,22,23
deep
water
along
coasts
9
(
fish
and
crustaceans)
13,14,16
(
cephalopods,
teleosts,
chondrichthyans,
marine
mammals)
24
(
sharks,
rays,
bony
fish,
chimaeras,
squid,
shrimp,
crabs,

carrion,
and
seals)
5
marine
and
benthic,
neritic,
on
the
continental
shelves
2
carnivores
(
primarily
cartilagenous
fish,
bony
fish,
ambush
strategy,
stealth,
group
shallow
open
coast
and
bays
9,10
and
mammals)
4,5,22,23,25,26
hunting,
bursts
of
speed
9
shallow
sandy
areas
7,11,16
benthic
carnivore
(
crustaceans,
fish,
and
clams)
4,10,11,13,27
pursuit
and
capture
over
mudflats
4
found
in
rocky­
bottom
kelp­
bed
habitat16
(
worms
and
crustaceans)
51
bottom
feeder
10
hunts
with
snouts
above
the
seafloor
11
sand
or
mud
bottom
off
beaches
and
in
bays
and
sloughs
1,12,16
carinvores
bottom
feeders
10
found
in
sand­
bottom
kelp
holdfast
and
rocky­
bottom
kelp­
bed
habitat
16
(
clams,
ghost
shrimp,
polychaetes,
and
crabs)
4,11,27,28
browsers,
crush
shells
12
(
worms,
small
crustaceans,
and
mollusks)
51
digs
by
flapping
of
rostrum
and
pectoral
fins
13
(
pelecypods,
polychaetes,
and
crustaceans)
55
carnivore
(
feeds
on
fast
moving
fish,
such
as
corbina,
queenfish,

and
California
halibut)
16,19,51
(
mature
herring,
young
salmon,
sticklebacks,
sand
lance,
pipefish,

cephalopods,
and
schizopods)
49
benthic
in
nearshore,
soft
bottom
habitats,
with
highest
concentration
found
near
protected
inlets
and
embayments
were
water
temperatures
17
Page
229
of
1167
5­
2220e
­
Elasmobranchs
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CC
CD
CE
CF
CG
CH
day
and
night
feeder
2
northern
elephant
seals
3
Santa
Rosa
Island
NF
NG
NG
nocturnal
3,4,5
Santa
Cruz
Island
NG
NG
San
Clemente
Island
1
NG
NG
nocturnal
and/
or
northern
elephant
seals,
bald
eagles
3
NF
NF
NF
NF
crepuscular
feeder
6,7,8
primarily
nocturnal
feeder
3,7,8,9,16,17
northern
elephant
seals
3
NF
NF
NF
NF
may
be
a
nocturnal
feeder
8
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
continuously
throughout
day
1
NF
NF
NF
NF
NF
continuously
throughout
day
1,10,11
elephant
seals
3
NF
NF
NF
NF
Page
230
of
1167
5­
2220e
­
Elasmobranchs
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CI
CJ
CK
CL
CM
CN
CO
CP
1
10.7%
NG
0.052
allozymes
NG
NF
NF
NF
1
10.7%
NG
0.037
NG
14.30%
NG
0.078
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
231
of
1167
5­
2220e
­
Elasmobranchs
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53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CQ
CR
CS
CT
CU
CV
CW
CX
NF
NF
NF
4
M=
0.15­
0.20
NG
NG
Hoenig's
equation
NG
4
M=
0.14
NG
NG
Pauly's
method
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
232
of
1167
5­
2220e
­
Elasmobranchs
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CY
CZ
DA
DB
DC
NG
NG
flatfish,
California
lizardfish,
parasites:
8
families,
8
spp.
(
see
reference)
10
7
2­
9
km
(
tracked
for
periods
of
13­
25
hrs)

NG
NG
lingcod
2
8
96
km
in
46
days
NF
NF
NF
parasites:
5
families,
5
spp.
(
see
reference)
10
NF
NF
NF
NF
parasites:
1
family,
1
spp.
(
see
reference)
10
NF
NF
NF
NF
9
<
10
km,
swam
at
depths
between
600­
1,100
m
parasites:
9
families,
25
spp.
(
see
reference)
10
NF
NF
NF
Parasites:
isopod(
Lironeca
vulgaris
),
copepod
(
Pandarus
bicolor
),
10
0­
539
km
and
the
leech
(
Branchellion
lobata
)
3
parasites:
5
families,
6
spp.
(
see
reference)
10
NF
NF
NF
parasites:
8
families,
13
spp.
(
see
reference)
10
NF
NF
NF
benthic
flatfish
3
guitarfish,
California
halibut,
thornback,
benthic
flatfish1
11
<
4.75
nautical
miles
(
at
liberty
for
208
days)

parasites:
19
families,
40
spp.
(
see
reference)
10
parasites:(
Otodistomum
veliporum
),
and
the
copepod
(
Pandarus
bicolor)
3
7,13
tagging
revealed
angel
sharks
have
a
home
range
which
they
return
to
after
a
night
of
foraging
11
most
rays
moved
1­
2
nautical
miles
(
at
liberty
for
2­
284
days)

Page
233
of
1167
5­
2220e
­
Elasmobranchs
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
DD
DE
DF
DG
majority
are
residential
6,13
NF
NF
NF
may
migrate
seasonally
7,8
move
offshore
in
winter
6
NF
sensitive
to
electrical
fields
in
gen.
solitary,
sometimes
NF
water,
may
be
used
in
orientation
aggregate
4,5
or
locating
food
2
NF
NF
gen.
solitary,
sometimes
NF
aggregate
4
NF
NF
NF
NF
NF
Yes
1
NF
NF
137
NF
NF
often
found
in
NF
aggregations
4
observed
swimming
with
leopard
sharks
4
Resident
9
sensitive
to
electrical
fields
in
NF
NF
water,
may
be
used
in
orientation
2
Page
234
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
A
B
C
D
E
Myliobatidae
1,2
Myliobatis
californica
1,2
bat
ray
Oregon
to
Gulf
of
California
1,3,4,5
2
intertidal
to
108
m
eagle
ray,
California
stingray,

stingaree,
bat
stingray,
batfish
1,3
Torpedinidae
1,2
Torpedo
californica
1,2
Pacific
electric
ray
northern
British
Columbia
to
Bahia
de
Sebastian
2
3­
453
m
California
electric
ray,
electric
ray,
central
Baja
California
1,3,4,5
torpedo
fish,
California
torpedo,

crampfish,
torpedo
1,3,4
Platyrhinidae
Platyrhinoidis
triseriata
1,2
thornback
Tomales
Bay,
northern
California
to
Gulf
of
California
4
2
surfline
to
137
m
Platyrhininae
2
round
skate,
thornback,
guitarfish,

California
thornback,
shovelnose,

banjo
shark
1,3,5
Rajidae
1,2
Raja
binoculata
1,2
big
skate
Bering
sea
and
se.
Alaska
to
Cabo
Falsa,
1
3­
800
m
big
skate
of
California,
gangiei
rui
1,3,5
southern
Baja
California
1,4
Rajidae
1,2
Raja
inornata
1,2
California
skate
Washington
(
Straits
of
Juan
de
Fuca)
to
southern
1
18­
670
m
common
skate
1,3
Baja
California
and
Gulf
of
California
4
Rajidae
1,2
Raja
rhina
1,2
longnose
skate
Bering
Sea
and
Aleutian
Is.
to
Isla
Cedros,
central
Baja2
13­
800
m
California
longnose
skate
1,3
California
4
Rajidae
1
Bathyraja
kincaidii
2
sandpaper
skate
Sea
of
Japan
to
Bering
Sea
to
Cortes
Bank,
southern
2
17­
1,380
m
(
Bathyraja
interrupta
)
1,2
black
skate,
Bering
skate
1,4,6
California
4
Rajidae
1
Raia
stellulata
2
starry
skate
Bering
Sea
and
Aleutian
Is.
to
Coronado
Bank,
2
18­
732
m
Raja
stellulata
1
prickly
skate,
cheap
skate
1,7
northern
Baja
California
4
(
Raja
asterias
)
2
Page
235
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
F
G
H
I
J
NF
1
TL=.
758
DW
+­
44.413
(
r=.
9757)
2,3
1.8
m
(
DW)
Elkhorn
Slough
(
DW)
9,10
100.4(
204)

(
95%
confidence
intervals)

NF
NF
2,3
1.37+
m
NF
NF
NF
NF
2,3
91
cm
NF
NF
7,8
Both:
Log
W=
log
a
+
b
log
TL;
p<
0.001,
r2=
0.96
2
DW=
15.5
+
0.72
TL:
p<
0.001,
r2=
0.99
1,2,3
2.4
m
NF
NF
Y=­
35.1
+
44.2*
Log(
x)
158,
169
NF
NF
1,3
76
cm
NF
NF
7,8
Both:
Log
W=
log
a
+
b
log
TL;
r2=
0.94
2
DW=
17.3
+
0.69
TL:
p<
0.001,
r2=
0.98
1,3,5
137
cm
Monterey
Bay(
TL)
11,12
96.7(
10)

y=
­
20.4
+
3.2*
log
(
TL)
(
95
%
confidence
intervals)

NF
NF
3
86
cm
NF
NF
NF
NF
3
76
cm
NF
NF
Page
236
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
K
L
M
N
O
P
Q
R
S
T
U
158.7(
140)
NG
0.229(
0.11142)
0.0995(
0.0188)
NG
­
1.58(
0.541)
­
2.059(
0.354)
NG
NG
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
106.9(
13.1)
104.7(
9.1)
0.25(
0.10)
0.16(
0.05)
0.17(
0.05)
0.73(
1.1)
­
0.3(
0.08)
­
0.16(
0.62)
NG
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
237
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
V
W
X
Y
Z
AA
AB
vertebral
centra;
x­
radiography
tetracycline
mark­
recapture
compared
techniques
and
NF
3
>
24
yrs.
NF
dioecious
and
oil
clearing
intrareader
comparison
8
24
yrs.
(
150
cm)

1224
yrs
(
100
cm
DW)

NF
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
8,9
3
12
yrs.
NF
dioecious
10
12
yrs
(
160.7
cm)

NF
NF
NF
NF
6
20­
30
yrs
NF
dioecious
vertebral
centra;
parafin
edge
analysis
betw.
reader
comparison
NF
6
20­
30
yrs.
NF
dioecious
wax­
sectioning
10
13
yrs
(
132.2
cm)

NF
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
NF
dioecious
Page
238
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
AC
AD
AE
AF
AG
AH
AI
AJ
promiscuous
viviparous2,19
No
females
grow
larger
1,2
males
have
claspers
3
NG2
105
cm
DW
NG
20,25
45.0­
62.2
cm
DW
(
2­
3)
NG
NG
6
61
cm
DW
91
cm
DW
NG
promiscuous
aplacental
viviparity
12,13
No
males
have
claspers
NF
NF
NF
promiscuous
aplacental
viviparity
4
No
NF
males
have
claspers
NF
NF
NF
viviparous
13
promiscuous
oviparous
1,4,7,12,14,17
No
females
grow
larger
3,4
males
have
claspers
4
21,26,29100­
110
(
7­
8
yrs)
130
cm
(
12
yrs)
NG
promiscuous
oviparous
7,14
No
NF
males
have
claspers
22
52
cm
52
cm
NG
promiscuous
oviparous
4,7,12,14
No
females
grow
larger
5
males
have
claspers
4
21,26
60cm
(
5)
70
cm(
8
yrs)
NG
promiscuous
oviparous
1
No
NF
males
have
claspers
NF
NF
NF
promiscuous
oviparous
1,12
No
NF
males
have
claspers
NF
NF
NF
Page
239
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1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
AK
AL
AM
AN
AO
AP
AQ
AR
NG
NG
NG
NG
NF
Elkhorn
Slough,
CA
1,16
88.1
cm
DW
(
5)
vertebral
centra;
x­
radiography
tetracycline
mark­
recapture
compared
techniques
and
Pacific
coast
2
and
oil
clearing
intrareader
comparison
NG
NG
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Pacific
coast
2
NG
vertebral
centra;
parafin
edge
analysis
betw.
reader
comparison
2
NF
wax­
sectioning
NG
NG
NG
NG
NF
NF
NG
vertebral
centra;
parafin
edge
analysis
betw.
reader
comparison
2
NF
wax­
sectioning
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
240
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
annual
breeding
cycle4
3
2­
6
offspring
21
2­
5
embryos
7,31
2­
12
offspring
single
spawner
single
spawner
single
spawner
2,9
<
7
eggs
25
1­
7
embryos
per
egg
case
25
3­
4
average
embryos
per
egg
case
single
spawner
25
1
egg
per
egg
case
single
spawner
9,25
1
egg
per
egg
case
single
spawner
single
spawner
Page
241
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1167
5­
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­
Elasmobranchs
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105
106
107
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109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BE
BF
BG
BH
NF
NF
2,17,23,269­
12
month
gestation
NA
NF
NF
NA
NF
NF
NA
NF
18
approx.
1
year
NA
NF
NF
NA
NF
NF
NA
NF
NF
NA
NF
NF
Page
242
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1167
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2220e
­
Elasmobranchs
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104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BI
BJ
BK
BL
BM
BN
NA
NF
3
22­
28
mm
(
internal
eggs)
NA
NA
NA
NA
NF
NA
NA
NA
NA
NA
NF
NA
NA
NA
NA
NA
NF
NF
demersal1,7
elephant
seals
1
NA
NA
NF
10­
12
cm
demersal
1
NF
NA
NA
NF
1
8­
12
cm
(
length)
demersal
1,6
NF
NA
NA
NF
NF
NF
NF
NA
NF
NF
NF
NF
NA
Page
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1167
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Elasmobranchs
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106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BO
BP
BQ
BR
BS
BT
BU
NA
NA
NA
NA
NA
3
approx.
23
cm
to
maturity
mud
and
sandy
bottom
4
10
approx.
50
cm
TL
and
35
cm
DW
NA
NA
NA
NA
NA
NF
fine
sand
bottom
or
mud,
also
found
in
rock
and
kelp
bed
areas
6
NA
NA
NA
NA
NA
NF
fine
sand
to
mud
bottoms
6
NA
NA
NA
NA
NA
8
<
130
cm
soft
substrate
4
NA
NA
NA
NA
NA
NF
NF
NA
NA
NA
NA
NA
NF
NF
NA
NA
NA
NA
NA
NF
NF
NA
NA
NA
NA
NA
NF
NF
Page
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1167
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Elasmobranchs
103
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105
106
107
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109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BV
BW
BX
BY
carnivore
(
clams)
8
NF
NF
NF
carnivore
NF
NF
NF
benthic
carnivore
16
NF
NF
NF
opportunistic
carnivore
(
decapod
crustaceans)
17
use
of
electroreception
in
finding
buried
preNF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
245
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1167
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­
Elasmobranchs
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105
106
107
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109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BZ
CA
CB
intertidal
mudflats
in
estuaries,
open
surf,
and
estuaries
11
opportunistic
bottom
feeders43
digs
burrowing
clams
and
worms
from
substrate4
kelp
bed
and
adjacent
kelpless
cobble
area
4
carnivore
(
echiroid
worm,
clams,
fish)
10
digs
by
flapping
of
rostrum
and
pectoral
fins
13
found
in
rocky­
bottom
kelp­
bed
habitat16,17
(
epifaunal
molluscs,
infaunal
worms,
and
molluscs)
29
benthic
in
nearshore,
soft
bottom
habitats
17
(
mollusks
and
crustaceans)
43
(
infaunal
bivalves,
epifaunal
gammarid
amphipods,
and
gastropods)
45
(
clams,
crabs,
shrimps,
and
the
echiuroid
worm
Urechis
caupo
)
54
(
pelecypods,
polychaetes,
and
crustaceans)
55
fine
sand
bottom
or
mud,
also
found
in
rock
and
kelp
bed
areas
7
carnivore
(
bottom
dwelling
fish)
19,30
ambush
predator
14
kelp
bed
and
adjacent
kelpless
cobble
area
4
preys
on
fish
using
powerful
electric
discharges
5,1
during
the
day
,
these
animals
are
often
found
buried
in
the
substratum
15
found
in
sand­
bottom
kelp
holdfast
envrionment
16
fine
sand
to
mud
bottoms7,16
benthic
carnivore
(
polychaetes,
mollusks,
small
crustaceans)
4,19
NF
kelp
bed
and
adjacent
kelpless
cobble
area4
(
sand
dwelling
worms)
51
found
in
sand­
bottom
kelp
holdfast
and
rocky­
bottom
kelp­
bed
habitat
16
soft
substrate
11
opportunistic
carnivore
(
bottom
dwelling
fishes)
31
use
of
elctroreception
in
finding
buried
prey
16
(
fish
and
crustaceans)
4,14,15,43
inshore
muddy
bottoms
5
carnivore
(
shrimp
and
other
invertebrates)
15,32
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
(
clams,
crabs,
shrimp,
fish,
innkeeper
worms,
snails,
abalone,
and
worms)
4
benthic
predator
which
forages
in
shallow
mudflats
and
seagrass
beds
ain
bays
and
20
are
elevated
and
sediments
are
primarily
soft
sand
and
mud
17
Page
246
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1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CC
CD
CE
CF
CG
CH
NF
NF
NF
NF
NF
NF
diurnal
and
nocturnal
feeder
12
NF
NF
NF
NF
NF
nocturnal
3,13,15
northern
elephant
seal
3
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
247
of
1167
5­
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­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CI
CJ
CK
CL
CM
CN
CO
CP
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
248
of
1167
5­
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­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CQ
CR
CS
CT
CU
CV
CW
CX
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
249
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CY
CZ
DA
DB
DC
NF
NF
NF
NF
NF
NF
parasites:
3
families,
4
spp.
(
see
reference)
10
NF
NF
NF
NF
parasites:
4
families,
4
spp.
(
see
reference)
10
NF
NF
NF
NF
parasites:
copepod
(
Charopinus
dentatus
)
4
NF
parasites:
trematodes(
Otodistomum
veliporum
)
6
parasites:
11
families,
22
spp.
(
see
reference)
10
NF
NF
NF
parasites:
11
families,
21
spp.
(
see
reference)
10
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
parasites:
2
families,
2
spp.
(
see
reference)
10
NF
15tracked
moving
an
average
of
0.74
km/
hr
and
frequenting
the
shallow
parts
of
the
bay
15all
three
recaptures
caught
within
1
km
of
their
tagging
location
white
seaperch
and
senoritas
commonly
aggregate
and
feed
where
bat
ray
are
stirring
up
the
bottom
with
their
wings9
Page
250
of
1167
5­
2220e
­
Elasmobranchs
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
DD
DE
DF
DG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
tend
to
move
out
of
bays
and
estuaries
in
winter15
travel
inshore
to
spawn
and
breed
in
spring
and
summer2,14
yes,
large
schools
of
bat
rays
have
been
sighted
off
southern
and
central
12
found
solitary
and
in
aggregations4
Page
251
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1167
5­
2220e
­
Elasmobranchs
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156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
A
B
C
D
E
Chimaeridae
1,2
Hydrolagus
colliei
1,2
spotted
ratfish
se.
Alaska
to
Baja
and
n.
Gulf
of
California;
1,2
inshore
to
965
m
Pacific
ratfish,
chimaera,
ratfish,
common
off
British
Columbia
to
n.
California
1,4
goatfish,
king­
of­
the­
herrings,
water
hare,
elephant
fish,
rattail,
rabbitfish
1,3,8
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Gates
and
Frey
1974
1­
Eschmeyer
and
Herald
1983
1­
Eschmeyer
and
Herald
1983
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Leet
et
al
1992
2­
Kato
et
al
1967
2­
Love
et
al
2000
3­
Eschmeyer
and
Herald
1983
3­
Love
1996
3­
Love
1996
4­
Hart
1973
4­
Love
et
al
2000
5­
Love
1996
5­
Miller
and
Lea
1972
6­
Love
et
al
2000
7­
Chamberlain
1979
8­
Halstead
and
Bunker
1952
9­
Holts
1988
10­
Reilly
et
al
1994
Page
252
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
F
G
H
I
J
9
Males:
W=
0.000047676L3.0546
(
r2=.
9835)
NF
3,5
100
cm
NF
NF
Females:
W=
0.000067733L2.9720(
r2=.
9886)

1­
Ackerman
1971
1­
Martin
1982
1­
Eschmeyer
and
Herald
1983
1­
Kusher
et
al
1992
2­
Sandell
1973
2­
Zeiner
1991
2­
Love
1996
2­
Yudin
and
Caillet
1990
3­
Nammack
et
al
1985
3­
Love
et
al
2000
3­
Nammack
et
al
1985
4­
Jones
and
Geen
1977a
4­
Compagno
1984
4­
Jones
and
Geen
1977b
5­
Natanson
1984
5­
Hart
1973
5­
Saunders
and
McFarlane
1993
6­
Ebert
1984
6­
Castro
1996
6­
Holden
and
Meadows
1962
7­
Zeiner
1991
7­
Ebert
1986a
7­
Bedford
1987
8­
Zeiner
and
Wolf
1993
8­
Timmons
and
Bray
1997
9­
Johnson
and
Horton
1972
9­
Martin
1982
10­
Ripley
1946
10­
Martin
and
Caillet
1988a
11­
Saunders
et
al
1984
11­
Zeiner
1991
12­
Zeiner
and
Wolf
1993
13­
Yudin
1987
14­
Ketchen
1975
Page
253
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
K
L
M
N
O
P
Q
R
S
T
U
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
254
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
V
W
X
Y
Z
AA
AB
NF
NF
NF
NF
NF
NF
dioecious
1­
Weldon
et
al
1987
1­
Ackerman
1971
1­
Emmett
et
al
19
2­
Jones
and
Geen
1977b
2­
Kusher
et
al
1992
3­
Beamish
and
McFarlane
1985
3­
Love
1996
4­
Kirnosova
1989
4­
Castro
1996
5­
McFarlane
and
Beamish
1987
5­
Hart
1973
6­
Natanson
and
Caillet
1990
6­
Casillas
et
al
1998
7­
Natanson
1984
7­
Yudin
1987
8­
Zeiner
1991
8­
Martin
and
Caillet
1988a
9­
Fraser
1923
9­
Timmons
and
Bray
1997
10­
Smith
1984
10­
Zeiner
1991
11­
Ackerman
1971
11­
Ketchen
1975
12­
Babel
1967
12­
Hopkins
1993
13­
Soldat
1982
14­
Alverson
and
Stansby
1963
15­
Beamish
and
Smith
1976
16­
Holland
1957
Page
255
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
AC
AD
AE
AF
AG
AH
AI
AJ
promiscuous
oviparous
1,2,7,12
No
females
grow
larger
5
Males:
possess
frontal
tenaculum,
6
20
cm(
snout
to
vent)
25
cm(
snout
to
vent)
NG
prepelvic
tenacula,
and
claspers
2
991
1­
Eschmeyer
and
Herald
1983
1­
Love
1996
1­
Norwell
1994
1­
Ackerman
1971
2­
Love
1996
2­
Martin
1982
2­
Johnson
and
Horton
1972
2­
Talent
1985
3­
Emmett
et
al
1991
3­
Zeiner
1991
3­
Martin
1982
3­
Kusher
et
al
1992
4­
Roedel
and
Ripley
1950
4­
Zeiner
and
Wolf
1993
4­
Zeiner
1991
4­
Emmett
et
al
1991
5­
Castro
1996
5­
Casillas
et
al
1998
5­
Babel
1967
5­
Compagno
1984
6­
Ripley
1946
6­
de
Wit
1975
6­
Love
1996
7­
Casillas
et
al
1998
7­
Yudin
1987
7­
Ford
1921
8­
Hishaw
and
Albert
1947
8­
Ford
1921
8­
Hart
1973
9­
Compagno
1984
9­
Ketchen
1972
10­
Ebert
1986
©
10­
Jones
and
Geen
1977c
11­
Herald
and
Ripley
1951
11­
Saunders
and
McFarlane
1993
12­
Hart
1973
12­
Natanson
1984
13­
Zorzi
and
Martin
1995
13­
Bedford
1987
14­
Zorzi
1994
14­
Crow
et
al
1996
15­
Compagno
1984
15­
Ebert
1986a
16­
Breder
and
Rosen1966
16­
Ebert
1986b
17­
Fraser
1923
17­
Ebert
1984
18­
Daugherty
1960b
18­
Ebert
1996
19­
Hopkins
1993
19­
Timmons
and
Bray
1997
20­
Martin
and
Caillet1988b
21­
Zeiner
and
Wolf
1993
22­
Martin
and
Zorzi
1993
23­
Yudin
1987
24­
Holt
1988
25­
Martin
1982
26­
Zeiner
1991
27­
Babel
1967
28­
Ketchen
1975
29­
Fleming
1999a
30­
Saunders
et
al
1984
31­
Daugherty
1960b
32­
Hopkins
1993
Page
256
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
AK
AL
AM
AN
AO
AP
AQ
AR
NG
NG
NG
NG
NF
California
12,13
Pacific
coast
2
Unspecified
location
11
1­
Compagno
1984
1­
Talent
1985
2­
Casillas
et
al
1998
2­
Love
1996
3­
Love
1996
3­
Eigenmann
1891
4­
Hart
1973
4­
Emmett
et
al
1991
5­
Alverson
and
Stansby
1963
5­
Ketchen
1972
6­
Sandell
1973
7­
Babel
1967
8­
Jones
and
Geen
1977c
9­
Compagno
1984
10­
Ebert
1986a
11­
Hart
1973
12­
Yudin
1987
13­
Dean
1906
14­
Ripley
1946
15­
Holts
1988
16­
Martin
and
Caillet
1988b
17­
Alverson
and
Stansby
1963
Page
257
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
*****
*****
*****
*****
annual
breeding
cycle
1
2,7
2
eggs
*****
*****
*****
*****
*****
*****

*****
*****
*****
*****
1­
Casillas
et
al
1998
1­
Ackerman
1971
2­
Bedford
1987
2­
Casillas
et
al
1998
3­
Roedel
and
Ripley
1950
3­
Talent
1985
4­
Martin
1982
4­
Smith
et
al
1998
5­
Fleming
1999a
5­
Compagno
1984
6­
Emmett
et
al
1991
7­
Love
1996
8­
Castro
1996
9­
Hart
1973
10­
Roedel
and
Ripley
1950
11­
Nammack
et
al
1985
12­
Ketchen
1972
13­
Kirnosova
1989
14­
Natanson
1984
15­
Ebert
1986a
16­
Eschmeyer
and
Herald
1983
17­
Ebert
1989
18­
Ebert
1986a
19­
Ebert
1984
20­
Babel
1967
21­
Martin
and
Caillet
1988b
22­
Yudin
1987
23­
Ripley
1946
24­
Holts
1988
25­
Martin
and
Zorzi
1993
26­
Alverson
and
Stansby1963
27­
Ketchen
1972
28­
Fleming
1999a
29­
Ford
1921
30­
Daugherty
1960b
31­
Hopkins
1993
Page
258
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
BE
BF
BG
BH
NF
NF
2,18
1
yr.
NA
19
9­
12
months
1­
de
Wit
1975
1­
Nammack
et
al
1985
1­
Talent
1985
2­
Ackerman
1971
2­
Kirnosova
1989
2­
Love
1996
3­
Yudin
1987
3­
Ketchen
1972
4­
Ebert
1984
4­
Sandell
1973
5­
Castro
1996
6­
Roedel
and
Ripley
1950
7­
Hart
1973
8­
Jones
and
Geen
1977c
9­
Hishaw
and
Albert
1947
10­
Compagno
1984
11­
Kirnosova
1989
12­
Feder
et
al
1974
13­
Eschmeyer
and
Herald
1983
14­
Ebert
1986a
15­
Ebert
1984
16­
Babel
1967
17­
Martin
and
Caillet
1988b
18­
Casillas
et
al
1998
19­
Dean
1906
20­
Ripley
1946
21­
Holts
1988
22­
Alverson
and
Stansby
1963
23­
Fleming
1999a
24­
Ford
1921
25­
Daugherty
1960b
26­
Hopkins
1993
Page
259
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
BI
BJ
BK
BL
BM
BN
NA
NF
2
egg
capsule:
2.9
by
1.9
by
1.3
cm
demersal
2,3
northern
elephant
seals
1
NA
egg:
35
by
20
by
12
mm
planted
vertically
in
mud
or
attached
to
rocks
1,4
found
in
polyhaline
and
euhaline
waters
1
1­
Casillas
et
al
1998
1­
Casillas
et
al
1998
1­
Love
1996
2­
Dean
1906
2­
Allen,
M.
J.
1982
3­
Yudin
1987
3­
Sathyanesan
1966
4­
Roedel
and
Ripley
1950
4­
Love
1996
5­
Ebert
1996
5­
Compagno
1984
6­
Roedel
and
Ripley
1950
7­
Fraser
1923
Page
260
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
BO
BP
BQ
BR
BS
BT
BU
NA
NA
NA
NA
NA
NF
demersal
9
muddy
bottom
10
generally
on
rocky
botom
in
deepwater,
also
common
in
estuaries
11
1­
Emmett
et
al
1991
1­
Eschmeyer
and
Herald
1983
2­
Compagno
1984
2­
Compagno
1984
3­
Love
1996
3­
Emmett
et
al
1991
4­
Ketchen
1972
4­
Love
1996
5­
Hart
1973
5­
Allen
1990
6­
Bedford
1987
6­
Feder
et
al
1974
7­
Ebert
1986a
7­
Ebert
1984
8­
Zeiner
and
Wolf
1993
8­
Babel
1967
9­
Yudin
1987
9­
Allen,
M.
J.
1982
10­
Martin
1982
10­
Hart
1973
11­
Beamish
and
Smith
1976
11­
Casillas
et
al
1998
12­
San
Filippo
1995
Page
261
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
BV
BW
BX
BY
opportunistic
carnivore
(
shrimp,
mollusks,
echinoderms)
18
NF
NF
spiny
dogfish
4
(
prim.
polychaetes)
19
1­
Ackerman
1971
1­
Compagno
1984
1­
Sandell
1973
1­
Leet
et
al
1992
2­
Barry
et
al
1996
2­
Ackerman
1971
2­
Segura­
Zarosa
et
al
1997
2­
Ketchen
1986
3­
Compagno
1984
3­
Russo
1975
3­
Love
1996
3­
Love
1996
4­
Talent
1976
4­
Talent
1976
4­
Ketchen
1972
4­
Quinn
et
al
1980
5­
Castro
1996
5­
Talent
1982
5­
Babel
1967
6­
Emmett
et
al
1991
6­
Fouts
and
Nelson
1999
6­
San
Filippo
1995
7­
Casillas
et
al
1998
7­
Allen
1990
8­
Talent
1982
8­
Segura­
Zarosa
et
al
1997
9­
Galvan­
Magana
et
al
1989
9­
Ketchen
1972
10­
Sandell
1973
10­
Babel
1967
11­
Taylor
1970
11­
Badkin
(
unpublished)

12­
Roedel
and
Ripley
1950
13­
Hart
1973
14­
Love
1996
15­
Ebert
1994
16­
Feder
et
al
1974
17­
Badkin
(
unpublished)

18­
Johnson
and
Horton
1972
19­
Quinn
et
al
1980
20­
Beamish
and
Smith
1976
21­
Saunders
et
al
1984
22­
Fraser
1923
Page
262
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
BZ
CA
CB
demersal
1
opportunistic
carnivore
(
shrimp,
mollusks,
echinoderms)
33
benthic
searcher,
uses
smell,

found
in
sand
and
mud
to
rocky
reef
11
(
brittle
stars,
myodocopine
ostracods,
gammarid
amphipods)
34
electroreception,
and
lateral
line
to
detect
prey
17
generally
on
rocky
botom
in
deepwater,
also
common
in
estuaries
5
(
clams,
crustaceans,
shrimp,
fish)
14
(
shrimp,
clams,
worms,
fishes,
brittlestars,
gammarid
amphipods,

and
seastars)
4
omnivore
(
molluscs,
crustaceans)
35
(
bivalves,
fish,
decapods,
limpets,
barnacles)
36
(
fish,
mollusks,
squid,
nudibranchs,
opisthobranchs,
annelid,
and
crustaceans)
38
1­
Eschmeyer
and
Herald
1983
1­
Ackerman
1971
1­
Compagno
1984
2­
Compagno
1984
2­
Leet
et
al
1992
2­
Russo
1975
3­
Emmett
et
al
1991
3­
Emmett
et
al
1991
3­
Talent
1976
4­
Larson
and
DeMartini
1984
4­
Love
1996
4­
Talent
1982
5­
Casillas
et
al
1998
5­
Compagno
1984
5­
Love
1996
6­
Allen
1990
6­
Russo
1975
6­
Fouts
and
Nelson
1999
7­
Feder
et
al
1974
7­
Talent
1976
7­
Allen
1990
8­
Castro
1996
8­
de
Wit
1975
8­
Segura­
Zarosa
et
al
1997
9­
Ebert
1984
9­
Haesker
and
Czech
1993
9­
Ebert
1991b
10­
Ebert
1986a
10­
Talent
1982
10­
Ketchen
1972
11­
Love
1996
11­
Sandell
1973
11­
Zorzi
and
Martin
1995
12­
Babel
1967
12­
Ripley
1946
12­
Babel
1967
13­
Fleming
1999a
13­
Roedel
and
Ripley
1950
13­
Van
Blaricom
1976
14­
Turner
et
al
1969
14­
Hart
1973
14­
Lowe
et
al
1994
15­
Lowe
et
al
1994
15­
Casillas
et
al
1998
15­
Bray
and
Hixon
1978
16­
Limbaugh
1955
16­
Castro
1996
16­
Badkin
(
unpublished)

17­
O'Brien
and
Valle
2000(
in
press)
17­
Jensen
1965
17­
Allen,
M.
J.
1982
18­
Ebert
et
al
1992
18­
Ripley
1946
19­
Feder
et
al
1974
19­
Brodeur
et
at
1987
20­
Segura­
Zarosa
et
al
1997
20­
Hopkins
1993
21­
Ebert
1986
c
22­
Ebert
1984
23­
Ebert
1986a
24­
Ebert
1994
25­
Ebert
1989
26­
Ebert
1991
(
a)

27­
Ketchen
1972
28­
Babel
1967
29­
Barry
et
al
1996
30­
Bray
and
Hixon
1978
31­
Badkin
(
unpublished)

32­
Zorzi
1994
33­
Johnson
and
Horton
1972
34­
Allen,
M.
J.
1982
35­
Halstead
and
Bunker
1952
36­
Quinn
et
al
1980
37­
Alverson
and
Stansby
1963
38­
Dean
1906
39­
Taylor
1970
40­
Herald
and
Ripley
1951
41­
Alverson
and
Stansby
1963
42­
Bonham
1954
43­
Fleming
1999a
Page
263
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
CC
CD
CE
CF
CG
CH
nocturnal
14
soupfin
sharks,
dogfish,
Pacific
halibut,
NF
NF
NF
pigeon
guillemots,
northern
fur
seals,

and
northern
elephant
seals
3
spiny
dogfish
2
1­
Sandell
1973
1­
Emmett
et
al
1991
1­
Gaida
1997
2­
Fouts
and
Nelson
1999
2­
Casillas
et
al
1998
3­
Love
1996
3­
Love
1996
4­
Standora
and
Nelson
1977
4­
Ketchen
1972
5­
Bedford
1987
5­
Ketchen
1986
6­
Segura­
Zarosa
et
al
1997
6­
Castro
1996
7­
Nelson
and
Johnson
1970
8­
Compagno
1984
9­
Castro
1996
10­
Ketchen
1972
11­
Babel
1967
12­
Lowe
et
al
1994
13­
Bray
and
Hixon
1978
14­
Allen,
M.
J.
1982
15­
Caillet
et
al
1979
16­
Ebeling
and
Bray
1976
17­
Turner
et
al
1969
Page
264
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
CI
CJ
CK
CL
CM
CN
CO
CP
NF
NF
NF
NF
NF
NF
NF
1­
Gaida
1997
1­
Smith
and
Abramson
1990
Page
265
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
CQ
CR
CS
CT
CU
CV
CW
CX
NF
NF
NF
NF
NF
NF
NF
NF
1­
Smith
and
Abramson
1990
2­
Au
and
Smith
1997
3­
Wood
et
al
1978
4­
Bedford
1987
Page
266
of
1167
5­
2220e
­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
CY
CZ
DA
DB
DC
NF
NF
NF
venomous
dorsal
spine
capable
of
inflicting
injury
2
NF
reproductive
organs
toxic
when
comsumed8
parasites:
copepod
(
Caligus
gurnardi
)
4
parasites:
12
families,
15
spp.
(
see
reference)
10
1­
Casillas
et
al
1998
1­
Babel
1967
1­
Smith
and
Abramson
1990
2­
Allen
1990
2­
Halstead
and
Bunker
1952
2­
Compagno
1984
3­
Babel
1967
3­
Ebert
1984
3­
Herald
and
Ripley
1951
4­
Fraser
1919
4­
Hart
1973
5­
Compagno
1984
5­
Jensen
1965
6­
Lloyd
1938
6­
McFarlane
and
Beamish
1986
7­
Beamish
and
Smith
1976
7­
Standora
and
Nelson
1977
8­
McAllister
1968
8­
Bedford
1987
9­
Bray
and
Ebeling
1975
9­
Carey
and
Clark
1995
10­
Love
and
Moser
1983
10­
Ebert
1996
11­
Babel
1967
12­
Ackerman
et
al
2000
13­
Holts
1988
14­
Holland
1957
15­
Hopkins
1993
Page
267
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1167
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­
Elasmobranchs
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
DD
DE
DF
DG
migratory
movement
10
NF
NF
NF
1­
Compagno
1984
1­
Van
Dykhuizen
et
al
1998
1­
Eschmeyer
and
Herald
1983
2­
Love
1996
2­
Knudtson
1977
2­
Castro
1996
3­
Ketchen
1986
3­
Leet
et
al
1992
4­
Hart
1973
4­
Love
1996
5­
Hishaw
and
Albert
1947
5­
Compagno
1984
6­
Bedford
1987
6­
Casillas
et
al
1998
7­
Galvan­
Magana
et
al
1989
7­
Jensen
1965
8­
Standora
and
Nelson
1977
8­
De
Wit
1975
9­
Babel
1967
9­
Breder
and
Rosen
1966
10­
Mathews
1975
10­
Ford
1921
11­
Casillas
et
al
1998
11­
Limbbaugh
1955
12­
Quinn
et
al
1980
12­
Hopkins
1993
13­
Holts
1988
14­
Martin
1982
15­
Hopkins
1983
seasonal
and
diel
migrations:
move
to
shallow
waters
in
spring
and
return
to
deep
waters
in
summer
and
2
11
nocturnal
onshore
movemements
of
small
ratfish11,12
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268
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210
211
212
213
214
215
216
217
218
219
220
A
B
C
D
E
Page
269
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­
Elasmobranchs
209
210
211
212
213
214
215
216
217
218
219
220
F
G
H
I
J
Page
270
of
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­
Elasmobranchs
209
210
211
212
213
214
215
216
217
218
219
220
K
L
M
N
O
P
Q
R
S
T
U
Page
271
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Elasmobranchs
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210
211
212
213
214
215
216
217
218
219
220
V
W
X
Y
Z
AA
AB
Page
272
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Elasmobranchs
209
210
211
212
213
214
215
216
217
218
219
220
AC
AD
AE
AF
AG
AH
AI
AJ
Page
273
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210
211
212
213
214
215
216
217
218
219
220
AK
AL
AM
AN
AO
AP
AQ
AR
Page
274
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Elasmobranchs
209
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211
212
213
214
215
216
217
218
219
220
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
Page
275
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Elasmobranchs
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215
216
217
218
219
220
BE
BF
BG
BH
Page
276
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1167
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­
Elasmobranchs
209
210
211
212
213
214
215
216
217
218
219
220
BI
BJ
BK
BL
BM
BN
Page
277
of
1167
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­
Elasmobranchs
209
210
211
212
213
214
215
216
217
218
219
220
BO
BP
BQ
BR
BS
BT
BU
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278
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Elasmobranchs
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215
216
217
218
219
220
BV
BW
BX
BY
Page
279
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Elasmobranchs
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211
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215
216
217
218
219
220
BZ
CA
CB
44­
Brodeur
et
al
1987
45­
Barry
et
al
1996
46­
Saunders
et
al
1984
47­
Ford
1921
48­
San
Filippo
1995
49­
Fraser
1923
50­
Reilly
et
al
1994
51­
Limbaugh
1955
52­
Holden
1966
53­
Daugherty
1960b
54­
Hopkins
1993
55­
O'Brien
and
Valle
2000
Page
280
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1167
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­
Elasmobranchs
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211
212
213
214
215
216
217
218
219
220
CC
CD
CE
CF
CG
CH
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281
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Elasmobranchs
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210
211
212
213
214
215
216
217
218
219
220
CI
CJ
CK
CL
CM
CN
CO
CP
Page
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211
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213
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215
216
217
218
219
220
CQ
CR
CS
CT
CU
CV
CW
CX
Page
283
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Elasmobranchs
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211
212
213
214
215
216
217
218
219
220
CY
CZ
DA
DB
DC
Page
284
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Elasmobranchs
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212
213
214
215
216
217
218
219
220
DD
DE
DF
DG
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Elasmobranchs
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Embiotocidae1
Amphistichus
argenteus
1
1surf
to
151
Amphistichus
koelzi
1
1surf
to
9.1
Amphistichus
rhodoterus
1
1surf
to
7.3
Cymatogaster
aggregata
1
1inshore
to
209
TAXONOMY
RANGE
calico
surfperch;
California
porgie;

humback
perch1,2
barred
surfpech;
barred
perch;
silver
perch;
surf
perch;
sand
perch;
silver
surf
fish1,2,5
n.
WA
to
n.­
c.
Baja
CA
(
Bahia
Santa
Rosalia)
1
WA
(
Cape
Flattery)
n.
Baja
CA
(
Arroyo
San
Isidro)
1
redtail
surfperch;
rosy
surf
fish;
redtail
seaperch;
porgy;
Oregon
porgie1,2
shiner
surfperch;
shiner;
sparada;
minny;

shiner
seaperch;
yellow
shiner;
bay
perch;

seven­
eleven
perch1,2,6
B.
C.
(
Vancouver
I.)
to
c.
CA
(
Avila
Beach)
1
s.
e.
Alaska
(
Wrangell)
to
c.
Baja
CA
(
Punta
Abreojos)
1
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
F
G
H
I
JKLM
Length
weight
equation
Max.
reported
Geographic
Area
Linf
in
cm
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
1Male:
W=
0.0000214SL3.1025
2TL=
1.2906(
SL)
+
0.0312
(
for
sizes
42
to
178
mm)
2
43
s.
California
(
SL)
3
28.1
33.9
NG
NG
1Female:
W=
0.0000386SL2.9914
2TL=
1.2054(
SL)
+
15.01
(
for
sizes
180
to
350
mm)

5TL=
2.93
+
1.24(
SL);
r2=
0.998;
SE=
2.37
5TL=
­
3.62
+
1.06(
FL);
r2=
0.996;
SE=
4.23
5SL=
­
2.03
+
0.805(
TL);
r2=
0.998;
SE=
1.91
5SL=
­
4.00
+
0.877(
FL);
r2=
0.998;
SE=
1.77
5FL=
­
2.52
+
0.943(
TL);
r2=
0.996;
SE=
3.99
5FL=
4.88
+
1.14(
SL);
r2=
0.998;
SE=
2.01
NF
5TL=
1.23
+
1.28(
SL);
r2=
0.996;
SE=
2.79
1
30
NF
NF
NF
NF
NF
5TL=
­
4.12
+
1.12(
FL);
r2=
0.994;
SE=
3.27
5SL=
­
0.42
+
0.778(
TL);
r2=
0.996;
SE=
2.17
5SL=
­
5.14
+
0.88(
FL);
r2=
0.996;
SE=
2.01
5FL=
4.77
+
0.887(
TL);
r2=
0.994;
SE=
2.91
5FL=
6.36
+
1.133(
SL);
r2=
0.996;
SE=
2.28
*
13Male:
W=
0.000011TL3.08
(
r2=
0.986)
5TL=
0.321
+
1.26(
SL);
r2=
0.996;
SE=
3.97
2
41
NF
NF
NF
NF
NF
*
13Female:
W=
0.00000813TL3.14
(
r2=
0.988)
5TL=
2.88
+
1.06(
FL);
r2=
0.998;
SE=
3.12
*
13Both:
W=
0.00000912TL3.12
(
r2=
0.986)
5SL=
0.308
+
0.791(
TL);
r2=
0.996;
SE=
3.14
5SL=
­
0.77
+
1.172(
FL);
r2=
0.998;
SE=
2.04
5FL=
­
2.47
+
0.941(
TL);
r2=
0.998;
SE=
2.93
5FL=
0.828
+
0.852(
SL);
r2=
0.998;
SE=
1.74
11SL=
0.81TL
­
5.29
(
r2=
0.998)

11FL=
0.95TL
­
5.09
(
r2=
0.998)

1Male:
W=
0.0000158SL3.111
1TL=
1.245(
SL)
+
1.771
1
19.3
s.
California
(
SL)
1
NG
NG
12.87
(
0.285)
NG
1Female:
W=
0.000009697SL3.212
3,9TL=
1.3194(
SL)
(
sizes
<
99
mm)

2,11Male:
W=
0.00000554SL3.38776
3,9SL=
0.7579(
TL)
(
sizes
<
99
mm)

2,11Female:
W=
0.0000117SL3.19885
3,9TL=
1.3036(
SL)
(
sizes
>
100
mm)

2,11Both:
W=
0.0000106SL3.22607
3,9SL=
0.7671(
TL)
(
sizes
>
100
mm)

3,5Both:
W=
0.000491SL3.05
3,9TL=
1.3133(
SL)

7Both:
W=
0.000017SL3.125
(
r2=
0.992)
3,9SL=
0.7615(
TL)

8Both:
W=
0.00407TL3.45
(
r2=
0.98)
4,6TL=
1.26(
SL)
+
1.73
(
r2=
1.0)

15Both:
W=
0.0000229FL3.02
5TL=
5.27
+
1.22(
SL);
r2=
0.958;
SE=
3.08
5TL=
4.4
+
1.06(
FL);
r2=
0.996;
SE=
2.61
5SL=
­
1.35
+
0.785(
TL);
r2=
0.958;
SE=
2.47
5SL=
­
3.93
+
0.914(
FL);
r2=
0.96;
SE=
2.29
5FL=
­
3.74
+
0.939(
TL);
r2=
0.996;
SE=
2.45
5FL=
6.92
+
1.05(
SL);
r2=
0.96;
SE=
2.46
13TL=
0.54
+
1.05FL;
r2=
0.92
13SL=
­
0.34
+
0.92FL;
r2=
0.94
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)
equation.
*

denotes
TL
=
mm
or
FL,
SL=
cm
relationships,
but
no
equations,
given
for
males
and
females
in
Gordon
(
1965)
Page
287
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1167
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67
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
N
O
P
Q
R
S
T
U
V
W
X
Growth
Parameters
k
(
SE)
to
(
SE)

Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
NG
NG
NG
NG
NG
NG
NG
NG
scales;
pelvic
bones
No
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NG
0.063
(
0.003)
NG
NG
­
0.45
mo.
(
0.023)
NG
NG
NG
Peterson
length
frequency
distribution
No
No
intrareader
comparison/
multiple
structure
comparison
AGE
&
GROWTH
r2
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
Y
Z
AA
AB
AC
Age
composition
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
NF
6,13
9
NF
Dioecious1
NF
107
to
9
16
6
(
Males);
9
(
Females)

NF
NF
NF
Dioecious
NF
7
6
8
NF
Dioecious
NF
16
9
1,2,4,9,11
1,15,16
8
2,4
Dioecious2
promiscuous1,2,4
9
9
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12
13
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15
16
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AD
AE
AF
AG
AH
AI
Sexual
dimorphism
Mode
of
reproduction
Parental
Care
Male
vs.
Female
length
Secondary
Characteristics
Male
Female
Viviparous2,18,28,31
NF
NF
1,13,26
NG
(<
1)
NG
(
2)

213
(
2)
13
(
2)

Males
have
a
tubercular
penis4
8NG
(
NG)
NG
(
NG)

Males
have
modified
anal
fin11
2213
(
NG)
­
sex
unspecified
Viviparous2,18,28
NF
NF
Males
have
a
tubercular
penis4
22NG
(
NG)
NG
(
NG)

Males
have
modified
anal
fin11
Viviparous2,18,28,31
NF
NF
Males
have
a
tubercular
penis4
8NG
(
NG)
NG
(
NG)

Males
have
modified
anal
fin11,15
11,26
NG
(
2)
24
(
3)

2224
(
NG)­
sex
unspecified
Ovoviviparous1,2,4,5,27
NF
1,2,4,6,8,9Male<
Female
3NG
(
at
birth)
NG
(
2)

Viviparous16,18,22,24,26,28,30,31
6,15,16,24NG
(
at
birth)
NG
(
at
birth)

Males
have
a
tubercular
penis4
8NG
(
NG)
NG
(
NG)

19
NG
(
at
birth)
NG
(
at
birth)

20
NG
(
at
birth)
NG
(
2)

215
(<
1)
<
5
(<
1)

229.2
(
NG)­
sex
unspecified
23,26
NG
(
at
birth)
NG
(
1)

TL,
FL*,
or
SL^
in
cm
(
age
in
yrs
discussed
as
viviparous,
but
ovoviviparity
was
disqualified
for
sake
of
discussion
(
p.

509)
17
Males
have
dark
head,
females
dark
underside
during
mating1,3
Breeding
Behavior
1st
maturity
Males
have
a
modified
anal
fin
appendage
for
copulation
6,7,14,17,18,19,21,24,28
Males
turn
dark
during
breeding
season1,2,7,8,17,18,20,25,26,28
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Embiotocidae
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17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AJ
AK
AL
AM
AN
AO
AP
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
NG
(<
1)
NG
(
2)
scales
NG
NG
NF
s.
California2,4,32,45,51,56,59
NG
(
NG)
NG
(
NG)
scales;
pelvic
bones
No
intrareader
comp./
multiple
structure
comp.

NG
(
NG)
NG
(
2)
NG
NG
NG
24
(
NG)­
sex
unspecified
NA
NA
NA
23.1
(
NG)­
sex
unspecified
NA
NA
NA
NF
NF
NG
(
NG)
NG
(
4)
NG
NG
NG
NF
Oregon26,59
NG
(
2)
27
(
4)
scales
Marginal
increment
analysis
intra/
interreader
comparison
California7,57
27.7
(
NG)­
sex
unspecified
NA
NA
NA
at
birth
NG
(
2)
NG
NG
NG
NF
British
Columbia
1,6,18,39,53,59
NG
(
at
birth)
NG
(
at
birth)
NG
NG
NG
Washington
1,8,21,38,47
NG
(
NG)
NG
(
1)
NG
NG
NG
Oregon
8,17,50
NG
(
at
birth)
NG
(
at
birth)
scales
tracked
first
annulus
growth
comparison
to
otoliths
California
1,15,40,49,52,53,54
NG
(
at
birth)
NG
(
2)
scales
No
multiple
scale
comparison
n.
California
8,10,42,54
5
(<
1)
<
5
(<
1)
direct
observations
NA
NA
c.
California
8,14
13.2
(
NG)­
sex
unspecified
NA
NA
NA
s.
California
5,8,35,36,54,59
NG
(
NG)
NG
(
NG)
NG
NG
NG
50%
maturity
s)
at
Fertiliza
REPRODUCTION
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Embiotocidae
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Fertilization
of
eggs
Spawning
females
present
Both
reproductive
states
present
Spawning
Frequency
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
(
per
year)

#####
#####
######
Once1,4
Once1,4
*****
*****
Once1,4
^^^^^
Once1,4
*****
*****
*****

*****
*****
*****

*****
*****

*****
*****

*****

ation
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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Embiotocidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BD
BE
BF
BG
BH
Size
specific
fecundity
equation
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
Larval
duration
9
33.4
(
NG)
2,54
(
17.8
SL)
to
113
(
28.5
SL)
1F=
0.335SL
­
49.65
None
None
15,16
33
(
up
to
113)
14
8.4
(
17.2
SL;
2yrs.)
to
62.4
(
33.7
SL;
9
yrs.)

20
20
(
2
yrs.)
to
70
(
4yrs.)

2347
(
NG)
to
80
(
NG)

3225
(<
25.4
?
L);
45
(>
25.4
?
L);
up
to
113
4123
(
25.4
?
L)
to
46
(>
25.4
?
L)

brood
size
increases
with
size/
age2,14,20
NF
NF
NF
None
None
4,
13.3
(
1
to
39)
5,42
8
(
NG)
to
45
(
NG)
1F=
0.232SL
­
40.86
None
None
9
25
(
NG)
14
8.7
(
21.8
SL;
4
yrs.)
to
33.7
(
29.2
SL;
8
yrs.)
3F*=
0.218TL
­
51.32
(
r2=
0.757)

17
7.4
(
25.8
TL;
3
yrs.)
to
33.7
(
36.7
TL;
8
yrs.)
3F*=
0.218TL
­
46.32
(
r2=
0.757)

3410
(
NG)
to
12
(
NG)

brood
size
increases
with
size/
age14,17
1
8.94
±
3.01
(
4
to
17)
3
6
(
9.0
SL)
to
16
(
13.9
SL)
1F=
0.0.171SL
­
9.2
None
None
210.8
±
3.5
(
NG)
6,43
5
(
NG)
to
36
(
NG)
5F=
0.0128FL2.735
(
r2=
0.79)

3
8.27
(
2
to
18)
14
5.8
(
8.23
SL;
1
yr)
to
20
(
12.2
SL;
6
yrs.)

8
10
(
5
to
17)
2239
6
(
9.3
FL;
1
yr.)
to
20
(
13.7
FL;
6
yrs.)

9
9.5
(
NG)
23
16
(
5
[
10
SL]
to
22
[
138
SL])

128
(
NG)
27
4
(
6.5
SL;
<
1
yr.)
to
24
(
12.8
SL;
3
yrs.)

28
7
(
9.5
to
9.9
BL?)
to
13
(
12.0
to
12.4)

30
6
(
NG)
to
10
(
NG)

335
(
NG)
to
17
(
NG)

376
(
10
?
L;
1
yr.)
to
15
(
13
?
L;
4
to
6)

40
20
(
15.2
?
L)

brood
size
increases
with
size/
age4,8,9,10,14,22,27,28,30,37,38
Fecundity
range
in
offspring
#(@
cm,
grams,
or
age)
DISPERS
Fecundity
Mean
(
range)
fecundity
in
offspring
#
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BI
BJ
BK
BL
BM
BN
BO
BP
BQ
Size
at
settlement
Size
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
Size
(
in
mm)
Habitat
Foraging
guild
Foraging
method
NF
NF
3
0.4
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
4
0.65
to
0.68
NF
NF
NF
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
NF
NF
NF
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
NF
NF
1,4,7
0.3
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
6
0.25
Eggs
Larvae
SAL
&
RECRUITMENT
Food
habits
Page
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Embiotocidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BR
BS
BT
BU
BV
BW
Fish
Size
Food
habits
hen
/
FrequenPredators
(
SL
mm)
Habitat
Foraging
guild
Foraging
method
NA
NA
44.5
to
maturity3
NF
NF
NF
>
40
to
maturity18
<
51
to
maturity39
NA
NA
NF
NF
NF
NF
NA
NA
50
to
maturity14
estuaries
and
bays9
carnivore
NF
NA
NA
34
to
50
to
maturity1
omnivore
NF
56
to
78
to
maturity2
29
to
maturity6
39
to
49
to
maturity7
eelgrass
beds16,20
34
to
38
to
maturity10
surfgrass
beds17
(
copepods
and
cyprid
larvae)
19
30
to
maturity22
43.7
to
maturity23,36
27
to
38
to
maturity24
33
(
27
to
38)
to
maturity26
27<
50
35
to
40
to
maturity28,32
36
to
maturity31
41
to
43
to
maturity33
27
to
36
to
maturity34
38
to
maturity38
(
small
fish;
crustaceans,
such
as
decapods
and
isopods;
polychaetes)
8
(
small
crustaceans,
such
as
copepods
and
amphipods;
algae;
polychaetes;

mollusks)
1,15,16,17,21
spring/
summer­
shallow
bays
and
estuaries;
mud/
sand
flats;
kelp
beds;
winter­
neritic1,6,15,19
HABITAT
ASSOCIATION
&
TR
Life
Stag
Juveniles
Page
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BX
BY
BZ
CA
Adults
Food
habits
When
/
Frequency
Predators
Habitat
Foraging
guild
NF
kelp
bass1
carnivore
NF
kelp
bass1
carnivore
NF
kelp
bass1
shallow
surf
and
sandy
bottoms
7,28,40,64
carnivore
gulls
and
cormorants8
diurnal3
kelp
bass1
(
micro)
carnivore13;
omnivorous38,50/
planktivory,
benthivory
(
crustaceans,
such
as
gammarids,
sand
crabs,
and
various
shrimps)
5,65
(
small
crustaceans,
such
as
gammarids,
barnacles;

polychaetes;
chaetognaths;
gastropods
and
bivalves;
fish
eggs;
algae?)
2,5,7,11,13,17,38,41,49,50,52,54,56,57,59,63,64,65
rocks
and
pilings;
surfzone;
bays
and
sandy
areas;
artificial
reefs5,10,14,19,28,32,40,48,52,60,63,64,65
(
mollusks;
small
fish,
such
as
smelt;
crustaceans,
such
as
Emerita
;
polychaetes)
5,28,61,65
shallow,
sandy
surf;
artificial
reefs;
rocky
outcrops
7,9,19,28,40,52,63,64
(
crustaceans,
especially
Emerita
;
mussels
and
other
bivalves;
echinoders,
such
as
Dendraster
)
2,5,6,10,15,44,60,65,66
kelp,
surfgrass,
and
eelgrass
beds;
bays
and
estuaries;
docks
&
pilings;
sandy/
muddy
bottoms;
deeper
water
in
winter;

artificial
reefs
5,6,7,14,17,18,19,20,22,26,34,35,36,39,41,44,45,47,48,50,51,52,53,54,55,56,61,64
ROPHIC
INTERACTIONS
ge
Page
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Embiotocidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CB
CC
CD
CE
CF
Haplotype
diversity
Foraging
method
When
/
Frequency
Predators
Substocks
(
Geographic
Area)

NF
NF
birds,
such
as
least
terns11,28
NF
NF
NF
NF
pinnipeds,
such
as
harbor
seals20,28
NF
NF
NF
NF
possibly
river
otters13
NF
NF
gulls
and
cormorants;
lingcod25
grazer/
picker­
nibbler2
diurnal/
nocturnal2,8,19
NF
picker/
persuer3
nocturnal10
suction
feeder16
diurnal13
possibly
river
otters13
harbor
seals14,21,23
great
blue
herons16,17,28
pinnipeds20
Caspian
and
Forster's
terns22,24
Brandt's
and
double­
crested
cormorants;

bald
eagles;
least
terns28
birds;
marine
mammals,
such
as
harbor
seals
and
porpoises;
fishes,
such
as
salmon,
sturgeon,
kelp
and
barred
sand
bass,
California
halibut;
rockfishes;
crabs
1,2,3,5,8,9,10,15,16,28
mainlander,
may
exist,
but
of
only
one
species
(
no
sub­
species)
3
Page
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CG
CH
CI
CJ
CK
CL
CM
Genetic
variance
Mean
number
of
%
Polymorphic
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
2NG
0.0573
(
range=
0.038
to
0.072)
NF
NF
GENETICS
Mean
heterozygosity
Nucleotide
diversity
(#)
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15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CN
CO
CP
CQ
CR
CS
CT
CU
CV
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
First
Recruitment
Full
Recruitment
Fishery
/
Total
Length
(
age)
Fishery
/
Total
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
2
0.013
NG
0.013
NF
NF
NF
NF
NF
NF
NF
NF
NF
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Embiotocidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CW
CX
CY
CZ
DA
DB
s
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Interactions
/
associations
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
NF
NF
NF
NF
NF
infested
by
parasites
from
the
genus
Lironeca
23
NF
NF
NF
NF
NF
direct
count
embryos
NF
NF
NF
NF
NF
NF
NF
NF
infested
by
parasites
from
the
genus
Lironeca
23
infested
by
parasites
from
the
families
Echinorhynchidae,
Polymorphidae,
Argulidae,

Amphicotylidae,
Lacistorhynchidae,

Phyllobothriidae,
Tetraphyllidea,
Bomolochidae,

Caligidae,
Ergasilidae,
Pennellidaem
Philichthyidae,
Acanthocolpidae,
Diplangidae,

Hemiuridae,
Heterophyidae,
Monorchidae,

Opecoelidae,
Zoogonidae,
Cymothoidae,

Capsalidae,
Gyrodactylidae,
Gyrodactyloidea,

Anisakidae,
Cucullanidae,
Myxidiidae22
parasites
Bomolchus
cuneatus
and
Ergasilus
turgidus
found
on
gills13
parasitized
by
Lasiotocus
sp.,
Genitocotyle
acirrus
,
and
other
digenetic
trematode11
parasitized
by
Genitocotyle
acirrus,

Pseudopecoelus
umbrinae,
and
several
other
parasites11
may
contain
intestinal
parasite
Telolecithus
pugetensis
4
parasitized
by
Diclidophora
and
other
internal
parasites9,10
infested
by
parasites
from
the
family
Philichthyidae22
infested
by
parasites
from
the
families
Opecoelidae
and
Diclidophoridae22
infested
by
parasites
from
the
families
Philichthyidae,
Diplangidae,
Monorchiidae,

Opecoelidae,
Cymothoidae,
and
Capsalidae22
MORTALITY
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Embiotocidae
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
DC
DD
DE
DF
DG
DH
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
NF
NF
Yes3,5,30,35
NF
NF
NF
NF
Yes5,35
NF
NF
NF
NF
Yes5,35
NF
3<
3
miles
NF
Yes4,5,8,9,10,11,13,14,20,21,22,24,26,29,32,33,35
Possibly2
solitary
at
night11
Transient
(
emigrates
out
of
bay)
21
Residential23
yearlings
non­
schooling
late
summer/
early
fall20
also
exhibit
diurnal
migrations
to
deeper
waters17,18,19
Transient
(
moves
to
deeper
waters
in
the
winter
and
spring)

1,3,4,5,6,7,13,16,17,18,19
Transient
(
moves
to
shallow
waters
to
mate
in
spring,
deeper
waters
in
the
fall/
winter)
20,22,26
4up
to
57
km,
but
usually
less
than
3.7
km
BEHAVIOR
2most
<
2
miles,
but
up
to
31
miles
Movement
Page
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68
69
70
71
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73
74
75
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77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
A
B
C
D
E
Embiotoca
jacksoni
1
11
to
50
Embiotoca
lateralis
1
11
to
45
Taeniotoca
lateralis
2
Hyperprosopon
argenteum
1
11
to
182
Hyperprosopon
ellipticum
1
silver
surfperch;
shiner1,2
11
to
110
B.
C.
(
Vancouver
I.)
to
c.
Baja
CA
(
Punta
San
Rosarito
and
Isla
Guadalupe)
1
striped
surfperch;
blue
perch4;
rainbow
perch;
squawfish;
crugnoli;
blue
surf
fish;

striped
surf
fish;
blue
seaperch1,2,4,5
s.
e.
Alaska
(
Wrangell)
to
n.
Baja
CA
(
Punta
Cabras)
1
distribution
gap
in
the
southern
California
Bight3
n.
CA
(
Fort
Bragg)
to
c.
Baja
CA
(
Punta
Abreojos
and
Isla
Guadalupe)
1
walleye
surfperch;
surf
fish;
silver
perch;

walleye
surf
fish;
walleye
seaperch;
china
pompano;
white
perch1,2,5
B.
C.
(
Brooks
Peninsula)
to
n.

Baja
CA
(
Rio
San
Vicente)
1
black
surfperch;
common
surf
fish;
bay
perch;
porgy;
blue
perch;
bay
blackperch;

pogie1,2,3,5
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71
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80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
F
G
H
I
JKLM
1Male:
W=
0.00003122SL3.044
1TL=
1.238(
SL)
+
2.911
2
39
s.
California
(
SL)
1
NG
NG
22.5
(
0.093)
NG
1Female:
W=
0.00001472SL3.2
5TL=
5.24
+
1.25(
SL);
r2=
1.0;
SE=
1.99
6Both:
W=
0.0000083SL3.31
5TL=
0.456
+
1.09(
FL);
r2=
0.996;
SE=
4.03
7Both:
W=
0.00002SL3.149
(
r2=
0.995)
5SL=
­
4.07
+
0.799(
TL);
r2=
1.0;
SE=
1.59
9Both:
W=
0.034TL(
in.)
3.137
(
r2=
0.99)
5SL=
­
5.7
+
0.882(
FL);
r2=
1.0;
SE=
1.45
5FL=
0.317
+
0.916(
TL);
r2=
0.996;
SE=
3.7
5FL=
6.58
+
1.13(
SL);
r2=
1.0;
SE=
1.64
7TL=
1.26SL
8TL=
7.0
=
1.2406SL
(
SE=
1.88)

4Male:
W=
­
618.95
+
46.03(
SL)
(
r
2=
0.884)
5TL=
5.35
+
1.26(
SL);
r2=
0.998;
SE=
2.78
2
38
c.
California
(
TL)
3
38.5
39.8
NG
0.263
4Female:
W=
­
2,118.62
+
106.66(
SL)
(
r
2=
0.941)
5TL=
1.03
+
1.09(
FL);
r2=
0.998;
SE=
2.67
10Female:
W=
0.000021FL3.007
(
r2=
0.964)
5SL=
­
3.93
+
0.794(
TL);
r2=
0.998;
SE=
2.21
10Male:
W=
0.000014FL3.079
(
r2=
0.975)
5SL=
­­
5.08
+
0.879(
FL);
r2=
0.998;
SE=
1.49
16Female:
W=
2.01046
+
3.16214
(
logFL
­
2.23934)
5FL=
­
0.695
+
0.917(
TL);
r2=
0.998;
SE=
2.45
16Male:
W=
1.82624
+
3.27164
(
logFL
­
2.18213)
5FL=
5.94
+
1.14(
SL);
r2=
0.998;
SE=
1.7
1Male:
W=
0.00009448SL3.238
1TL=
1.228(
SL)
+
4.32
1
30
s.
California
(
SL)
1
NG
NG
17.37
(
0.031)
NG
1Female:
W=
0.00001714SL3.114
3,9TL=
1.3214(
SL)
(
sizes
<
99
mm)
2
31
2,11Male:
W=
0.000011SL3.22343
3,9SL=
0.7568(
TL)
(
sizes
<
99
mm)

2,11Female:
W=
0.0000474SL2.90774
3,9TL=
1.2919(
SL)
(
sizes
>
100
mm)

2,11Both:
W=
0.000032SL2.99467
3,9SL=
0.774(
TL)
(
sizes
>
100
mm)

6Both:
W=
0.0000107SL3.22
3,9TL=
1.2925(
SL)

3,9SL=
0.7737(
TL)

5TL=
4.7
+
1.23(
SL);
r2=
0.996;
SE=
4.13
5TL=
0.036
+
1.09(
FL);
r2=
0.996;
SE=
3.5
5SL=
­
3.21
+
0.81(
TL);
r2=
0.996;
SE=
3.36
5SL=
­
3.98
+
0.888(
FL);
r2=
0.998;
SE=
2.54
5FL=
0.581
+
0.916(
TL);
r2=
0.996;
SE=
3.21
5FL=
4.87
+
1.12(
SL);
r2=
0.998;
SE=
2.86
*
14Male:
W=
0.00000783TL3.131
(
r2=
0.976)
5TL=
14.49
+
1.17(
SL);
r2=
0.99;
SE=
2.57
2
27
NF
NF
NF
NF
NF
*
14Female:
W=
0.0000214TL2.936
(
r2=
0.966)
5TL=
5.62
+
1.06(
FL);
r2=
0.996;
SE=
1.82
Page
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
N
O
P
Q
R
S
T
U
V
W
X
NG
0.39
(
0.001)
NG
NG
­
0.705
(
0.008)
NG
NG
NG
otoliths/
whole
No
No
0.263
NG
­
0.1
­
0.2
NG
NG
NG
NG
scale
impressions
No
intrareader
comparison/
multiple
scales
NG
0.538
(
0.001)
NG
NG
­
0.257
(
0.011)
NG
NG
NG
scales
No
No
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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Embiotocidae
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
Y
Z
AA
AB
AC
6
6,7
7
1
Dioecious
NF
84
(
at
Catalina
I.)

12
10
14
6
(
at
Santa
Cruz
I.)

16
9
3,5
2,4,5,67
to
10
NF
Dioecious
NF
14
6
(
at
Santa
Cruz
I.)

16
8
1,10
3,16
6
NF
Dioecious
promiscuous
(
polygynous?)
3
polygymous5
8
6
5
NF
Dioecious
NF
16
7
12
(
contains
validation
via
marginal
increment
and
Petersen's
length­
frequency
analyses)
Page
305
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Embiotocidae
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63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
AD
AE
AF
AG
AH
AI
Viviparous2,6,14,15,18,20,24,28,31
NF
NF
1,9NG
(
1)
12.1
(
1)

8NG
(
NG)
NG
(
NG)

14NG
(
NG)
14.6
(
2)

18NG
(
NG)
NG
(
1)

2214.6
(
NG)­
sex
unspecified
Males
have
a
tubercular
penis4
2312.7
(
1)
12.7
(
1)

Viviparous2,3,6,7,8,9,10,11,18,19,21,25,28,29,31
NF
3Male<
Female
Males
have
a
tubercular
penis4
4,5,26
23
to
25
(
2)
23
to
25
(
2)

Ovoviviparous27
7,24
NG
(
2)
NG
(
3)

8NG
(
NG)
NG
(
NG)

22NG
(
NG)
NG
(
NG)

Viviparous2,13,18,28,31
NF
4,7Male<
Female
Males
have
a
tubercular
penis4
8NG
(
NG)
NG
(
NG)

17,256.5
to
9.5^
(<
1)
9.5
(<
1)

22NG
(
NG)
NG
(
NG)

26
11.4
(
1)
11.4
(
1)

Breeding
males
darker
than
females1
Viviparous2,18,28,31
NF
5Male>
Female
Males
have
a
tubercular
penis4
8NG
(
NG)
NG
(
NG)

1214.1
(
1)
17.1
(
2)

modified
anal
fin
used
as
a
copulatory
organ23,26
male
anal
and
pelvic
fins
become
barred
and
shaded
yellow,
orange,

and
blue
during
breeding
season2,5
(
this
is
questioned
by
Isaacson
&

Isaacson)
13
Males
have
a
modified
anal
fin
appendage;
Females
have
drab
coloration
when
pregnant10,28
Males
with
yellow
pelvic
fins
and
modified
anal
fin
appendage;
females
with
dark
bars12
Males
with
modifed
anal
fin
appendage14,16
Males
with
modifed
anal
fin
appendage
for
copulation16,22,27
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
AJ
AK
AL
AM
AN
AO
AP
NG
(
NG)
NG
(
NG)
NG
NG
NG
NF
n.
California34
NG
(
NG)
NG
(
2)
NG
NG
NG
c.
California14
NG
(
NG)
NG
(
NG)
scales
NG
NG
s.
California
2,16,29,41
NG
(
NG)
NG
(
2)
NG
NG
NG
California54
23.4
(
NG)­
sex
unspecified
NA
NA
NA
NG
(
NG)
NG
(
NG)
NG
NG
NG
NG
(
3)
NG
(
3)
scales
NG
intrareader
comparison
NF
British
Columbia
3,6,13
NG
(
3)
NG
(
3)
scales
NG
NG
Washington9,27,47,53,59
NG
(
NG)
NG
(
3)
NG
NG
NG
Oregon9,20,50,59
28.6
(
NG)­
sex
unspecified
NA
NA
NA
c.
California11,12
s.
California2,43
NG
(
NG)
NG
(
1)
NG
NG
NG
NF
Oregon37,50
6.5
to
9.5
(<
1)
9.5
(<
1)
length
frequency;
otoliths
No
interreader
comparisons
s.
California7,22,31,37,46,54,55,58,59
20.6
(
NG)­
sex
unspecified
NA
NA
NA
NG
(
NG)
NG
(
NG)
NG
NG
NG
NG
(
NG)
NG
(
2)
NG
NG
NG
NF
Oregon28,59
14.1
(
1)
17.1
(
2)
scales
NG
interreader
comparison
s.
California48
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Embiotocidae
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63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
*****
Once1,2,3,4
#####
#####
#####
*****
#####
Once1,4
*****

#####
#####
#####
Once1,4
Once1,4
Page
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­
Embiotocidae
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
BD
BE
BF
BG
BH
1
11.28
±
3.01
(
7
to
18)
22
8
(
NG)
to
60
(
NG)
1F=
0.12SL
­
12.18
None
None
6,9,16
14.4
(
7
to
31)
14
4.3
(
14.9
SL;
2
yrs.)
to
30.7
(
29.6
SL;
7
yrs.)

1013
(
5
to
26)
21
7
(
15.3
SL;
2
yrs)
to
31
(
22.7
SL;
4
yrs.)

26
5
(
NG)
to
15
(
NG)

33
1
(
NG)
to
26
(
NG)

34up
tp
60
(
NG)

brood
size
increases
with
size/
age14,21,26,36
9
18
(
NG)
1
18
(
3
yrs.)
to
32
(
7
yrs.)
1F=
0.173SL
­
27.79
None
None
4
up
to
44
(
NG)

7
18.4
(
21.6
SL;
3
yrs.)
to
30.9
(
27.7
SL;
6
yrs.)

11,43
11
(
26.3
FL;
4
yrs.)
to
36
(
33.4
FL;
7
yrs.)

14
18
(
21.6
SL;
3
yrs.)
to
32
(
29.7
SL;
7
yrs.)

23
21
(
NG)
to
80
(
NG)

29,43
18
(
NG)
to
92
(
NG)

35
22
(
NG)
to
54
(
NG)

brood
size
increases
with
size/
age24,37
7,16
8.6
(
1
to
19)
4,13
5
(
NG)
to
12
(
NG)
1F=
0.135SL
­
9.0
None
None
9
9
(
NG)
14
6
(
11.0
SL;
1
yr.)
to
11
(
15.1
SL;
4
yrs.)
6F=
0.1352SL
­
9.0024
(
r2=
0.348)

23
7
(
NG)
to
12
(
NG)

25
1
(
10.5
SL)
to
19
(
15.3
SL)

33,38
5
(
NG)
to
12
(
NG)

40
3
(
NG)
to
5
(
NG)

brood
size
increases
with
size/
age14,25
9
10
(
NG)
14
3.4
(
12.8
SL;
2
yrs.)
to
16.3
(
18.4
SL;
5
yrs.)
1F=
0.229SL
­
25.8
None
None
18,43
4
(
17.4
TL;
2
yrs.)
to
17
(
18.4
TL;
3
yrs.)
4F=
0.178TL
­
25.962
(
r2=
0.462)

Page
309
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1167
5­
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­
Embiotocidae
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
BI
BJ
BK
BL
BM
BN
BO
BP
BQ
NF
NF
4
0.53
to
0.92
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
NF
NF
5
0.67
to
0.81
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
6,70.3
to
0.81
1
3
(
TL)
to
birth
size
NF
NF
2,6
0.7
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
1
1.5
to
2.0
to
birth
size
NF
NF
NF
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
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Embiotocidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
BR
BS
BT
BU
BV
BW
NA
NA
45
to
maturity6
microcarnivore9,11,14;
mesocarnivore13
picker/
non­
winnower2,4
52
to
maturity13
surfgrass
beds17
(
harpacticoids
and
tanaids)
19
54
to
maturity19
55
to
maturity20,21,25,29
45
to
57
to
maturity33
NA
NA
50
to
maturity4,30
carnivore
grazer2
70
to
maturity5
43
to
58
to
maturity8
55
to
maturity29
51
to
maturity34
NA
NA
40
to
maturity2,35
sandy
bottoms5
carnivore
NF
>
45
to
maturity17
surfgrass
beds17
(
small
crustaceans)
4
>
38
to
maturity38
NA
NA
40
to
maturity15
NF
NF
NF
(
crustaceans,
such
as
gammarids
and
isopods;
gastropods
and
bivalves;

bryzoans)
6,9,10,11,12,14
(
small
crustaceans,
such
as
gammarids
and
isopods;
polychaetes;
fish
eggs;

bivalves
and
gastropods;
sea
urchins)

2,3,5,6,16,17,18
kelp
bed
understory;
non­
planar
algal
patches,
occasionally
turf;

sandy
bottoms;
rocky
areas
3,5,7,8,10,11,12,13,14,18
bays
and
estuaries;
eelgrass;
kelp
bed
understory
and
algal
patches4,6,8,10,16
31
to
48
to
maturity
(
some
pugheaded);
48
to
56
(
normal)
11
picker;
larger
juveniles
also
may
winnow3
Page
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Embiotocidae
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
BX
BY
BZ
CA
diurnal1,2
microcarnivore8,13,31,34,48,53;
mesocarnivore36,46
birds,
such
as
comorants7
NF
kelp
bass1
microcarnivore44;
mesocarnivore46
NF
kelp
bass1
carnivore
NF
NF
carnivore
(
crustaceans,
such
as
gammarids)
5,65
kelp
forest;
midwater
over
sandy
bottoms;
surf
and
surfgrass;

pilings;
artificial
reefs
3,4,5,7,14,19,20,26,38,42,45,48,54,55,60,63,64
kelp
canopy;
pilings
and
docks;
rocky
shores
and
reefs;
reefs
with
foliose
red
algae;
eelgrass;
artificial
reefs
2,5,7,8,11,16,19,23,25,27,28,33,37,39,40,41,42,44,45,46,47,53,56,57,59,63,64
shallow
surf
and
sandy
bottoms;
rocky
outcroppings
and
jetties;
mouths
of
estuaries;
artificial
reefs
5,7,19,28,30,40,45,63,64
within
same
age
group;
smaller
individuals
are
gammarid
specialists,
largest
are
caprellid
specialists;
intermediate
sizes
generalists.
Caprellids
taken
mostly
in
Gelidium
beds47
(
crustaceans,
such
as
gammarids
and
mysids;
polychaetes;

bivalves;
small
fish;
squid
eggs)
1,2,5,10,15,17,21,45,52,56,57,59,65
kelp
bass;
grass
rockfish
1,2,3,4,5,6
(
crustaceans,
such
as
caprellids,
free
living
gammerids,
and
isopods;
gastropods
and
bivalves;
polychaetes;
tunicates;

ophiuroids;
fish
eggs;
bryozoans)
1,3,5,8,11,20,22,23,24,25,26,29,30,31,37,39,40,46,55,65
mixed
rock
and
sand;
turf;
kelp
forest
bottoms
with
or
without
foliose
algae;
eelgrass
and
surfgrass
beds;
rocky
coastlines;

pilings;
artificial
reefs;
bays
and
estuaries;
in
sandy
areas
north
of
Santa
Barbara;
soft
substrate
in
southern
CA
Bight
1,2,5,7,14,18,19,20,21,22,23,24,26,27,29,31,33,40,
42,45,46,48,49,51,52,54,55,57,58,60,62,64
(
polychaetes;
bryozoans;
mollusks;
crustaceans,
such
as
tubicolous
gammerids
and
caprellids;
ophiuroids;

gastropods;
external
fish
parasites)
2,4,5,10,12,13,14,17,18,19,21,23,24,25,26,29,

30,33,34,35,37,43,45,46,50,52,54,55,56,59,65
Page
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Embiotocidae
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
CB
CC
CD
CE
CF
grazer1
diurnal1,4,8,10,11,15,17,20
Santa
Cruz
Island;
mainland
California1
NF
searcher3
possibly
river
otters13
harbor
seals19
picker15
pinnipeds20
Caspian
and
Forster's
tern24
moray
eels26
grazer4
diurnal1,3,8,11,16
NF
NF
day
and
night17
Caspian
and
Forster's
tern24
picker11,12
Brandt's
cormorant28
picker/
persuer3
nocturnal1,14,10,18,19
birds3
NF
NF
kelp
bass;
scorpionsfish;
halibut12,18,26
possibly
river
otters13
pinnipeds20
Caspian
and
Forster's
tern24
NF
NF
possibly
river
otters13
NF
NF
Caspian
and
Forster's
tern24
harbor
seals;
Brandt's
and
double­
crested
cormorants28
Brandt's
and
double­
crested
cormorants;

least
terns28
electric
rays
and
sharks;
serranid
basses;

rockfishes;
pinnepeds
6,7,8
north
of
Santa
Monica
Bay;
south
of
Santa
Monica
bay;
Channel
Is.
and
mainland
2
birds;
electric
rays
and
sharks;
serranid
basses;
pinnepeds3,6,15
searcher/
picker/
oral
winnower
5,7,8,9,10,14,18
suction
feeder/
picker/
winnower16
searcher/
picker/
browser;

nonwinnower
5,7,8,10,16
Page
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Embiotocidae
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
CG
CH
CI
CJ
CK
CL
CM
NF
NF
NF
3NG
0.015
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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5­
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Embiotocidae
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52
53
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56
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
CN
CO
CP
CQ
CR
CS
CT
CU
CV
NF
NF
NF
NF
NF
NF
4
0.288
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
315
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5­
2220e
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Embiotocidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
CW
CX
CY
CZ
DA
DB
NG
eggs
NG
NG
possibly
with.
E.
lateralis
1,2,3,6
cleaned
by
Oxyjulis
californica
6
exploitation
competition
with
sea
urchins8
parasitized
by
copepod
family
Clavella15
competes
with
Hypsypops
rubicundus
for
food10
infested
by
parasites
from
the
genus
Lironeca
23
NF
NF
NF
NF
possibly
with.
E.
jacksoni
1,2,3,6
competes
with
E.
jacksoni
2
schools
with
D.
vacca
5
NF
NF
NF
NF
NF
infested
by
parasites
from
the
genus
Lironeca
23
NF
NF
NF
NF
NF
infested
by
parasites
from
the
families
Bomolochidae,
Lernaeopodidae,
Pennellidae,

Philichthyidae,
Acanthocolpidae,
Diplangidae,

Monorchiidae,
Opecoelidae,
Zoogonidae,
and
Cymothoidae22
infested
by
parasites
from
the
families
Argulidae,

Lernaeopodidae,
Philichthyidae,
Diplangidae,

Hemiuridae,
Monorchiidae,
Opecoelidae,

Zoogonidae,
Myxidiidae,
and
Sinuolineidae22
infested
by
parasites
from
the
families
Argulidae,

Philichtyidae,
Acanthocolpidae,

Cryptogonimidae,
Haplosplanchnidae,

Monorchiidae,
Opecoelidae,
Cymothoidae,

Diclidophoridae,
and
Gyrodactylidae22
parasitized
by
Diclidophora
and
other
internal
parasites10
infected
by
ectoparasites
from
the
families
Caligidae
and
Lernaeidae6
competes
with
E.
lateralis
for
resources;
competition
may
affect
growth
and
birth
rates
9
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants16
parasitized
by
Diclidophora
and
other
internal
parasites10
parasitized
by
Genitocotyle
acirrus
,

Schikhobalotrema
girellae
,
and
other
digenetic
trematodes11
kept
from
shallow
algal
areas
by
competitive
interference
with
dominant
E.
lateralis
4,8,11
cleaned
by
Oxyjulis
californica
and
Brachyistius
frenatus
7,17
parasitized
by
Telolecithus
pugetensis,

Neozoogonus
californicus
,
and
others
trematodes11
competes
with
E.
jacksoni
for
resources;
competition
may
affect
growth
and
birth
rates
9
constant
competetitive
interference
and
seasonal
exploitation
competition
with
E.
jacksoni
5
seasonal
competition
with
E.
jacksoni
for
Gelidium
feeding
habitat
in
shallow
reef
areas
(<
5
m)
7,11
negatively
affect
by
presence
of
Macrocystis
on
the
understory18
constant
competitive
interference
and
seasonal
exploitation
competition
with
E.
lateralis
5
parasitized
by
Telolecithus
pugetensis,

Neozoogonus
californicus
,
and
others
trematode11
seasonal
competition
with
E.
lateralis
for
Gelidium
feeding
habitat
in
shallow
reef
areas
(<
5
m)
7,11
removes
external
parasites
from
conspecifics
and
other
fishes1,7,12,20
competitive
interference
with
E.
jacksoni
,
keeping
E.

jacksoni
out
of
the
shallow
algal
areas
4,8,11
indirectly
derives
positive
benefit
from
presence
of
Macrocystis
on
the
understory18
Page
316
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5­
2220e
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Embiotocidae
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
DC
DD
DE
DF
DG
DH
NF
Residential2,8,23
NF
Yes2,5,7,8,9,12,14,25,27,35
Yes3,4
2(~
50
m)
No19
1,3<
3
miles
Residential2,21
Possibly1
Yes2,5,6,13,32,35
Yes1,4
winter
migration
to
deeper
water19
2(~
50
m)
somewhat
solitary3
Yes­
defend
foraging
microhabitats6
1<
6
miles
Transient6,23
NF
Yes1,2,5,9,11,12,14,15,30,34,35
NF
3<
3
miles
NF
NF
NF
Yes3,5,25,35
NF
winter
migration
to
lower
estuary
and
ocean25
school
during
the
day,
disperse
offshore
at
night31
Yes,
temporarily
during
breeding
season,

but
also
present
year
round
5
3(
292
m2
[
187
to
481
m2]:
during
breeding
season);
(
25.4
m2
[
21.2
to
30.7
m2]:
rest
of
year)
Page
317
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5­
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
A
B
C
D
E
Hypsurus
caryi
1
11
to
40
Phanerodon
furcatus
1
12
to
70
Rhacochilus
toxotes
1
13
to
47
Racochilus
vacca
1
11
to
209
pile
surfperch;
forktail
perch;
porgee;
white
n.
CA
(
Cape
Mendocino)
to
n.

Baja
CA
(
Bahia
San
Carlos)
1
s.
e.
Alaska
(
Wrangell)
to
c.
Baja
n.
CA
(
Mendocino
County)
to
s.

Baja
CA
(
Cabo
Thurloe
and
Isla
Guadalupe)
1
white
surfperch;
Pacific
white
perch;

splitail
perch;
forktail
perch;
white
surf
perch;

shiner1,2,5
rubberlip
surfperch;
porgee;
alfione;
pile
perch;
sprat;
liverlip;
buttermouth
1,5
B.
C.
(
Vancouver
I.)
to
c.
Baja
CA
(
Punta
Escarpada)
1
rainbow
surfperch;
rainbow
seaperch;

striped
perch;
bugara;
moharra1,2,5
Page
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1167
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Embiotocidae
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115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
F
G
H
I
JKLM
*
14Immatures:
W=
0.00000652TL3.148
(
r2=
0.937)
5SL=
­
10.82
+
0.843(
TL);
r2=
0.99;
SE=
2.18
5SL=
­
6.88
+
0.897(
FL);
r2=
0.996;
SE=
1.41
5FL=
­
4.47
+
0.941(
TL);
r2=
0.996;
SE=
1.72
5FL=
8.32
+
1.11(
SL);
r2=
0.996;
SE=
1.56
12SL*=
0.782TL
­
1.155
(
r2=
0.996)

12FL*=
0.9004TL
+
2.429
(
r2=
0.974)

6Both:
W=
0.000002SL3.56
5TL=
8.7
+
1.19(
SL);
r2=
0.998;
SE=
3.12
1
30
NF
NF
NF
NF
NF
5TL=
1.3
+
1.09(
FL);
r2=
0.998;
SE=
2.6
5SL=
­
6.7
+
0.836(
TL);
r2=
0.996;
SE=
2.61
5SL=
­
6.05
+
0.913(
FL);
r2=
1.0;
SE=
1.3
5FL=
­
0.735
+
0.916(
TL);
r2=
0.998;
SE=
2.39
5FL=
6.78
+
1.09(
SL);
r2=
1.0;
SE=
1.42
1Male:
W=
0.00002655SL2.978
1TL=
1.296(
SL)
+
1.584
1
46
s.
California
(
SL)
1
NG
NG
20.2
(
0.056)
NG
1Female:
W=
0.00002254SL3.013
3,9TL=
1.3571(
SL)
(
sizes
<
99
mm)

2,11Male:
W=
0.0000185SL3.07131
3,9SL=
0.7368(
TL)
(
sizes
<
99
mm)

2,11Female:
W=
0.0000147SL3.11733
3,9TL=
1.3526(
SL)
(
sizes
>
100
mm)

2,11Both:
W=
0.00000765SL3.24848
3,9SL=
0.7393(
TL)
(
sizes
>
100
mm)

6Both:
W=
0.0000141SL3.12
3,9TL=
1.3504(
SL)

7Both:
W=
0.000019SL3.065
(
r2=
0.996)
3,9SL=
0.7405(
TL)

*
17Male:
W=
0.0000891TL2.98
5TL=
5.58
+
1.27(
SL);
r2=
0.978;
SE=
6.16
*
17Female:
W=
0.000117TL3.06
5TL=
10.39
+
1.09(
FL);
r2=
0.974;
SE=
7.51
5SL=
­
0.741
+
0.774(
TL);
r2=
0.978;
SE=
4.82
5SL=
­
8.02
+
0.927(
FL);
r2=
0.996;
SE=
1.89
5FL=
­
3.95
+
0.897(
TL);
r2=
0.974;
SE=
6.82
5FL=
9.24
+
1.08(
SL);
r2=
0.996;
SE=
2.03
6Both:
W=
0.00000457SL3.36
5TL=
2.67
+
1.26(
SL);
r2=
1.0;
SE=
2.99
1
47
NF
NF
NF
NF
NF
5TL=
­
7.53
+
1.14(
FL);
r2=
1.0;
SE=
1.57
5SL=
­
2.03
+
0.79(
TL);
r2=
1.0;
SE=
2.37
5SL=
­
8.51
+
0.909(
FL);
r2=
1.0;
SE=
21.3
5FL=
6.61
+
0.874(
TL);
r2=
1.0;
SE=
1.37
5FL=
9.44
+
1.1(
SL);
r2=
1.0;
SE=
2.34
6Both:
W=
0.00000017SL4.18
(<
80
mm)
5TL=
4.82
+
1.29(
SL);
r2=
0.998;
SE=
4.6
2
44
Oregon
(
TL)
4
43.2
49
NG
NG
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Embiotocidae
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105
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107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
N
O
P
Q
R
S
T
U
V
W
X
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NG
0.425
(
0.001)
NG
NG
­
0.782
(
0.003)
NG
NG
NG
scales
No
No
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NG
NG
NG
NG
NG
NG
NG
NG
scales
No
No
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Embiotocidae
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105
106
107
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109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
Y
Z
AA
AB
AC
NF
NF
NF
Dioecious
NF
1
3,6,16
7
5
Dioecious
NF
13
8
NF
2,16
7
to
10
NF
Dioecious
NF
13,14­
juveniles
2,67
to
10
NF
Dioecious
NF
Page
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1167
5­
2220e
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
AD
AE
AF
AG
AH
AI
2216.5
(
NG)­
sex
unspecified
26
15.2
(
1)
17.8
(
2)

Viviparous2,14,18,24,28,31
NF
NF
Males
have
a
tubercular
penis4
14,26
NG
(
NG)
13.1
(
NG)

2213.1
(
NG)­
sex
unspecified
Viviparous2,12,18,28,29,31
NF
4Male<
Female
Males
have
a
tubercular
penis4
8NG
(
NG)
NG
(
NG)

Ovoviviparous27
22NG
(
NG)
NG
(
NG)

Viviparous2,3,18,23,28,31
NF
NF
Males
have
a
tubercular
penis4
22NG
(
NG)
NG
(
NG)

Viviparous2,3,10,11,18,28,29,31
NF
NF
Males
have
a
tubercular
penis4
8NG
(
NG)
NG
(
NG)

Males
turn
darker
during
breeding
season29
Males
with
modifed
anal
fin
appendage
for
copulation26
Males
with
modifed
anal
fin
appendage
for
copulation14
Page
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1167
5­
2220e
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Embiotocidae
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102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
AJ
AK
AL
AM
AN
AO
AP
19.7
(
NG)­
sex
unspecified
NA
NA
NA
NG
(
NG)
NG
(
NG)
NG
NG
NG
NG
(
NG)
NG
(
NG)
scales
NG
NG
NF
n.
California34,59
19.7
(
NG)­
sex
unspecified
NA
NA
NA
NG
(
NG)
NG
(
2)
NG
NG
NG
NF
n.
California30,59
25.4
(
NG)­
sex
unspecified
NA
NA
NA
c.
California14,33,54
s.
California
24,54,59
California53
29
(
NG)­
sex
unspecified
NF
NF
NF
NF
California3,7,45
NG
(
NG)
NG
(
4)
NG
NG
NG
NF
British
Columbia3
Page
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1167
5­
2220e
­
Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Once1,4
Once1,4
Once1,4
Once1,4,5
Page
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1167
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2220e
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
BD
BE
BF
BG
BH
brood
size
increases
with
size/
age14
6,9
11.6
(
9
to
22)
21
9
(
13.1
SL)
to
22
(
21.0
SL)
1F=
0.154SL
­
15.18
None
None
16
NG
(
5
to
22)
23,24
8
(
NG)

brood
size
increases
with
size/
age21
1
12.51
±
6.49
(
5
to
34)
offspring
15
7.3
(
4
to
11);
6.8
(
3
to
11)
1F=
0.062SL
­
3.75
None
None
11,16
NG
(
8
to
33)
14
10.1
(
16
SL;
2
yrs.)
to
28.5
(
23.4
SL;
7
yrs.)
2F=
0.115SL
­
13.07
19
8
(
19.6
SL)
to
33
(
26.2
SL)

23
10
(
NG)
to
23
(
NG)

37,38
12.1
(
16.5
to
21.5)
to
24.6
(
25.2)

brood
size
increases
with
size/
age14,19
NF
32
21
(
NG)
1F=
0.112SL
­
18.37
None
None
1,5,38
12
(
4
yrs.)
to
60
(
10
yrs.)
1F=
0.347SL
­
66.05
None
None
5,16
42
(
30
to
50)
for
600
g
mother;
65
Page
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1167
5­
2220e
­
Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
BI
BJ
BK
BL
BM
BN
BO
BP
BQ
NF
NF
NF
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
NF
NF
NF
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
NF
NF
NF
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
NF
NF
NF
No
external
eggs
NA
No
external
larval
stage
NA
NA
NA
Page
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1167
5­
2220e
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
BR
BS
BT
BU
BV
BW
NA
NA
55
to
maturity19
NF
NF
48
to
maturity31
NA
NA
34
to
maturity6
kelp
beds;
bays
and
estuaries3,6
NF
NF
44
to
53
to
maturity12
surfgrass
beds17
43.5
to
maturity16,34
NA
NA
NF
kelp
bed
understory3,8,18
carnivore
NF
NA
NA
13
to
maturity9
carnivore
grazer1
(
polychaetes;
bryozoans;
amphipods;

mussels)
21
over
sandy
bottoms;
kelp
bed
understory2,3,8
Page
327
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1167
5­
2220e
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
BX
BY
BZ
CA
NF
NF
carnivore
NF
kelp
bass1
(
micro)
carnivore13,42/
benthivorous
carnivore64
NF
kelp
bass1
(
meso)
carnivore9
NF
kelp
bass1
mesocarnivore8
rocky
shores;
pilings;
kelp
forest
bottoms;
surfgrass
beds;
soft­
(
crustacenas;
mollusks,
such
as
gastropods;
algae;
small
fish;
fish
eggs)
2,3,5,10,12,15,25,26,44,45,52,56,65
(
crustaceans,
such
as
gammerids
and
isopods;
ophiuroids;

mollusks;
polychaetes)
4,5,25,56,65
(
substrate
oriented
crustaceans,
such
as
gammerids,

isopods,
and
decapods;
ophiuroids;
mollusks,
such
as
gastropods
and
bivalves;
polychaetes;
bryozoans;
fish
eggs)

2,4,5,10,11,12,13,17,18,42,52,56,57,65
rocky
shores
and
reef
bottoms;
midwater;
kelp
beds
and
forest;
artficial
reefs;
soft
substrata
5,7,13,14,19,20,21,24,42,45,49,52,54,58,60,63,64
piers
and
docks;
sandy/
muddy
shallows;
eelgrass
and
surfgrass
beds;
rocky
reefs;
sand­
rock
interface;
artificial
reefs;

kelp
bed
bottoms,
occasionally
midwater
5,7,8,14,15,18,19,20,21,23,24,42,43,45,47,48,49,52,53,54,55,60,62,63,64
kelp
forest
edge
near
bottom;
holdfast
area;
rocky
reef;

surfgrass;
artififcial
reefs;
soft
substrata
7,12,14,19,20,23,24,40,42,45,49,60,63,64
Page
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1167
5­
2220e
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
CB
CC
CD
CE
CF
grazer1
diurnal1,4,11
NF
NF
NF
searcher;
oral
winnower5
grazer1
diurnal1,4,8,14
NF
NF
searcher3
picker13
possibly
river
otters13
plucker18
pinnipeds20
Caspian
terns22
sea
lions27
nocturnal1,3,5,6,11
birds3
NF
NF
diurnal/
nocturnal12
possibly
river
otters13
pinnipeds20
Caspian
terns24
harbor
seal;
Brandt's
cormorants28
grazer1,4
diurnal1,4,6,7,11
birds;
rockfishes3,8
Santa
Cruz
Island;
mainland
California1
NF
Brandt's
and
double­
crested
cormorants;

least
terns28
searcher/
plucker;
oral
winnower5,18
birds;
rockfishes
and
seabasses,
such
as
barred
sand
bass3,4,5
Page
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1167
5­
2220e
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
CG
CH
CI
CJ
CK
CL
CM
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1
6
NF
NF
NF
NF
Page
330
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1167
5­
2220e
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Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
CN
CO
CP
CQ
CR
CS
CT
CU
CV
NF
NF
NF
NF
NF
NF
3
0.117
0
0.117
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1
0.0387
0
0.0387
Page
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1167
5­
2220e
­
Embiotocidae
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
CW
CX
CY
CZ
DA
DB
eggs
NF
NF
NF
NF
removes
ectoparasites
from
fish
12
NF
NF
NF
NF
NF
parasitized
by
copepod
Argulus
borealis
13
parasitized
by
copepod
family
Clavella15
infested
by
parasites
from
the
genus
Lironeca
23
NF
NF
NF
NF
NF
mutual
association
with
R.
vacca
3
direct
counting
eggs
NG
NG
NF
cleaned
by
Oxyjulis
californica
2,6,7
infested
by
parasites
from
the
families
Hemiuridae,
Opecoelidae,
Diclidophoridae,
and
Gyrodactylidae22
infested
by
parasites
from
the
families
Pennellidae,
Philichthyidae,
Zoogonidae,
and
Myxidiidae22
infested
by
parasites
from
the
families
Argulidae,

Bomolochidae,
Caligidae,
Lernaeopodidae,

Pennellidae,
Diplangidae,
Monorchiidae,

Opecoelidae,
Zoogonidae,
Cymothoidae,

Diclidophoridae,
and
Myxiidae22
infested
by
parastites
from
the
families
Caligidae,
Philichthyidae,
Acanthocolpidae,

Monorchiidae,
Zoogonidae,
and
Ceratomyxidae22
cleaned
by
Phanerodon
atripes
and
Oxyjulis
californica
2,6
parasitized
by
digenetic
trematode
Neozoogonus
californicus
11
may
be
indirectly
negatively
affected
by
chlorine
discharge
plums
from
power
plants16
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants16
parasitized
by
Telolecithus
pugetensis,

Neozoogonus
californicus
,
and
others
trematodes11
removes
ectoparasites
from
fish
such
as
Sebastes
mystinus
and
Chromis
punctipinnis
6,14,21
parasitized
by
Telolecithus
pugetensis,

Neozoogonus
californicus
,
and
others
trematodes11
share
same
resources
as
S.
politus
and
Phanerodon
furcatus
19
infected
by
ectoparasites
from
the
families
Caligidae
and
Lernaeidae;
also
monogenetic
trematodes6
Page
332
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5­
2220e
­
Embiotocidae
101
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105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
DC
DD
DE
DF
DG
DH
NF
Transient11,12,14
NF
Yes3,5,9,12,35
NF
aggregate
for
breeding
in
fall
25
NF
Residential3,10,23
NF
Yes2,3,8,9,11,12,17,18,30,35
NF
Adults­
occasionally23,29
Yes­
juveniles29
NF
NF
NF
Yes2,5,12,17,35
NF
not
usually28
NF
Residential2,23,24
NF
Yes2,5,7,9,12,13,16,17,25,30,32,35
Yes1
Page
333
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2220e
­
Embiotocidae
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152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
A
B
C
D
E
Damalichthys
vacca
2
1­
Robins
et
al.
1991
1­
Eschmeyer
1998
1­
Gates
&
Frey
1974
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Cassano
1998
2­
Robins
et
al.
1991
2­
Haldorson
&
Moser
1979
3­
Limbaugh
1955
4­
Morgan
1961
5­
Roedel
1948
6­
Fitch
&
Lavenberg
1975
perch;
splittail
perch;
pile
seaperch;
dusky
seaperch4;
silver
perch1,2,4,5
CA
(
Bahia
Playa
Maria
and
Isla
Guadalupe)
1
Page
334
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
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179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
F
G
H
I
JKLM
6Both:
W=
0.0000269SL3.03
(>
80
mm)
5TL=
2.31
+
1.14(
FL);
r2=
0.992;
SE=
7.35
9Both:
W=
0.027TL(
in.)
3.039
(
r2=
0.98)
5SL=
­
3.24
+
0.774(
TL);
r2=
0.998;
SE=
3.57
*
12Male:
W=
0.00000871TL3.08
(
r2=
0.9996)
5SL=
­
6.43
+
0.905(
FL);
r2=
0.998;
SE=
2.67
*
12Female:
W=
0.00000513TL3.19
(
r2=
0.993)
5FL=
0.008
+
0.871(
TL);
r2=
0.992;
SE=
6.42
5FL=
7.4
+
1.1(
SL);
r2=
0.998;
SE=
2.95
10SL=
0.75TL
­
1.51
1­
Eckmayer
1979
1­
Eckmayer
1979
1­
Love
et
al
2000
1­
Eckmayer
1979
2­
Anderson
1969
2­
Carlisle
et
al.
1960
2­
Baltz
1984
2­
Carlisle
et
al.
1960
3­
Odenweller
1971
3­
Anderson
1969
3­
Bryant
1978
4­
Swedberg
1965
4­
Odenweller
1971
4­
Wares
1971
5­
Odenweller
1975
5­
Karpov
&
Kwiecien
1988
6­
Quast
1968b
6­
Odenweller
1975
7­
Antrim
1981
7­
DeMartini
et
al.
1994
8­
Bayer
1985
8­
Isaacson
&
Isaacson
1966
9­
DeMartini
et
al.
1994
9­
Anderson
&
Bryan
1970
10­
Gnose
1967
10­
Wares
1971
11­
Anderson
&
Bryan
1970
11­
Bennet
&
Wydoski
1977
12­
Wares
1971
12­
Wydoski
&
Bennett
1973
13­
Bennet
&
Wydoski
1977
13­
Wilson
&
Millemann
1969
14­
Wydoski
&
Bennet
1973
15­
Suomela
1931
16­
Sivalingam
1953
17­
Beardsley
1969
Page
335
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
N
O
P
Q
R
S
T
U
V
W
X
Page
336
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
Y
Z
AA
AB
AC
13,16
9
to
10
1­
Anderson
et
al.
1969;
Anderson
&
Bryan
1970
1­
Emmett
et
al.
1991
1­
Isaacson
&
Isaacson
1966
1­
Carlisle
et
al.
1960
1­
Darling
et
al.
1980
2­
Odenweller
1971
2­
Fritzsche
et
al.
1989
2­
Bane
1970
2­
Hubbs
1917
2­
Hubbs
1921
3­
Swedberg
1965
3­
Anderson
1969
3­
Bane
1970
3­
Rechnitzer
&
Limbaugh
1952
4­
Odenweller
1975
4­
Swedberg
1965
4­
DeLeon
1999b
4­
Warner
&
Harlan
1982
5­
Gnose
1967
5­
Gnose
1967
5­
Beardsley
1969
5­
DeLeon
1999b
6­
Isaacson
&
Iasaacson
1966
6­
Baltz
1984
7­
Bennet
&
Wydoski
1977
7­
Holbrook
&
Schmitt
1984
8­
Wydoski
&
Bennet
1973
8­
Schmitt
&
Holbrook
1985
9­
Bane
1970
9­
Suomela
1931
10­
DeMartini
et
al.
1983
10­
Fitch
&
Lavenberg
1971
11­
Suomela
1931
11­
Sivalingam
1953
12­
Sivalingam
1953
12­
Fitch
&
Lavenberg
1975
13­
Garrison
&
Miller
1982
13­
DeLeon
1999b
14­
Wares
1971
14­
Holbrook
&
Schmitt
1995
15­
Cassano
1998
16­
Love
1996
Page
337
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
AD
AE
AF
AG
AH
AI
Ovoviviparous27
10,24,25,26
NG
(
2?)
NG
(
2)

22NG
(
NG)
NG
(
NG)

1­
Emmett
et
al.
1991
1­
Emmett
et
al.
1991
1­
Feder
et
al
1974
1­
DeMartini
1969
2­
Tarp
1952
2­
Hart
1973
2­
De
Martini
1969
2­
Carlisle
et
al.
1960
3­
Fritzsche
et
al.
1989
3­
Swedberg
1965
3­
Carlisle
et
al.
1960
3­
Hart
1973
4­
Gardiner
1978
a,
b
4­
Anderson
1968;
Anderson
&
Bryan
1970
4­
Blake
1869
4­
Fritzsche
et
al
in
Leet
et
al
1992
5­
Turner
1938
5­
Wydoski
&
Bennett
1973
5­
Carlisle
et
al.
1964
5­
Swedberg
1965
6­
Holbrook,
Schmitt,
&
Ambrose
1990
6­
Eigenmann
1892
6­
Wiebe
1968a
6­
Turner
1938
7­
Ingermann
&
Terwilliger
1981
7­
DeMartini
et
al.
1983
7­
Wiebe
1968b
7­
Gnose
1967
8­
Ingermann
et
al.
1984
8­
Gordon
1965
8­
Shaw
&
Allen
1977
8­
Baltz
1984
9­
Ingermann
&
Terwilliger
1982
9­
Boothe
1967
9­
Turner
et
al.
1969
9­
Isaacson
&
Isaacson
1966
10­
Webb
&
Brett
1972a
10­
Gnose
1967
10­
Wares
1971
11­
Webb
&
Brett
1972b
11­
Hubbs
1934
11­
Bennet
&
Wydoski
1977
12­
Banerjee
1971
12­
Rechnitzer
&
Limbaugh
1952
12­
Wydoski
&
Bennett
1973
13­
Engen
1968
13­
Isaacson
&
Isaacson
1966
13­
Triplett
1960
14­
Behrens
1977
14­
Anderson
&
Bryan
1970
14­
Behrens
1977
15­
Holbrook
&
Schmitt
1988
b
15­
Bennet
&
Wydiski
1977
15­
Bane
1970
16­
Wilson
&
Millemann
1969
16­
Hixon
1980
16­
Wilson
&
Millemann
1969
17­
Turner
1947
17­
Bane
1970
17­
DeMartini
et
al.
1983
18­
Eigenmann
1892
18­
Eigenmann
1892
18­
Schmitt
&
Holbrook
1985
19­
Cowen
1979
19­
Suomela
1931
19­
Suomela
1931
20­
Lagios
1965
20­
Gordon
1965
20­
Gordon
1965
21­
Blanco
1938
21­
Shaw
1971
21­
Shaw
1971
22­
Wiebe
1967
22­
Blanco
1938
22­
Karpov
et
al.
1995
23­
Fitch
&
Lavenberg
1971
23­
Hixon
1981
23­
Fitch
&
Lavenberg
1975
24­
Shrode
et
al.
1983
24­
Wiebe
1967
24­
Garrison
&
Miller
1982
25­
Sivalingam
1953
25­
Borton
1982
25­
DeLeon
1999b
26­
Boothe
1967
26­
Limbaugh
1955
26­
Love
1996
27­
Garrison
&
Miller
1982
27­
Sivalingam
1953
Males
turn
darker
during
breeding
season29
Male
pelvic
and
anal
fins
bright
yellow
and
snouts
with
black
spots
during
breeding
season9
Page
338
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
AJ
AK
AL
AM
AN
AO
AP
NG
(
NG)
NG
(
3)
scales
No
No
Oregon25,53,59
32.8
(
NG)­
sex
unspecified
NA
NA
NA
California6,54
c.
California23
s.
California19,54,59
1­
Young
1963
1­
Emmett
et
al
1991
2­
Eigenmann
1891
3­
Fritzsche
et
al.
1989
4­
Carlisle
et
al.
1960
5­
Odenweller
1971,1975
6­
Hart
1973
7­
Fritzsche
et
al.
1992
8­
Monaco
et
al.
1990
9­
Swedberg
1965
10­
Boothe
1967
11­
Bryant
1978
12­
Miller
&
Geibel
1973
13­
Fraser
1923
14­
Antrim
1981
15­
Turner
1938
16­
Terry
and
Stephens
1976
17­
Bayer
1985
18­
Wiebe
1968a
19­
Turner
et
al.
1969
20­
Gnose
1967
21­
Anderson
&
Chew
1972
22­
Rechnitzer
&
Limbaugh
1952
23­
Wales
1929
24­
Goldberg
1978
25­
Wares
1971
26­
Bennet
&
Wydoski
1977
27­
Blanco
1938
Page
339
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
^^^^^
^^^^^
*****
*****
^^^^^
1­
Baltz
1984
2­
Holbrook
&
Schmitt
1984
3­
Schmitt
&
Holbrook
1985
4­
Haldorson
1980
5­
Haldorson
1973
Page
340
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
BD
BE
BF
BG
BH
14
18
(
22.4
SL;
4
yrs.)
to
52
(
33.2
SL;
10
yrs.)

9
30
(
NG)
16
61
(
44.9
TL;
9
yrs.)

13NG
(
7
to
61)
brood
size
increases
with
size/
age12,14,16,38
14
NG
(
12
to
50)

1­
Antrim
1981
1­
Fritzsche
et
al.
1989
1­
Baltz
1984
2­
Darling
et
al.
1980
2­
Carlisle
et
al.
1960
2­
Goldberg
1978
3­
Bane
1970
3­
Odenweller
1971,1975
3­
Bennet
&
Wydoski
1977
4­
DeMartini
1987
4­
Hart
1973
4­
Wydoski
&
Bennett
1973
5­
Webb
&
Brett
1972a
5­
Fritzsche
et
al
in
Leet
et
al
1992
5­
Wilson
&
Millemann
1969
6­
Behrens
1977
6­
Crooke
in
Leet
et
al.
1992
6­
DeMartini
et
al.
1983
7­
DeMartini
et
al.
1983
7­
Swedberg
1965
8­
Gordon
1965
8­
Turner
1938
9­
Karpov
et
al.
1995
9­
Darling
et
al.
1980
10­
Fitch
&
Lavenberg
1975
10­
Wiebe
1968b
11­
Garrison
&
Miller
1982
11­
Gnose
1967
12­
DeLeon
1999b
12­
Webb
&
Brett
1972a
13­
Wares
1973
13­
Rechnitzer
&
Limbaugh
1952
14­
Haldorson
1973
14­
Baltz
1984
15­
Carlisle
1960a
15­
Goldberg
1978
16­
Love
1996
16­
Wares
1971
17­
Bennet
&
Wydiski
1977
18­
Wydoski
&
Bennett
1973
19­
Banerjee
1971
20­
Triplett
1960
21­
Behrens
1977
22­
Wilson
&
Millemann
1969
23­
Eigenmann
1892
24­
Agassiz
1853
25­
DeMartini
et
al.
1983
26­
Schmitt
&
Holbrook
1985
27­
Suomela
1931
(
)
g
(
45
to
80)
for
a
1200
g
mother
Page
341
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
BI
BJ
BK
BL
BM
BN
BO
BP
BQ
1­
Emmett
et
al.
1991
1­
Wang
1981
2­
Engen
1968
3­
Triplett
1960
4­
Eigenmann
1892
5­
Blanco
1938
6­
Wang
1981
7­
Garrison
&
Miller
1982
Page
342
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
BR
BS
BT
BU
BV
BW
76
to
85.7
to
maturity35,37
eelgrass
beds16
surfgrass
beds17
1­
Strategic
Assessment
Branch
1990
1­
Emmet
et
al.
1991
1­
Emmet
et
al.
1991
1­
Hueckel
&
Stayton
1982
2­
Hart
1973
2­
Terry
and
Stephens
1976
2­
Hart
1973
2­
Schmitt
&
Holbrook
1984a
3­
Fritzsche
et
al
in
Leet
et
al
1992
3­
Larson
&
DeMartini
1984
3­
Gnose
1967
3­
Schmitt
&
Holbrook
1984b
4­
Swedberg
1965
4­
Gnose
1967
4­
Hobson
&
Chess
1986
4­
Chu
1989
5­
Fraser
1923
5­
Hobson
&
Chess
1986
5­
Hueckel
&
Stayton
1982
6­
Antrim
1981
6­
Pearcy
&
Myers
1974
6­
Holbrook
et
al.
1985
7­
Bayer
1985
7­
Holbrook,
Carr,
et
al.
1990
7­
Wares
1971
8­
Hubbs
1921
8­
Ebeling
&
Laur
1985
8­
Bennet
&
Wydoski
1977
9­
Wales
1929
9­
Bennet
&
Wydoski
1977
9­
Schmitt
&
Holbrook
1984a
10­
Turner
1952
10­
Hixon
1980
10­
Schmitt
&
Holbrook
1984b
11­
Isaacscon
1965
11­
Schmitt
&
Holbrook
1984b
11­
Holbrook
&
Schmitt
1984
12­
Goldberg
1978
12­
Holbrook
&
Schmitt
1984
12­
Holbrook
&
Schmitt
1988
a
13­
Isaacson
&
Isaacson
1966
13­
Holbrook
&
Schmitt
1988
a
13­
Holbrook
&
Schmitt
1988
b
14­
Bennet
&
Wydoski
1977
14­
Holbrook
&
Schmitt
1988b
14­
Schmitt
&
Holbrook
1985
15­
Wydoski
&
Bennet
1973
15­
Gordon
1965
15­
Gordon
1965
16­
Banerjee
1971
16­
Borton
1982
16­
Borton
1982
17­
Engen
1968
17­
DeMartini
1981
17­
Wingert
et
al.
1979
18­
Triplett
1960
18­
Love
1996
18­
Holbrook
&
Schmitt
1992
19­
Behrens
1977
19­
Allen,
L.
G.
1982
19­
Shrode
et
al.
1983
20­
Holbrook
&
Schmitt
1988
a
20­
Hoffman
1986
20­
Garrison
&
Miller
1982
21­
Holbrook
&
Schmitt
1988
b
21­
DeLeon
1999b
22­
Bane
1970
22­
Love
1996
23­
Wilson
&
Millemann
1969
24­
Suomela
1931
25­
Schmitt
&
Holbrook
1985
26­
Suomela
1931
27­
Shaw
1971
(
benthic
crustaceans;
bivalves;

gastropods)
5,7,20,21,22
kelp
bed
understory;
sandy
bottoms;
estuaries
&
bays
3,5,6,7,8
Page
343
of
1167
5­
2220e
­
Embiotocidae
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
BX
BY
BZ
CA
1­
Holbrook
&
Schmitt
1988b
1­
Ebeling
et
al
1980b
1­
Ebeling,
Larson,
Alevizon,
&
Bray
1980
1­
Hobson
&
Chess
1976
2­
Schmitt
&
Holbrook
1985
2­
Holbrook
et
al.
1985
2­
Bodkin
1986
2­
Feder
et
al
1974
3­
Gordon
1965
3­
Holbrook
&
Schmitt
1984
3­
Hobson
&
Chess
1976
3­
Fritzsche
et
al.
1989
4­
Holbrook
&
Schmitt
1988a
4­
Ebeling
&
Bray
1976
4­
Ellison
et
al.
1979
5­
Holbrook
&
Schmitt
1988b
5­
Feder
et
al
1974
5­
De
Martini
1969
6­
Schmitt
&
Holbrook
1985
6­
Emmet
et
al.
1991
6­
Carlisle
et
al.
1960
7­
Fitch
&
Lavenberg
1975
7­
DeMartini
1969
7­
Odenweller
1971,1975
8­
Miller
1960d
8­
Smith
1967
8­
Bryant
1978
9­
Burge
&
Schultz
1973
9­
Quast
1968a
10­
Fritzsche
et
al
in
Leet
et
al
1992
10­
Wales
1928
11­
Bryant
1978
11­
Fraser
1923
12­
Miller
&
Geibel
1973
12­
Quast
1968d
13­
Alevizon
1975
13­
Antrim
1981
14­
Allen
1985
14­
Carlisle
et
al.
1964
15­
Terry
and
Stephens
1976
15­
Stephens
et
al.
1957
16­
Holbrook,
Carr,
Schmitt,
&
Coyer
1990
16­
Brett
1979
17­
Bayer
1985
17­
Allen,
M.
J.
1982
18­
Allen,
M.
J.
1982
18­
Turner
et
al.
1969
19­
Carlisle
et
al.
1964
19­
Johnson
et
al.
1994
20­
Turner
et
al.
1969
20­
Gnose
1967
21­
Johnson
et
al.
1994
21­
Hobson
&
Chess
1986
22­
Valle
et
al.
1999
22­
Hueckel
&
Stayton
1982
23­
Bodkin
1988
23­
Schmitt
&
Coyer
1983
24­
Larson
&
DeMartini
1984
24­
Holbrook
et
al.
1985
25­
Gnose
1967
25­
Laur
&
Ebeling
1983
26­
Hobson
&
Chess
1986
26­
Stouder
1987
27­
Holbrook,
Schmitt,
&
Ambrose
1990
27­
Wares
1971
bottoms;
artificial
reefs
5,7,9,14,19,20,21,23,24,27,39,41,42,45,53,54,55,56,60,63,64
(
crustaceans,
such
as
barnacles
and
decapods;
gastropods;

bivalves,
such
as
mussels
and
geoduck
clams;
chitons;

ophiuroids
and
Dendraster
;
polychates;
cephalopods;

external
fish
parasites)
2,3,4,5,12,16,18,22,25,26,27,32,45,51,52,56,57,58,62,65
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173
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185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
CB
CC
CD
CE
CF
harbor
seals14,23
pinnipeds20
grazer/
ambusher6
Caspian
terns24
crusher18
1­
Ellison
et
al.
1979
1­
Ebeling
&
Bray
1976
1­
Emmett
et
al.
1991
1­
Haldorson
1980
2­
Boothe
1967
2­
Emmett
et
al.
1991
2­
Odenweller
1971
2­
Bernardi
2000
3­
Allen,
M.
J.
1982
3­
Fritzsche
et
al.
1989
3­
Hubbs
et
al.
1970
3­
Cassano
1998
4­
Hueckel
&
Stayton
1982
4­
Ellison
et
al.
1979
4­
Carlisle
et
al.
1964
5­
Laur
&
Ebeling
1983
5­
Alevizon
1975a
5­
Turner
et
al.
1969
6­
Ebeling,
Larson,
Alevizon,
&
Bray
6­
Alevizon
1975b
6­
Hixon
1980
7­
Schmitt
&
Coyer
1982
7­
Brett
1979
7­
Alevizon
1975a
8­
Schmitt
&
Holbrook
1986
8­
Allen,
M.
J.
1982
8­
Buckley
&
Hueckel
1985
9­
Schmitt
&
Holbrook
1984b
9­
Gnose
1967
9­
Merkel
1957
10­
Holbrook
&
Schmitt
1989
10­
Hobson
&
Chess
1986
10­
Sekiguchi
1987
11­
Alevizon
1975a
11­
Laur
&
Ebeling
1983
11­
Fitch
&
Lavenberg
1971
12­
Cowen
1979
12­
Stouder
1987
12­
Limbaugh
1955
13­
Bray
&
Ebeling
1975
13­
Gordon
1965
13­
Stevens
et
al.
1984
14­
Schmitt
&
Holbrook
1990b
14­
Bray
&
Ebeling
1975
14­
Olesiuk
et
al.
1990
15­
Davis
&
Anderson
1989
15­
Hixon
1981
15­
Fitch
&
Lavenberg
1975
16­
Chu
1989
16­
Holbrook
&
Schmitt
1992
16­
DeLeon
1999b
17­
Holbrook
&
Schmitt
1992
17­
Sikkel
1995
17­
Bayer
1985
18­
Hobson
1994
18­
Helvey
1985
18­
DeLeon
1999b
19­
Hobson
1994
19­
Haaker
et
al.
1984
20­
Love
1996
20­
Morejohn
et
al.
1978
21­
Treacy
1985
22­
Parkin
1998
23­
Torok
1994
24­
Harvey
1982
25­
Miller
1960d
26­
Feder
et
al.
1974
27­
Hawes
1983
harbor
seals;
northern
elephant
seals;

Brandt's
and
double­
crested
cormorants28
picker/
cruncher;
non­
winnower5
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­
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163
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165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
CG
CH
CI
CJ
CK
CL
CM
1­
Haldorson
1973
1­
Beckwitt
1983
2­
Cassano
1998
3­
Waples
&
Rosenblatt
1987
Page
346
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­
Embiotocidae
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163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
CN
CO
CP
CQ
CR
CS
CT
CU
CV
1­
Wares
1971
2­
Bennet
&
Wydoski
1977
3­
Behrens
1977
4­
Behrens
&
Behrens
1973
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347
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­
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165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
CW
CX
CY
CZ
DA
DB
mutual
association
with
Racochilus
toxotes
3
schools
with
E.
lateralis
5
infested
by
parasites
from
the
genus
Lironeca
23
parasitized
by
copepod
family
Clavella15
1­
Schmitt
&
Coyer
1983
1­
Feder
et
al
1974
2­
Holbrook
et
al.
1985
2­
Fritzsche
et
al.
1989
3­
Fritzsche
et
al.
1989
3­
Alevizon
1975b
4­
Hixon
1980
4­
Lloyd
1938
5­
Schmitt
&
Holbrook
1986
5­
Richards
et
al.
1985
6­
Schmitt
&
Coyer
1982
6­
Hobson
1971
7­
Holbrook
&
Schmitt
1989
7­
Limbaugh
1955
8­
Hixon
1981
8­
Wares
1971
9­
Schmitt
&
Holbrook
1990b
9­
Bennet
&
Wydoski
1977
10­
Sikkel
1995a
10­
Wydoski
&
Bennet
1973
11­
Holbrook
&
Schmitt
1995
11­
Arai
1967
12­
Gotshall
1966
13­
Fraser
1919
14­
Bray
&
Ebeling
1975
15­
Dojiri
1981
16­
Hose
et
al.
1983
17­
Limbaugh
19??

18­
Holbrook,
Carr,
Schmitt,
&
Coyer
1990
19­
Ware
1979
20­
Carlisle
1960d
21­
Hobson
1994
22­
Love
&
Moser
1983
23­
Bennett
1993
infested
by
parasites
from
the
families
Argulidae,

Bomolochidae,
Caligidae,
Lernaeopodidae,

Philichthyidae,
Bucephalidae,
Diplangidae,

Hemiuridae,
Monorchiidae,
Opecoelidae,

Zoogonidae,
Cymothoidae,
Heteraxinidae,

Cucullanidae,
Myxidiidae,
Myxobolidae,
ab
indirectly
derives
positive
benefit
from
presence
of
Macrocystis
on
the
understory18
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants16
parasitized
by
Telolecithus
pugetensis,

Neozoogonus
californicus
,
and
others
internal
parasites8,11
removes
ectoparasites
from
fish,
like
the
blacksmith7
infected
by
ectoparasites
from
the
families
Bomolochidae,
Caligidae,
Lernaeide,
and
Lerneopodidae;
also
monogenetic
trematodes6
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168
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170
171
172
173
174
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176
177
178
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180
181
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185
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187
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193
194
195
196
197
198
199
200
201
202
203
DC
DD
DE
DF
DG
DH
Transient9,13,15
winter
migration
to
deeper
water19,25
1­
Smith
1967
1­
Emmett
et
al.
1991
1­
Gnose
1967
1­
Hobson
&
Chess
1976
1­
Hart
1973
2­
Fitch
&
Lavenberg
1971
2­
Miller
&
Geibel
1973
2­
Hixon
1980
2­
Ebeling
&
Bray
1976
2­
Darling
et
al.
1980
3­
Morgan
1961
3­
Antrim
1981
3­
Hixon
1981
3­
Feder
et
al
1974
3­
Hixon
1980
4­
CDF
&
G
1982
4­
Carlisle
et
al.
1964
4­
Emmett
et
al.
1991
4­
Schmitt
&
Holbrook
1986
5­
Turner
1938
5­
De
Martini
1969
5­
Hixon
1981
6­
Terry
and
Stephens
1976
6­
Bryant
1978
6­
Holbrook
&
Schmitt
1992
7­
Wiebe
1968a
7­
Miller
&
Geibel
1973
8­
Johnson
et
al.
1994
8­
Carlisle
et
al.
1964
9­
Gascon
&
Miller
1981
9­
Terry
and
Stephens
1976
10­
Larson
&
DeMartini
1984
10­
Wiebe
1968b
11­
Laur
&
Ebeling
1983
11­
Allen,
M.
J.
1982
12­
Stouder
1987
12­
Turner
et
al.
1969
13­
Anderson
&
Chew
1972
13­
Richards
et
al.
1985
14­
Baltz
1984
14­
Hobson
&
Chess
1986
15­
Wares
1971
15­
Rechnitzer
&
Limbaugh
1952
16­
Bane
1970
16­
Wales
1929
17­
Gordon
1965
17­
Baltz
1984
18­
Conley
1977
18­
Banerjee
1971
19­
Walton
1979
19­
Schmitt
&
Holbrook
1985
18­
Shaw
et
al.
1974
20­
Gordon
1965
19­
Wiebe
1967
21­
Walton
1979
20­
Shrode
et
al.
1983
22­
Shaw
et
al.
1974
21­
Morgan
1961
23­
Bray
&
Ebeling
1975
22­
DeLeon
1999b
24­
Wiebe
1967
23­
Helvey
1985
25­
Limbaugh
1955
24­
Klingbeil
1972
26­
Moyle
1976
25­
Beardsley
1981
27­
Davis
&
Anderson
1989
Page
349
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2220e
­
Embiotocidae
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208
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211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
A
B
C
D
E
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211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
F
G
H
I
JKLM
Page
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Embiotocidae
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205
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208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
N
O
P
Q
R
S
T
U
V
W
X
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Embiotocidae
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208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
Y
Z
AA
AB
AC
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211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
AD
AE
AF
AG
AH
AI
28­
DeLeon
1999b
28­
Garrison
&
Miller
1982
29­
Beardsley
1969
29­
Love
1996
30­
Cassano
1998
31­
Love
1996
Page
354
of
1167
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2220e
­
Embiotocidae
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205
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207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
AJ
AK
AL
AM
AN
AO
AP
28­
Wydoski
&
Bennett
1973
29­
Schmitt
&
Holbrook
1984b
30­
Banerjee
1971
31­
Engen
1968
32­
Triplett
1960
33­
Clark
1930a
34­
Behrens
1977
35­
Banes
1970
36­
Eigenmann
1892
37­
DeMartini
et
al.
1983
38­
Suomela
1931
39­
Gordon
1965
40­
Shaw
1971
41­
Lagios
1965
42­
Shaw
et
al,
1974
43­
Blanco
1938
44­
Wiebe
1967
45­
Fitch
&
Lavenberg
1971
46­
Wang
1981
47­
Borton
1982
48­
Limbaugh
1955
49­
Moyle
1976
50­
Morgan
1961
51­
Breder
&
Rosen
1966
52­
CDF
&
G
1982
53­
Garrison
&
Miller
1982
54­
DeLeon
1999b
55­
Klingbeil
1972
56­
Carlisle
1960a
57­
Miller
1960d
58­
Feder
et
al.
1974
59­
Love
1996
Page
355
of
1167
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2220e
­
Embiotocidae
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205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Page
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Embiotocidae
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211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
BD
BE
BF
BG
BH
28­
Gordon
1965
29­
Blanco
1938
30­
Wiebe
1967
31­
Shaw
et
al.
1974
32­
Fitch
&
Lavenberg
1971
33­
Wang
1981
34­
Breder
&
Rosen
1966
35­
Sivalingam
1953
36­
Fitch
&
Lavenberg
1975
37­
Garrison
&
Miller
1982
38­
DeLeon
1999b
39­
Cassano
1998
40­
Carlisle
1960d
41­
Carlisle
1960a
42­
Miller
1960d
43­
Love
1996
Page
357
of
1167
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2220e
­
Embiotocidae
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205
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208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
BI
BJ
BK
BL
BM
BN
BO
BP
BQ
Page
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216
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219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
BR
BS
BT
BU
BV
BW
28­
Shaw
et
al.
1974
29­
Schmitt
&
Holbrook
1990b
30­
Holbrook
&
Schmitt
1992
31­
Shrode
et
al.
1983
32­
Boothe
1967
33­
Fitch
&
Lavenberg
1975
34­
Garrison
&
Miller
1982
35­
DeLeon
1999b
36­
Cassano
1998
37­
Haldorson
1973
38­
Carlisle
1960d
39­
Carlisle
1960a
Page
359
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221
222
223
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225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
BX
BY
BZ
CA
28­
Baltz
1984
28­
Bennet
&
Wydoski
1977
29­
Hixon
1980
29­
Hixon
1980
30­
Wydoski
&
Bennett
1973
30­
Schmitt
&
Coyer
1982
31­
Schmitt
&
Holbrook
1984b
31­
Schmitt
&
Holbrook
1984a
32­
Triplett
1960
32­
Haldorson
&
Moser
1979
33­
Holbrook
&
Schmitt
1989
33­
Schmitt
&
Holbrook
1984b
34­
Bane
1970
34­
Holbrook
&
Schmitt
1984
35­
Wilson
&
Millemann
1969
35­
Holbrook
&
Schmitt
1988b
36­
Eigenmann
1892
36­
Brooks
et
al.
2000
37­
Cowen
1979
37­
Schmitt
&
Holbrook
1990a
38­
DeMartini
et
al.
1983
38­
Bane
1970
39­
Buckely
&
Hueckel
1985
39­
Alevizon
1975a
40­
Karpov
et
al.
1995
40­
Cowen
1979
41­
Walton
1979
41­
Gordon
1965
42­
Ebeling.
Larson,
&
Alevizon
1980
42­
Bray
&
Ebeling
1975
43­
Bray
&
Ebeling
1975
43­
Hixon
1981
44­
Borton
1982
44­
Fitch
&
Lavenberg
1971
45­
Limbaugh
1955
45­
Limbaugh
1955
46­
Schmitt
&
Holbrook
1990b
46­
Schmitt
&
Holbrook
1990b
47­
Dewees
&
Gotshall
1974
47­
Holbrook
&
Schmitt
1992
48­
Love,
Stephens,
et
al.
1996
48­
Davis
&
Anderson
1989
49­
Stephens
et
al.
1984
49­
Boothe
1967
50­
Boothe
1967
50­
Fitch
&
Lavenberg
1975
51­
Fitch
&
Lavenberg
1975
51­
Garrison
&
Miller
1982
52­
CDF&
G
1982
52­
DeLeon
1999b
53­
Garrison
&
Miller
1982
53­
Sikkel
1995a
54­
DeLeon
1999b
54­
Chu
1989
55­
DeMartini
1981
55­
Holbrook
et
al.
1994
56­
Richards
et
al.
1986
56­
Holbrook
et
al.
1997
57­
Holbrook
&
Schmitt
1995
57­
Klingbeil
1972
58­
Winget
1968
58­
Haldorson
1973
59­
Grossman
1982
59­
Carlisle
1960d
60­
Love
et
al.
1986
60­
Carlisle
1960a
61­
Cassano
1998
61­
Miller
1960d
62­
Carlisle
1960d
62­
Goodwin
&
Pease
1989
63­
Miller
1960c
63­
Birtwell
et
al.
1984
64­
Love
1996
64­
Hobson
1994
65­
O'Brien
&
Valle
2000
(
in
press)
65­
Love
1996
66­
O'Brien
&
Valle
2000
(
in
press)

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222
223
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227
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229
230
231
232
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240
241
242
CB
CC
CD
CE
CF
28­
Love
1996
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242
CG
CH
CI
CJ
CK
CL
CM
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242
CN
CO
CP
CQ
CR
CS
CT
CU
CV
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CW
CX
CY
CZ
DA
DB
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233
234
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236
237
238
239
240
241
242
DC
DD
DE
DF
DG
DH
26­
Cassano
1998
28­
CDF
&
G
1982
29­
Garrison
&
Miller
1982
30­
DeLeon
1999b
31­
Helvey
1985
32­
Richards
et
al.
1986
33­
Cassano
1998
34­
Hobson
1994
35­
Love
1996
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Embiotocidae
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22
23
24
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28
29
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31
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
A
B
C
D
E
Family
Name
Scientific
name
(
Synonym)
Common
Name(
s)
Geographic
range
Observed
depth
range
(
m)

Bothidae
1
Paralichthys
californicus
1,2
California
halibut
Quillayute
River,
northern
Wahington
to
Bahia
1,2
nearshore
to
183
m
Paralichthyidae
2
Monterey
halibut
Magdelena,
southern
Baja
California
1,2,3
southern
halibut,
chicken
halibut
bastard
halibut,
flyswaters,
barndoors,

alabato,
flatties,
door
mat
1,2,3,4,9
Bothidae
1
Citharichthys
sordidus
1,2
pacific
sanddab
Sea
of
Japan
(?)
to
Bering
Sea
(
Alaska
?)
to
Cabo
2
0­
549
m
Paralichthyidae
2
mottled
sanddab,
sole,
soft
flounder,
San
Lucas,
southern
Baja
California
2
megrim,
mottled
saddab,
Catalina
sanddab
2,3,4,5,6
Bothidae
1
Citharichthys
stigmaeus
1,2
speckled
sanddab
Prince
William
Sound,
Alaska
to
Bahia
Magdelena,
195
intertidal­
607
m
Paralichthyidae
2
Catalina
sanddab,
sandab
3,4,6,7
southern
Baja
California
2,3,4,5,7,8
3
majority
of
pop.

occurs
in
<
50
m
Pleuronectidae
1,2
Platichthys
stellatus
1,2
starry
flounder
Japan,
Korea,
Beaufort
Sea,
Canadian
Arctic,
2
nearshore­
375
m
diamond
back,
emery
wheel,
Bering
Sea
and
Aleutian
Is.
to
Los
Angeles
Harbor,

grindstone,
sole,
California
southern
California
2
flounder,
rough
jacket,

great
flounder,
founder
1,2,3,6
Pleuronectidae
1,2
Parophrys
vetulus
English
sole
Bering
sea
and
Aleutian
Island
to
Bahia
San
2
intertidal
to
555
m.

(
Pleuronectes
vetulus
)
2
California
sole,
lemon
sole,
Cristobal,
central
Baja
2,3,4
pointed
nose
sole,
common
RANGE
TAXONOMY
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10
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12
13
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15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
F
G
H
I
J
Total
Length
(
cm)
Max.
reported
Geographic
area
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
1
Both:
W=
0.000939SL3.088
4
TL=
8.15+
1.13
SL
1,2
152.4
cm
TL
Unspecified
location(
NG)
1
1137.43
2
Male:
W=.
000063293
L3.16951
1
TL=
SL
x
1.1878
1
41
cm
TL
NF
NF
2
Female:
W=.
000062851
L3.18012
1
SL=
TL
x
.8421
2
Both:
W=.
000038442
L3.25755
L=
SL,
W=
gm
3
Both:
W=
0.00000257
L
3.26
2
Both:
W=
0.000003934L3.25755
(
SL)

2
Both:
W=.
000019188
L3.44488
(
SL)
1
TL=
SL
x
1.2120
1,2,3
17
cm
TL
NF
NF
4
Both:
W(
dry)=
0.000000531L3.2953
1
SL=
TL
x
.8412
4
Both:
W(
wet)=
0.000002661L
3.2953
5
Females:
W=
0.00003284L2.95
NF
1,2,4,5
91
cm
TL
NF
NF
5
Males:
W=
0.00002959L2.96
13
Both:
W=
0.01917TL2.95410
16Females:
W=
0.0141L2.95
16Males:
W=
0.0102L3.05
6
Both:
W=
0.0000147L2.916
2
FL=
17.766
x
SL(
1.118)
SE:
4.968
1,2,4,6
57
cm
TL
Monterey
Bay
(
TL)
2
30
7
Females:
W=.
00024524
L3.082384
Males:
W=.
00034156
L2.96793
(
W=
ounce2
TL=­
7.324
x
FL(
1.029)
SE:
4.910
Puget
Sound
(
TL)
3
31.22
8,14
Both:
W=.
000034674
SL2.83
2
SL=
2.635
x
TL(.
833)
SE:
3.435
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
K
L
M
N
O
P
Q
R
S
T
U
V
r2
Female
Both
Male
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
1417.42
1217.51
0.1218
0.1194
0.1414
0.1004
0.3801
0.4073
NG
NG
NG
NG1
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
42.84
NG
­
0.3886
­
0.2584
NG
0.296
0.3125
NG
SE
est.
3.35
SE
est.
4.47
NG
interoperculum
2
43.22
NG
0.366494
0.25318
NG
­
0.8272
­
0.6754
NG
SE
est.
3.6450
SE
est.
5.2919
NG
interoperculum
3
k
(
SE)
to
(
SE)

Linf
in
cm
(
SE)
AGE
and
GROWTH
Growth
Parameters
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
W
X
Y
Z
AA
AB
Age
composition
Validation
Verification
Other
refs.
Longevity
(
Length)
information
Sexuality
NG
1
NG
1
1
,15,
20(
juveniles)
1,2
30
yrs.
1
dioecious
1
2,
14,19,22
NF
NF
3,5,22
2,3,9
10
yrs.
2
dioecious
NF
NF
4
4
approx.
36­
42
months
NF
dioecious
2
3.5
yrs.

NF
NF
6,16,21,23
2
24
yrs.
NF
dioecious
17(
juveniles),
18
No
2
betw.
reader
comparison
2
7,8,9,10,11,12
5
17
yrs.
(
52
cm)
3
dioecious
1,2
No
3
compared
structures,
2
22
yrs.(
49.2
cm)

betw.
reader
comparison
3
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AC
AD
AE
AF
AG
AH
AI
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
promiscuous
oviparous1,2
No
females
larger
1,2,3,10,11
No
16
19
cm
(
1)
16
36
cm
(
2)

1
NG
1
NG
2
20
cm(
2­
3)
2
37.5
cm(
4­
6)

3
20­
23
cm(
2)
3
43­
56
cm(
3­
4)

19NG
NG
promiscuous
oviparous
1,3,4,5
No
females
larger
3,4
No
4
NG
4
16.5
cm
TL
1
NG
1
NG
182yrs
2yrs.

19NG
NG
promiscuous
oviparous
1,3
No
males
more
slender
No
5
NG
5
7­
8
cm
TL
and
smaller
head
1
7
cm
1
7
cm
than
females
5
promiscuous
oviparous1,3,9
No
females
are
larger
3,6
No
1
6,17,1822­
28
cm.(
2
yrs.)
6,17,18
24­
36
cm.
(
3
yrs)

722
cm,
(
3
yrs.)
7
30
cm.
(
4
yrs.)

1NG
1
NG
approx.
47.5
cm
(
4yrs.)

promiscuous
oviparous
1,2,3,6
No
females
are
larger
3,712
No
8
2
yrs
8
3
yrs
9
25
cm.
(
3)
9
30
cm.

10
21
cm
10
26
cm
during
breeding
season
females
can
be
distinguished
from
males
by
distension
of
the
body
posterior
to
origin
of
the
anal
fin2
Breeding
Behavior
1st
maturity
Sexual
dimorphism
REPRODUCTION
Total
length
in
cm
(
age
i
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17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AJ
AK
AL
AM
AN
AO
AP
AQ
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.

16
22.7
cm
(
1.3)
16
47.1
cm(
4.3)
Otoliths/
soaked
and
dryed
16
No16
No16
cen.
California
28
1
22.9
cm
(
2)
1
48.3
cm
(
4­
5)
NG1
NG1
NG1
s.
California
4,5
*****

2
NG
2
NG
NG
2
NG
2
NG
2
California
current
region37
3
NG
3
NG
NG
3
NG
3
NG
3
California
1,2,3,14,42
*****

47.1
cm(
3­
4
yrs)
47.1
cm(
3­
4
yrs)
NG
NG
NG
Pacific
coast6
4
NG
4
19.0
cm
TL
(
3)
4
NG
4
NG
4
NG
1,2
Puget
Sound,
WA
6,9,11,12,36
1
NG
1
19
cm
(
2­
3)
1
NG
1
NG
1
NG
n.
California
10
NG
NG
NG
NG
NG
California
9,14,36
19
cm
(
2­
3
yrs)
19
cm
(
2­
3
yrs)
NG
NG
NG
California
to
Baja,
CA
6
5
NG
5
NG
5
scales
5
No
5
No
s.
California
13,2,15,35
1
NG
1
NG
California36
6,17,18NG
6,17,18NG
6
scales
6
No
6
No
1,3
Gulf
of
Alaska
17
7
NG
7
NG
7
NG
7
NG
7
NG
British
Columbia
6,9,11,28,36
1
36.8
cm
(
2­
3
yrs.)
1
40.6
cm(
3­
4
yrs.)
1
NG
1
NG
1
NG
Bering
Sea36
approx.
29
cm
(
3
yrs.)
Puget
sound,
WA
9,11,17,28,41
Washington
6
Montery
Bay,
CA35
CalCOFI
area
8
California
31,6,9,11,14,16,28,36
*****

8
26
cm.
8
29.5
cm.
8
NG
8
NG
8
NG
1,3
British
Columbia11,36
9
46
cm
9
46
cm
9
NG
9
NG
9
NG
British
Columbia
to
Oregon
6
10
26
cm
10
31
cm
10
NG
10
NG
10
NG
Puget
Sound,
WA36,41
50%
maturity
n
yrs)
at
Fertilization
and
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
states
present
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
Multiple
spawner
1
*****
*****
*****
*****
*****

*****
*****

*****
*****
*****
*****
*****
Multiple
spawner
2,3,4,5,6,10
*****
Multiple
spawner3,7
Single
spawners10,11
*****

*****
*****

*****
annual
spawner8
single
spawner12
*****
*****
multiple
spawner13,4
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BD
BE
BF
BG
Fecundity
Mean
Fecundity
Size
specific
eggs/
g
Number
of
eggs
fecundity
eq
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
1
avg.
#
eggs/
spawn:
313,000­
589,000
NF
NF
1,2
approx.
2
days
@
16
C
1
approx.
5.5
million
eggs/
spawning
season
12
1.4(
20
C)
to
3.1
days
(
12
C)

1
small
halibut:
300,000
eggs
every
7
days
1
large
halibut:
1
million
eggs/
day
when
spawning
NF
NF
NF
NF
NF
2
4,300(
85mm)­
30,800
(
148
mm)
NF
NF
2
900,000
­
11,000,000
eggs
per
season
3,7,101,1000,000
eggs
(
56.5
cm.
SL)
NF
3,9
2­
15
days
411,000,000
eggs
7,10900,000­
2,500,000
(
38­
48
cm)

3
1.1
million
eggs
(
average)
4,11150,000
(
30
cm)
to
1,900,000
(
44cm)
1
F=­
1,875,906
+
75,377
L
(
TL
in
cm)
4
3­
12
days
2,4150,000­
2,100,00
eggs
1,5
1
million
(
36­
38cm)­
2
milion.
(
43
cm)
2
F=.
0004558L3.6097
5,6
3­
4
days
6
327,600
(
30
cm)
­
1,566,200
(
43
cm)
3,7
4­
12
days
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22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BH
BI
BJ
BK
BL
Eggs
Size
at
settlement
Size
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
1,2,3
approx.
20­
29
days
1,2,3,19
0.75­
0.94
cm
(
bays
and
along
coast)
1
0.5
to
1
million
fish
(
age
1)
1
0.68­
0.83
mm
planktonic,
nearshore
waters
1,2,3,4
4
5­
6
weeks
4
<
1.7
cm(
bays
and
coast)
2
0.74­
0.84
mm
spawned
in
nearshore
coast
waters24,25
20
4.8­
9.0
mm
SL
(
27­
104
days)
3
0.74­
0.82
mm
3,5
>
271
days
19
20­>
39
mm
(
SL)
NF
1
0.78­
0.84
mm
planktonic,
coastal
waters
5,6
5
19­
38
mm
4
average:
0.1
mm
found
in
polyhaline
waters
7
18
25­
75
mm
5
0.55­
0.77
mm
3,5
>
324
days
6
2.4­>
3.6
cm
SL
NF
1
0.62­
0.66
mm
planktonic,
coastal
waters
6
4
<
3.5
cm
(
bays
and
coast)

5
2.5­
3.8
cm
(
bays
and
coast)

19
2.4­>
3.55
cm
212.6­
5.4
cm
TL
6,7
1.5
months
710­
11
mm
NF
1
0.88­
1.3
mm
pelagic
8
8
25­
30
days
8
10.5
mm
4
0.89­
0.94
mm
eggs
occur
over
water
20­
70
m
deep
7
3
37­
75
days
3,5
0.89­
1.01
mm
6,9,10
2­
2.5
months
9,10
20­
22
mm
(
estuaries)
NF
1
0.80­
1.1
mm
pelagic
4,9,10,22,27
3,11,14,156­
10
weeks
11
<
25
mm
(
estuaries)
6
0.93­
1.05
mm
nearshore
11
4
8­
10
weeks
12
20­
25
mm
(
bays
and
open
coast)
3,4
0.89­
0.93
mm
tolerant
of
a
wide
range
of
eggs
are
pelagic,
spawned
on
the
open
coast23
eggs
spawned
in
shallow,
coastal
marine
areas
near
river
and
slough
mouths23
DISPERSAL
&
RECRUITMENT
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22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BM
BN
BO
BP
BQ
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
NF
1
1.6­
9.4
mm
planktonic,
nearshore
waters
1,2,3,21,25
planktivore
1
NF
2
2.0­
8.0
mm
3
2.0­
9.4
mm
NF
1
<
2.6­
40
mm
4
planktonic,
coastal
waters4,5
planktivore
2
NF
3
2.0­>
39
mm
often
found
offshore2,6,24
found
inshore21,22,25
concentrated
in
midwater24
NF
1
1.3­
38
mm
planktonic,
coastal
waters
2,5,7
planktivore
NF
3
2.0­>
35.5
mm
82.0­>
40
mm
concentrated
in
midwater24
NF
1
1.9­
7.4
mm
planktonic
8
planktivore
3,6
NF
4
1.93­
10.5
mm
found
in
euhaline
to
fresh
water
6
(
copepods,
copepod
nauplii,
barnacle
larvae,
Cladocera)
4
3
1.9­>
6.0
found
in
estuaries
to
37
km
offshore
6
(
copepods,
eggs,
and
nauplii,
as
well
as
barnacle
larvae
and
82.0­
7.0
mm
SL
diatoms)
6
(
Cladocera
eggs)
12
(
copepod
nauplii)
13
NF
1
2.3­
11.5
mm
planktonic
9,
coastal
waters
10,11
planktivore
5,6
visual
feeders
1
5
<
26
mm
found
offshore20
(
copepods
and
other
small
4
>
2.8
mm
planktonic
organisms)
7
found
in
inshore
waters,
occasionally
found
offshore22,24,25
larvae
are
transported
to
inshore
nursery
occurs
throughout
water
column,
most
abundant
in
midwater23,24
Food
habits
Larvae
occur
throughout
the
water
column,
larger
larvae
occur
closer
to
shore16,26
(
Stenosomella
spp
.,
bivalve
veligers,
Euterpina
acutifrons
(
nauplii),
and
unidentified
nauplii)
14
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BR
BS
BT
BU
HABITAT
ASSO
When
/
Frequency
Predators
Fish
Size
Habitat
diurnal
feeder1,3
NF
1
<
2.0­
>
22.0
cm
sandy
substrate
of
bays
and
estuaries
1,2,3,4
2
0.9
cm
to
maturity
9,10
7
mm
to
maturity
sometimes
aggregate
near
rocks
and
sand
dollar
beds
6
bays
used
as
nursery
areas
during
first
year
of
life
25
NF
albacore,
salmon,
8
approx.
42
mm
to
maturity
sandy
to
mud
bottom
7,8,30
mackeral,
and
other
predators
1
shallow
coastal
waters,
bays,
and
estuaries
9
NF
NF
8
approx.
35
mm
to
maturity
nearshore
sandy
substrate
7,30
NF
larger
fish
and
herons
4
3
11.5
mm
to
maturity
nearshore
in
estuaries
and
sandy
intertidal
3
6
10
mm
to
maturity
found
in
mesohaline
to
fresh
water
9
found
in
estuaries
and
lower
reaches
of
major
coastal
rivers
9,2
diurnal
feeder
2
larger
fish
2,3,4
4,5
18­
26
cm
intertidal
zone
10,11
appendicularians
5
3
19.6
mm
to
maturity
benthic
in
bays
and
estuaries
3,5,9,12,13,14,28
shallow
muddy
habitat
15
estuaries
used
as
nursery
and
rearing
areas
before
migrating
to
the
open
coast
to
spawn26,27
juveniles
move
to
areas
of
higher
salinity,
but
remain
in
estuaries
through
their
2nd
year26
costal
embayments
and
estuaries,
also
in
shallow
open
costal
waters5
remain
in
shallow
water
embayments
until
15.0­
20.0
cm
SL26
juveniles
remain
in
bays
approx.
2
yrs,
then
emigrate
to
coast6
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BV
BW
BX
BY
OCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Foraging
guild
Foraging
method
When
/
Frequency
Predators
carnivores
1
ambusher
1
diurnal
feeder
1
shore
birds,
water
fowl,
and
fishes
in
(
small
fish,
mysids,
crustaceans)
2,3
bays
1
(
bottom
dwelling
invertebrates)
1,4
carnivores
5
Sit
and
wait
or
hunt
and
peck
2
active
both
day
and
night
2
larger
fish,
shorebirds,

(
crustaceans,
intervebrates)
6,7
and
marine
mammals
2
benthic
carnivores
9
ambusher
3
diurnal
feeder
3
larger
demersal
fish
particularly
(
copepods,
amphipods,
cumaceans,
mysids)
9
California
halibut
3
carnivore
5
ambusher
4
diurnal
feeder
4
seabirds
and
marine
mammals
12
(
copepods,
amphipods,
and
annelid
worms)
5
pinnepeds,
larger
fishes,
sharks,
marine
(
primarily
copepods)
10,11
mammals,
herons,
and
comorants
9
(
invertebrates)
12
(
nemerteans
and
priapulids)
13
(
polychaetes,
mysids,
tanaids)
14
benthic
carnivore
5,15
pursuit
and
capture
5
diurnal
feeder
5
lincod,
greenlings,
croakers,
rockfish,

(
Magelona
palps,
juvenile
bivalves,
bivalve
siphons,
copepods)
16
juveniles
>
6.5
cm
dig
for
prey
5
and
sharks
4
(
small
polychaetes,
molluscs,
and
crustaceans)
17
lg.
piscivorous
fishes,
big
skate
Food
habits
Juveniles
(
northern
anchovy
(
large
juveniles);
mysids,
and
larval
fish(
small
juveniles))
24
(
gammarid
amphipods,
mysids,
teleost
fishes,
and
harpacticoid
copepods)
23
(
calanoid
copepods,
glyceriform
polychaetes,
gammarid
amphipods)
8
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24
25
26
27
28
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33
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35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BZ
CA
CB
CC
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
carnivore
ambush
predator
1,2,3
diurnal
feeder
1,2
(
prim.
schooling
fish)
1,2,3
locates
prey
by
sight
2,3,4
active
both
day
and
night
3,4
(
Pacific
sardine,
squid,
white
croaker,
northern
anchovy)
4
(
fish,
squid
,
octupi)
5
piscivore
(
engraulids,
atherinids,
and
small
flatfish)
42
(
anchovies,
other
fishes,
and
squid)
44
sandy
to
mud
bottom
3,4,5
opportunistic
carnivores
5
Sit
and
wait
or
hunt
and
peck
5
active
both
day
and
night
3,4
estuaries
and
coastal
waters
to
as
deep
at
306
m
6
(
crustaceans,
intervebrates)
6,7
pelagic
feeder,
with
some
benthic
feeding
2,6,7,8
shallow
water,
sandy­
bottom
association27
(
anchovies)
8
locates
prey
by
sight
2
(
polychaetes,
amphipods,
bivalves,
tanaids,
mysids)
10
(
euphausiids,
squid)
2
(
krill,
shrimp,
squid,
small
fish,
worms)
5
nearshore
sandy
substrate
4,7,29
benthic
carnivores
off
the
bottom
pursuit
2
diurnal
feeder
3,5
spring­
summer
seasonal
of
rocky
intertidal28
(
amphipods,
mysids)
2,11
ambusher,
digger
9
(
decapods,
molluscs,
fishes)
12
locates
prey
by
sight
2
(
polychaetes,
fish,
crabs,
and
mysids)
13,14
(
mysids,
gammarid
amphipods,
reptantian
decapods,
and
echiurids)
2
(
mysid
shrimp,
gammarid
amphipods,
shrimps,
worms,

squid,
and
fish)
5
(
tiny
crustaceans,
polychaete
worms,
and
fish)
41,45
gravel,
sand,
and
mud
8,24
benthic
carnivore
(
bivalves,
Echiroids)
11
benthic
predator,
stirs
up
substrate
10
diurnal
feeders
6,7,8
euryhaline,
found
from
seawater
to
fresh
water
6,24,
(
clams,
crabs,
sand
dollar,
and
brittle
star)
13
Relies
heavily
on
olfactory
cues
11
don't
feed
during
spawning
most
adults
found
in
less
than
150
m
6
(
amphipods,
molluscs,
annelids,
crustaceans,
insects)
16
or
cold
water
periods
13
found
in
estuaries
or
their
freshwater
sources
6
(
bivalves,
polychaetes,
crabs,
callianssids)
17
(
annelids,
crabs,
shrimp,
clams)
18
(
clams,
brittle
stars,
fishes,
crabs)
5
(
crabs,
shrimp,
worms,
clams,
calm
siphons,

other
small
molluscs,
small
fish,
brittlestars)
19
(
adult
and
juvenile
geoduck
clams)
46
soft
sand
or
mud,
located
offshore
3,
9,10,12
benthic
carnivore
(
prim.
polychaetes)
2,8,20
pursuit
and
capture
predator
12
diurnal
feeders
2,3,9,10
sex
segregation
assoc.
with
sediment
12
(
sm.
whole
clams
and
amphipods)
15
capable
of
digging
into
sediment
12
shallow
muddy
habitat
12
(
sand
dollars,
gasteropods
and
bivalves)
13
searcher
2
feeds
opportunistically
on
Capitella
at
night20
Adults
Food
habits
occurs
nearshore,
with
larger
individuals
ocurring
deeper2
benthic,
sandy
substrate
often
aggregate
near
structures1
(
northern
anchovy,
mysids,
caridean
shrimp,
and
other
juvenile
and
adult
fish)
40
found
in
sand­
bottom
kelp
holdfast
and
rockybottom
kelp­
bed
habitat29
live
buried
in
the
sand,
near
rocks
or
seaweeds
at
depths
that
average
10
m29
found
in
sand­
bottom
kelp
holdfast
and
rockybottom
kelp­
bed
habitat29
found
in
sand­
bottom
kelp
holdfast
and
rockybottom
kelp­
bed
habitat29
(
shrimp,
crabs,
polychaetes,
squid,
octopus,
and
other
fishes)
44
(
amphipods,
isopods,
insects,
oligochaetes,
and
calanoid
copepods)
47
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24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CD
CE
CF
CG
Haplotype
diversity
Predators
Substocks
(
Geographic
Area)

Pacific
angel
shark,
Pacific
electric
eel,
California
sea
lion,
and
Mission
Bay
NF
NG
bottlenose
dolphin
1
Marina
del
Ray
1
electric
rays,
angel
sharks,
and
other
larger
predators
2,3
Northern
fur
seals,
California
sea
lions,
Pacific
white­
sided
NF
NF
NF
dolphins,
Dall's
porpoises,
common
murres,
pigeon,
guillemots,

Brandt's
and
double­
crested
cormorants
2
harbor
seals14
larger
demersal
fish,
particularly
California
halibut
4
NF
NF
NF
various
piscivores,
including
habor
seals,
Brandt's
cormorants,
pigeon
guillemots
2
cormorants,
seals,
sea
lions,
other
fish
and
crabs
11
harbor
seals14,15
Caspian
terns16
harbor
seals,
cormorants,
and
herons
2
NF
NF
NF
pinnepeds,
larger
fishes,
sharks,
and
marine
mammals
6
harbor
seals10,12,14
marine
mammals,
sharks,
and
larger
fishes
3,5
NF
NF
NF
arrowtooth
flounder,
California
sea
lions,
and
pelagic,

doublebreasted
and
Brandt's
cormorants2
Nucleotide
diversity
(#)

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23
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31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CH
CI
CJ
CK
CL
CM
Genetic
variance
Mean
number
of
%
Polymorphic
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
1
1.29
allele/
locus
23.7
%
of
loci
0.046
exp.
(.
019)
.046
obs.
(.
019)
allozymes
NG
1.68
allele/
locus
26.3
%
of
loci
0.058
exp.
(.
027)
.058
obs.
(.
027)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Mean
heterozygosity
GENETICS
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23
24
25
26
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CN
CO
CP
CQ
CR
CS
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Fishery
/
Length
(
age)
Fishery
/
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
1
8­
12
mm
SL
(
20­
29
days)
NG
small
seines,
otter
trawl,
beam
trawl
otoliths/
whole
NG
NG
2
NG
male:
avg.
57.6
cm
(
6)
gill
nets
and
trawl
nets
otoliths/
whole
No
No
female:
avg.
61.8
cm
(
6)
and
cut
and
burn
NF
NF
NF
NF
NF
NF
NF
NF
Nf
NF
NF
NF
5
20­
56
cm
NG
otter
trawl
NG
NG
NG
6
26
mm
NG
trawl
NG
NG
NG
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29
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31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CT
CU
CV
CW
Mortality
Estimates
Method
of
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
1
0.3
to
0.15
NG
NG
virtual
population
analysis
9
mean:
0.0124
per
day
SD=
0.001
NG
9
instantaneous
mortality:
0.044
(
youngest
juveniles),
NG
(
for
fish
31­
70
mm
or
70­
115
days)
mean:
0.0124
(
SD=
0.001)
(
juveniles<
70
and
older)

bay
model:
0.043
to
0.037
(
juveniles)

total
mortality
model:
0.011
to
0.014
(
juveniles)

13
NG
NG
Females:
approx.
0.30­
0.48
linear
regression
analysis
Males:
approx.
0.21
NF
NF
NF
NF
2
99.8­
99.9
%
(
egg
and
larval
period)
NG
NG
diff.
betw.
total
eggs
spawned
per
season
and
age
0
fish
present
in
study
area
7
99.8­
99.9%
(
egg
and
larval
period)
NG
NG
NG
3
0.294
NG
NG
NG
4
NG
0.775
NG
NG
5
Carr
Inlet
pop:
NG
NG
Beverton­
Holt
equation
MORTALITY
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CX
CY
CZ
DA
Rate
of
Exploitation
Age
range
used
Rate
of
Exploitation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
NG
NG
NG
California
lizardfish,
bigmouth
sole,
lingcod,

NG
NG
NG
speckled
sanddab
1
NG
NG
NG
NF
NF
NF
Citharichthys
stigmaeus
2
egg
and
larval
stages
NG
NG
Citharichthys
sordidus,
C.
xanthostigma,

Symphurus
atricauda,
Parophrys
vetulus
2
California
halibut
1
NG
NG
NG
competes
with
other
soft­
bottom
benthic
fishes
of
estuaries
and
shallow
nearshore
bays
3
ages
4­
13
NG
NG
slim
sculpin,
black
belly
eelpout,
Pacific
tomcod,

NG
NG
NG
ratfish,
Dover
sole,
and
white
croaker
3
0+
english
sole
NG
NG
speckled
sanddab
2
Page
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23
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25
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
DB
DC
Interactions
/
associations
with
other
organisms
Tagging
studies
Larvae:
associated
w/
larve
of
diamond
turbot
and
spotted
turbot
1
55%
of
tag
recoveries
showed
no
net
movement,
50%
of
Juveniles:
assoc.
w/
cheekspot
goby
and
giant
kelpfish
in
bays,
tagged
fish
showed
movement
>
1
km
w/
speckled
sanddab
and
spotted
turbot
along
coast.
2
13.4
km
(
avg.)

Adults:
schools
of
surf
perch
and
anchovy
1
64
most
recovered
within
5
miles
of
tag
site,
one
recovered
1
endoparasites:
trematodes
(
Tubulovesicula
linbergi
)(
Stephanostomum
casum
),
3
miles
away
after
39
days
cestodes(
Echeneibothrum
spp
.),
and
nematodes
(
Spirocamallanus
pereirai
)

1
ectoparasites:
copepods(
Lepeophtheirus
bufidis
)(
Taenicanthodes
haakeri
)

(
Acanthochondria
solea
)(
Holobomalochus
prolixus
)
and
isopods
(
Lironeca
vulgaris
)

parasites:
16
families,
21
spp.
(
see
reference)
14
parasite:
isopod
(
Lironeca
vulgaris
)
15
Dover
sole,
plainfin
midshipman,
pink
seaperch,
and
shortspine
combfish
2
NF
parasite:
isopod
(
Lironeca
vulgaris
)
13,15
parasites:
6
families,
8
spp.
(
see
reference)
14
California
tonguefish,
hornyhead
tubot,
and
english
sole
2
NF
parasites:
4
families,
5
spp.
(
see
reference)
14
parasite:
isopod
(
Lironeca
vulgaris
)
15
hybridizeds
with
a
related
flounder,
Kareius
bicoloratus
in
Japan
3
4,11
have
show
movements
up
to
200
km,
but
not
typical
hybridizes
with
english
sole
4
only
15
of
238
recaptures
within
Yaquina
Bay
moved
more
than
parasites:
isopod(
Livoneca
vulgara
),
cestode(
Lacistorhynchus
tenui
),

and
(
Corynesoma
sp.)

parasites:
copepods
(
Lepeophtheirus
hospitalis
),
(
Chondracanthus
rectangularis
)

(
Lepeophtheirus
nanaimoensis
),
and
(
Nectobrachia
indivisa
)
11
parasites:
trematode
(
Parahemiurus
platichthyi
)

parasites:
25
families,
46
spp.
(
see
reference)
14
parasite:
isopod
(
Lironeca
vulgaris
)
15
hybridizes
with
starry
flounder
5,6
majority
of
recaptures
in
vicinity
of
the
tagging
5,6,15
parasites:
nematode(
Philometra
americana
)
8
tagging
results
suppport
the
existance
of
separate
stocks
4,6,7,1
parasites:
acanthocephalan
(
Echinorhynchus
lagenifromis
),
nematode(
Philometra
extensive
migrations
more
characteristic
of
females
7
parasite:
digenea
(
Tubulovesicula
lindbergi
),
nematoda
(
Anisaka
sp.
larval)
16
parasites:
digenea
(
2
sp.),
monogenea
(
1
sp.),
cestoda
(
1
sp.),
nematoda(
4
sp.),

acanthocephala
(
1
sp.),
and
copepoda(
1
sp.)
16
BEHAVIOR
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
DD
DE
DF
DG
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
Resident
1,2
Yes
1
No
NF
Seasonal
onshore­
offshore
movements
related
to
spawning
3,4
Adults
in
California
migrate
inshore
from
February
to
July
for
spawning
5
not
highly
migratory,
movement
influenced
by
prey
6,7
NF
No
NF
postflexion
larvae
vertically
migrate
into
upper
water
column
at
night
18
not
highly
migratory,
movement
influenced
by
prey
6,7
NF
No
NF
postflexion
larvae
vertically
migrate
into
upper
water
column
at
night
18
Resident
8,9,19
No
NF
adults
move
inshore
in
late
winter­
early
spring
to
spawn
and
offshore
and
deeper
in
the
summer
and
fall,
these
movements
are
generally
less
than
5
km
16
Resident
9,20
Yes
2
No
Possibly
1
inshore­
offshore
seasonal
movement
2,10,11
tagged
flounder
transported
within
the
bay
tended
to
return
to
the
point
of
original
capture4
inshore­
offshore
seasonal
movement
associated
with
Movement
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67
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71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
A
B
C
D
E
sole,
sharpnose
sole,
igirisu
garei
1,2,3,4,6
Pleuronectidae
1,2
Eopsetta
jordani
1,2
petrale
sole
Kupreanof
Pt.,
western
Gulf
of
Alaska
to
Islas
Los
2
0­
550m.

brill,
California
sole,
cape
sole
Coronado,
northern
Baja
California
2
roundnose
sole,
Jordan's
flounder,
English
sole,
soglia,
petorau,

nameta,
tsubame
garei
2,3,4,6,8
Pleuronectidae
1,2
Hypsopsetta
guttulata
1,2
diamond
turbot
Cape
Mendocino,
northern
California
to
2
1.5­
82
m.

diamond
flounder,
turbot,
Bahia
Magdelena,
southern
Baja;
Gulf
of
halibut,
sole
1,2,3
California
2,3,4,6
Pleuronectidae
1,2
Lepidopsetta
bilineata
rock
sole
Southern
Japan
to
Korea
and
the
Sea
of
Okhotsk
2
0­
366
m.

Plueronectes
billineata
1
broadfin
sole,
roughscale
north
to
St.
Lawrence
I..
In
the
Bering
Sea
along
(
scientific
name
in
question)
sole,
two­
lined
flounder,
Aleutian
Is.
west
to
Stalemate
Bank
to
Tanner
slime
sole,
smear
dab,
Bank,
southern
California
2
tounge
sole,
rubber
sole,

short­
finned
sole,
gravel
sole,

roughback
sole,
flounder,
sole,

doublelined
founder
1,2,3,6
Pleuronectidae
1,2
Psettichthys
melanostictus
1,2
sand
sole
Southeastern
Bering
Sea
and
Aleutian
Is.
To
2
intertidal
to
325
m.

fringe
sole,
sand
flounder,
spotted
Redondo
Beach,
southern
California
2,3,4
flounder,
sand
dab,
karui­
rui
1,2,4,6
Page
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
F
G
H
I
J
9
Male:
W=.
1031
L2.970
Female:
W=.
1407
L3.077
1TL=­
7.324
x
FL(
1.029);
SE:
4.910
9
Male:
W=.
00001321
L3.033
Female:
W=.
00001168
L
3.012
1FL=
16.866
x
TL(
0.943);
SE:
4.701
13
Both:
W=
0.01429TL3.67544
10
Male:
W=
0.00272
TL3.135
NF
1,2,4,5
70
cm
TL
California
(
TL)
4
52
10
Female:
W=
0.000127
TL3.352
(
weight=
ounces)
Washington(
NG)
5
48.9
13
Both:
W=
0.00606
TL3.21749
British
Columbia
(
NG)
6
49
11
Both:
W=
2.213
x
10­
5
SL3.044
(
mm)
3
TL=
12.22
+
SL
(
1.100)
1,2,7
46
cm
TL
Anaheim
Bay,
CA(
NG)
7
NG
223
Both:
W=
.0078
L3.136
223
NF
1,2,4,5
60
cm
TL
British
Columbia
(
NG)
8
40.2
12
Female:
W=
0.000001145
L3.410(
mm)

12
Male:
W=
0.000008792L3.0626
(
mm)

13
Both:
W=
0.00552TL3.23131
NF
NF
1,2,5,4
63
cm
TL
NF
NF
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67
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73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
K
L
M
N
O
P
Q
R
S
T
U
V
63
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
tag
recovery
4
NG
NG
0.14
NG
NG
­
6.24
NG
NG
NG
NG
NG
estimated
from
Walford
plot
5
58.6
NG
0.16
0.167
NG
­
2.6
­
0.27
NG
NG
NG
NG
otolith
measurement
6
NG
19.83
NG
NG
0.1
NG
NG
0.31
NG
NG
NG
scales
and
otoliths
7
51.6
NG
0.26
0.146
NG
­
1.56
­
2.5
NG
NG
NG
NG
otolith/
measurement
and
length
frequency
analysis
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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67
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
W
X
Y
Z
AA
AB
No
4
No
4
22
2,9
25
yrs.
4
dioecious
No
5
No
5
7
females:
19
yrs(
57
cm)

No
6
No
6
7
males:
16(
55
cm)

No
7
compared
structures
and
2
8
yrs.
NF
dioecious
1
analysis
of
length
frequency
data
7
8
2
yrs.
(
18.0
cm)

No
8
No
8
13,22
2,6
22
yrs.(
52
cm)
5,
6
dioecious
NF
NF
5
NF
2
dioecious
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70
71
72
73
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79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
AC
AD
AE
AF
AG
AH
AI
11,1821
cm
(
2)
11,1826
cm(
3)

12
20­
25
cm(
2)
12
25­
36
cm
(
3)

202
yrs.
3­
4
yrs
promiscuous
oviparous
3,5,7
No
females
are
larger
8
No
9
32
cm.
(
4)
9
35
cm.
(
5
yrs.)

7
NG
7
32
cm
(
5)

13
NG
13
NG
12
29.7
cm
(
3)
12
31.8
cm
(
4)

10
29
cm.
10
31
cm.

5
32
cm
17
35
cm
1
NG
1
NG
19NG
NG
promiscuous
oviparous
1,2,3
No
NF
No
2
NG
2,1718.0
cm
(
2­
3)

1
NG
1
16.5
cm
(
2­
3)

promiscuous
oviparous
1,3,5
No
females
larger
3,9
No
9
28
cm
(
4)
9
31
cm
(
4)

14,18NG
14
NG
21
cm(
4)
(
AC)
25
cm
(
5)

182
yrs.
3­
4
yrs.

promiscuous
oviparous
1,3,4,5
No
NF
No
1
2
yrs
1
2­
3
yrs.

15
23
cm
(
2)
15
28
cm
(
3)

Page
390
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1167
5­
2220e
­
Flatfish
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65
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
AJ
AK
AL
AM
AN
AO
AP
AQ
11,18NG
11,18NG
11,18NG
11,18NG
11,18NG
Oregon
18,19,36
*****

12
NG
12
NG
12
NG
12
NG
12
NG
Monterey
Bay
7,28,36
NG
NG
NG
NG
NG
cen.
California
28
CalCOFI
area
8
California
32,35,36,43
*****

California
to
Baja,
California
6
*****

9
NG
9
NG
9
NG
9
NG
9
NG
1,3
British
Columbia
6,9,11,21
7
NG
7
NG
7
NG
7
NG
7
NG
Washington
9,17
13
36­
37
cm
(
7)
13
40­
44
cm.
(
8)
13
NG
13
NG
13
NG
Oregon
6,9,17,19
*****

12
40.6
cm(
7)
12
NG
12
NG
12
NG
12
NG
Washington
to
Oregon
29
*****

10
36
cm.
10
40
cm.
10
NG
10
NG
10
NG
British
Columbia
to
Oregon36
17
38
cm.(
7)
17
44
cm.
(
8)
17
Otolith/
measurements
17
No
17
No
CalCOFI
area
8
1
38
cm.(
7)
1
43
cm
(
8)
1
NG
1
NG
1
NG
California
17,20,40
*****

35.5
cm
35.5
cm
NG
NG
NG
throughout
range44
2
NG
2
NG
2
NG
2
NG
2
NG
San
Francisco
Bay,
CA
6
1
NG
1
NG
1
NG
1
NG
1
NG
cen.
California
28
Richardson
Bay,
CA
23,28
Anaheim
Bay,
CA
6,22,28
*****

San
Diego,
CA
24
9
NG
9
NG
NG
9
NG
9
NG
9
1,3,4
Bering
Sea6,9,36
14
32­
33
cm
(
4­
5)
14
<
30
cm
(
approx.
4)
NG14
NG14
NG14
British
Columbia26,36
NG
NG
NG
NF
NG
Puget
Sound,
WA
9,11,41
NG
NG
NG
NG
NG
CalCOFI
area
8
southern
California
9
Okhotsk
sea
coast
25
off
Kamchatka
6
North
America
6
1
NG
1
NG
NG1
NG1
NG1
3
northern
British
Columbia
9
15
NG
15
NG
whole
otolith/
dryed
15
No
15
No
15
Vancouver
Island,
B.
C.
9,27,30
Puget
Sound,
WA
9,11,12,30,34,36,38,41
CalCOFI
area
8
Pacific
coast
6
Page
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1167
5­
2220e
­
Flatfish
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
*****
*****
*****
*****
*****
*****
*****

*****

*****
*****

*****
*****
*****
multiple
spawner13
*****
*****

*****
NF
*****
*****

*****
NF
*****

*****
*****
NF
*****

*****
Page
392
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1167
5­
2220e
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Flatfish
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54
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BD
BE
BF
BG
10150,000
(
30
cm,
3
yrs)
to
300,000
(
43.7
cm)

121
million(
33
cm)­
2
million
(
43
cm)
11
approx.
4
days
173.75
days
@
55
F
NF
3,4,7,10400,000(
42
cm)­
1,200,000(
57cm)
NF
8,9
6.2­
13.5
days
3
8
days
6,13
8­
9
day
NF
NF
NF
NF
NF
3,4,7,8,11400,000
(
35
cm)
­
1,300,000
(
46
cm)
3
F=(
4,378)
10­
4l3.720
3,9
6­
25
days
9
80,000(
35
cm)­
920,000(
44
cm)
3
F=(
4,891)
10­
4l3.7355
6
9­
25
days
4F=
4.891
x
10­
4L3.720
(
L=
length
of
fish
in
cm)
1424
days
4F=­
178.7
+
0.695
W
(
W=
weight
of
fish,
g.)

NF
7
900,000
(
28
cm)­
1,400,000
(
37
cm)
NF
6,10,14
5­
7
days
9
3.5­
7
days
156­
7
days
10255,000
(
35­
36
cm),
305,000
(
37­
38
cm),
415,000
(
39­
40
cm),
435,000
(
41­
42
cm),
and
615,000
(
43­
44
cm)
2,8
3.5
days
@
12C
to
11.8
days
@
4C
Page
393
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1167
5­
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Flatfish
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BH
BI
BJ
BK
BL
12
4
months
721­
22
mm
2
average:
0.98
mm
salinities,
limited
in
distribution
82
months
by
temperature
12
13
6
months
(
including
egg
stage)
13
22
mm(
inner
continental
shelf)
NF
1,4
1.2­
1.3
mm
planktonic13,28
deep
water
14
3
1.21­
1.25
mm
2,3,4,145­
6
weeks
14,15
approx.
1.1
cm
SL
NF
1,2,3,12
0.78­
0.89
mm
pelagic,
in
estuaries
and
coastal
waters4,1
2
0.70­
0.88
cm
2
average:
0.84
mm
found
in
euhaline­
polyhaline
conditions4
16
0.44­<
1.12
cm
8
average:
0.80
mm
pawning
takes
place
within
2
km
of
shore2
4
<
1.7
cm
(
coast)
8
5
1.27
cm
(
shallow
areas)

210.44­<
0.88
cm
SL(
bays,
sloughs,
and
estuaries)

14
25­
50
days
716­
20
mm
NF
1,5,6,7
0.87­
1.0
demersal,
adhesive
eggs
8,11,16,17,18
5
25.4
mm
4,7
average:
0.92
mm
found
in
polyhaline
to
euryhaline
3
1.02­
1.09
mm
waters
7
860­
68
days
8
23­
27
mm
NF
1
0.83­
1.0
mm.
4
pelagic
7,8,16,19
7
25­
28
mm
3,4,6
average:
1.0
mm
at
sea
surface
20
5
25.4
mm
(
estuaries
and
open
coast)
Page
394
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1167
5­
2220e
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Flatfish
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63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BM
BN
BO
BP
BQ
2
2.0­
26
mm
(
appendicularians:
Oikopleura
spp.)
8,9
3
2.3­
20
mm
(
larvaceans,
eggs,
nauplii,
and
tintinnids)
10
planktivorous
invertebrates
1
2.8­
17.6
mm
planktonic
12
planktivore
6,10
NF
and
pelagic
fish
1
4,6
3.0­<
21
mm
found
up
to
150
km.
offshore
17
(
copepods,
their
eggs
and
nauplii)
6
(
plankton)
10
NF
1
2.2­
8.8
mm
pelagic,
in
estuaries
and
coastal
waters3,13
planktivorous
5,11
NF
2
average
1.6­
9.6
mm
SL
found
in
euhaline­
polyhaline
conditions3
(
zooplankton
and
phytoplankton)
7
23
3
1.7­>
8.8
mm
NF
4
5­<
20
mm
pelagic
12
planktivore
6
NF
3
3.4­>
17.7
mm
found
in
upper
30
m
of
water
column
6
1
3.1­>
17.7
mm
found
in
polyhaline
to
euryhaline
waters
6
3
3.4­<
20
mm
small
fishes
and
sea
birds
1
1
2.1­>
22.6
mm
pelagic
6,12,14
planktivore
NF
4
2.8­<
23
mm
planktonic
11
(
copepods,
their
eggs
and
nauplii)
6
3
<
3.0­>
22.6
mm
found
at
sea
surface
15,6
pelagic
in
coastal
waters
and
bays
18
found
in
euhaline
waters
18
abundant
in
neustonic
and
midwater
layers
nearshore24
areas
such
as
the
intertidal
zone,
bays
or
estuaries22
Page
395
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1167
5­
2220e
­
Flatfish
53
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BR
BS
BT
BU
found
in
polyhaline
and
euryhaline
waters
5,9
common
in
shallow
waters
along
the
coast
as
well
as
in
estuaries
24
NF
planktivorous
invertebrates
6
2.2
cm
to
maturity
sandy
or
sand­
mud
bottom
16,23
and
pelagic
fish
4
NF
NF
3
<
13.2
to
maturity
benthic
in
bays,
estuaries
and
sloughs
and
nearshore
coastal
waters
5,16,17
sand
and
mud
bottom
5,7,18
found
in
euhaline­
mesohaline
conditions
5
NF
larger
fish
and
sea
birds
4
3
19.1
mm
to
maturity
sand
or
mixed
sand
substrate
in
waters
<
50
m
15,19
6
17
mm
to
maturity
sand
and
muddy
sand
substrate
20
sand,
gravel
substrate,
rock
slopes,
or
soft
bottom
9
shallow
water
bays
over
the
continental
shelf
9
found
in
polyhaline
to
euryhaline
waters
9
NF
small
fishes
and
sea
birds
4
3
20.9
mm
to
maturity
sand
or
mud
bottom
21
6
23
mm
to
maturity
found
in
estuaries
along
the
west
coast
9
found
in
euhaline
waters
9
estuaries
are
used
and
nursery
grounds,
before
immigrating
to
the
open
coast(
6­
18
months)
26,27
Page
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1167
5­
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Flatfish
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BV
BW
BX
BY
and
sand
sole
5,6
piscivorous
birds,
fishes,
mammals
7,8
sharks
9
carnivore
NF
NF
adult
petrale
sole,
as
well
as
larger
fish
(
annelid
worms,
clams,
brittle
star)
9
(
mysids,
sculpin,
and
other
juvenile
flatfish)
5
benthic
carnivore
1,18
foraging
on
or
in
the
diurnal
feeder
6
Pacific
electric
ray,
Pacific
angel
shark,

substrate
6
and
other
piscivorous
fishes
10
carnivore
5
NF
diurnal
feeders
7
adult
rock
sole
and
other
larger
fish
9
(
copepods,
mysids,
gammarid
amphipods,
polychaetes)
20
(
small
polychaetes,
molluscs,
and
crustaceans)
17
(
polychaetes,
amphipods,
bivalves,
tanaids,
mysids)
14
(
cumaceans,
carideans,
and
gammarid
amphipods)
5
carnivore
NF
NF
larger
fishes
9
(
polychaetes,
clams
and
clam
siphons,
gastropods,
ghost
shrimp,

amphipods,
crustaceans
and
sm.
fish)
19
(
copepods,
their
eggs
and
nauplii,
small
crustaceans,
such
as
mysids
and
crangons,
worms,
and
mollusks)
5
(
harpacticoid
copepods,
gammarid
amphipods,
cumaceans,
mysids,

polychaetes,
small
bivalves,
clam
siphons,
and
other
benthic
invertebrates)
1
(
newly
settled
(<
50
mm):
polychaete
pieces,
amphipods,
bivalve
siphons,
and
mysids.
1
yr.
olds(
50­
100
mm):
cumaceans
and
14
Page
397
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1167
5­
2220e
­
Flatfish
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
BZ
CA
CB
CC
nearshore
coastal
waters
13
(
small
polychaetes,
molluscs,
and
crustaceans)
21
digs
into
substrate
using
pointed
snout
and
(
annelids,
crabs,
shrimp,
clams)
18
scoops
out
prey
13
(
amphipods,
cumaceans,
polychaetes,
mysids)
10
prey
located
by
sight,
smell,
and
perhaps
lateral
line
2
(
polychaetes,
brittle
stars,
gammerid
amphipods)
2
(
worms,
amphipods,
clams,
brittle
stars,
small
fish)
5
(
polychetes,
amphipods,
molluscs,
ophiouroids,

and
crustaceans)
3
(
Capitella
spp.)
43
(
adult
and
juvenile
geoduck
clams)
46
(
worms,
molluscs,
small
starfish,
small
crabs,
and
small
fish)
48
sandy
and
sand­
mud
bottom
14
carnivore
(
fishes
and
decapod
crustaceans)
8,22
NF
NF
(
fish,
pelagic
and
semipelagic
invertebrates)
12
(
fish,
large
crustaceans,
octupi,
squid)
5
(
bottom
fish,
invertebrates)
19
(
euphausiids,
shrimp,
anchovies,
and
other
fishes)
44
benthic
in
bays,
estuaries
and
sloughs
and
benthic
carnivore
3
foraging
on
or
in
the
substrate
3,14
diurnal
feeders
2,11,12
nearshore
coastal
waters
13,14,15
(
prim.
Molluscs,
poylchaetes,
and
crustaceans)
23
sand
and
mud
bottom
4,13,16,29
(
clam
siphons,
worms,
fishes,
small
crustaceans)
5
found
in
euhaline­
mesohaline
conditions
13
demersal
on
soft
to
hard
substrate
3,17
benthic
carnivore
(
polychaetes,
sand
lance)
24,25
bottom
feeder
16
diurnal
feeders
10,13,16
pebbly
or
semi­
rocky
bottoms
5
(
mollusks)
26(
ophiuroids)
8
rock
sole
doesn't
feed
in
the
sand,
gravel
substrate,
rock
slopes,
(
fish,
echiuroids,
ophiurids,
ascidians
and
shrimp)
27
wintering
grounds
17
or
soft
bottom
6
(
annelids,
crabs,
shrimp,
clams)
18
shallow
water
bays
over
the
continental
shelf
6
(
worms,
shrimps,
clams,
brittle
stars,
fish)
5,19
found
in
euryhaline
waters
6
(
ophiuroids,
polychaetes,
mollusks,
pelecypods,

sipunculids,
and
amphipods)
28
(
polychaetes,
echiuroids,
mollusks,
echinoderms,

benthic
fish,
and
urochordates)
29
(
clams,
polychaetes,
crabs,
shrimp,
scallops,
fish,

and
echinoderms)
24
(
primarily
polychaetes
and
molluscs)
39
(
polychaetes,
bivalves,
crabs,
and
brittle
stars)
44
(
adult
and
juvenile
geoduck
clams)
46
sand
or
mud
bottom
18
carnivore
NF
diurnal
feeder
6,8
affinity
for
shallow
waters
with
sandy
and
muddy
(
fish;
prim.
herring,
mysid,
shrimp,
squid)
30,31,32
substrates
all
along
the
Pacific
coast
6
(
fish,
mysids,
eggs)
13
found
in
euhaline
waters
6
(
epifaunal
molluscs,
infaunal
worms,
and
molluscs)
14
(
amphipods,
mysids,
fish)
33
(
sand
dwelling
worms,
isopods,
young
sand
crabs,

worm
mollusks,
and
barnacles)
45
(
euphausiids,
shrimp,
anchovies,
herring
sand
lance,

small
rockfish,
and
flatfish)
49
found
in
sand­
bottom
kelp
holdfast
and
rockybottom
kelp­
bed
habitat29
Page
398
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1167
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­
Flatfish
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54
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CD
CE
CF
CG
harbor
seals10,12,14,15
predacious
demersal
fishes,
sharks,
skates,
lingcod,
and
rockfish
17
harbor
seal
2,14
NF
NF
NF
sharks,
marine
mammals,
larger
flatfish,
and
pelagic
fish
6
Pacific
electric
ray,
Pacific
angel
shark,
and
other
NF
NF
NF
piscivorous
fishes
3,7
birds
3
sharks,
marine
mammals,
and
larger
fish
6
NF
NF
harbor
seals10
larger
fishes,
sharks,
and
marine
mammals
6
NF
NF
NF
Caspian
terns13
three
substocks:
Lepidopsetta
bilineata
bilineata
,
L.
bilineata
perarcuatus
,
and
L.
bilineata
mochigarei
2
Page
399
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1167
5­
2220e
­
Flatfish
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54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CH
CI
CJ
CK
CL
CM
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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400
of
1167
5­
2220e
­
Flatfish
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54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CN
CO
CP
CQ
CR
CS
3
3
yrs.
6­
7
yrs.
NG
NG
NG
NG
4
22.2
cm
(
2.2­
2.3
yrs.)
35.0
cm
(
5+
yrs.)
trawl
otolith
No
No
measurements
NF
NF
NF
NF
NF
NF
716
cm
(
2
yrs)
27
cm
(
4yrs)
small
mesh
trawl
otolith/
measurement
No
No
and
length
frequency
analysis
NF
NF
Nf
NF
NF
NF
Page
401
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1167
5­
2220e
­
Flatfish
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54
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56
57
58
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CT
CU
CV
CW
0.382­
0.51
(
males
ages
8­
10)

0.266­
0.357
(
females
ages
8­
10)

Golden
Gardens
pop:

0.329
(
males
3­
5)

0.357(
females
3­
5)
70
10
instantaneous
M:
0.00175
(
August)
to
NG
NG
Beverton­
Holt
equation
0.0075
per
day
(
September)

11females:
0.30
males:
0.38
(
annually)
NG
NG
NG
11
females(
3­
5
yrs):
0.36
males(
3­
5
yrs):
0.33
NG
NG
NG
11
females(
8­
10yrs):
0.27­
0.48
males(
8­
10yrs)
0.38­
0.5NG
NG
NG
14both:
0.1
(
Saratoga
Passage)
0.36­
0.46
(
Saratoga
PassageNG
NG
8
males:
0.25
males
0.45
males
0.70
males
CPUE
and
age
females:
0.20
0.25
females
0.45
females
composition
data
NF
NF
12
2.65
equation:
Nt2=
Nt1e­
z(
t1­
t2)
and
CPUE
data
6
female:
0.26
0.21
NG
CPUE
male:
0.53
NF
NF
NF
NF
Page
402
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1167
5­
2220e
­
Flatfish
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54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
CX
CY
CZ
DA
0+
english
sole
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
8­
14
years
NG
NG
NF
all
year
classes
NG
NG
round
stingray
4
6­
8
yrs
0.102
NG
NF
NF
NF
NF
NF
Page
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1167
5­
2220e
­
Flatfish
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
DB
DC
americana)
,
trematodes
(
Zoogonus
dextrociruus
)(
Otodistomum
veliporum
),
4
7.4
km
per
day
and
microsporidan
(
Glugea
spp.)
9
parasite:
isopod
(
Lironeca
vulgaris
)
13,15
parasites:
26
families,
35
spp.
(
see
reference)
14
parasites:
digenea
(
Neopodocotyloides
sinusaccus
)
16
coccurs
with
english
sole,
rex
sole,
Pacific
sanddab,
and
rock
sole
on
Oregon's
movements
up
to
350
miles
8
continental
shelf
7
seasonal
movement
to
deep
water
to
spawn
8
adults
in
California
move
offshore
in
fall
and
winter
to
spawn
9,1
summary
of
tagging
and
stocks
10
known
to
move
as
much
as
628
km
(
but
not
common)
17
parasites:
4
families,
5
spp.
(
see
reference)
14
NF
parasite:
isopod
(
Lironeca
vulgaris
)
15
parasites:
nematode
(
Philometra
spp
.)
10
5­
10
km
during
daytime,
up
to
30
km
at
night
11
parasites:
copepod
(
Argulus
borealis
),(
Lepeophtheirus
bifidus
),
and
majority
of
recoveries
made
Within
the
area
of
tagging,
support
(
Lepeophtheirus
parviventris)
11
discreteness
of
stocks
12
parasites:
28
families,
46
spp.
(
see
reference)
14
parasites:
digenea(
1
sp.),
cestoda
(
2
sp.),
copepoda
(
1
sp.)
and
nematoda(
3
sp.)
16
NF
parasite:
isopod
(
Lironeca
vulgaris
)
13,15
parasites:
8
families,
9
spp.
(
see
reference)
14
urban
bays
in
Puget
Sound,
WA
had
lower
conncentrations
of
sand
sole
eggs
than
rural
bays
due
to
the
presence
contaminants12
Page
404
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1167
5­
2220e
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Flatfish
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54
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
DD
DE
DF
DG
highly
migratory
12
Possibly
3
No
NF
juveniles
show
little
coastwise
or
bathymetric
movement
17
adults
move
inshore
and
northward
during
the
summer
to
feeding
grounds
and
offshore
and
southward
in
winter
to
spawning
grounds
17
Resident
4,13
NF
No
NF
Larger
fish
move
to
lower
regions
of
bays
and
estuaries,

adults
move
out
of
bays
to
spawn
4,13
Resident
14,15
NF
No
NF
migrate
to
shallow
water
in
summer
and
return
to
deeper
waters
in
winter
8,9,15
Resident
16
NF
No
NF
Adults
move
to
shallow
nearshore
waters
in
winter
to
spawn
16
no
pronounced
spawning
migration
noted
in
California,
a
distinct
spawning
mirgration
to
and
from
spawning
ground
has
been
noted
in
populaiton
in
the
straits
of
British
Columbia21
spawning9
Page
405
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1167
5­
2220e
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Flatfish
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104
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106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
A
B
C
D
E
Pleuronectidae
1,2
Pleuronichthys
coenosus
1,2
C­
O
turbot
southeastern
Alaska
(
Sitka
?)
to
Punta
Abreojos,
1,2
0
to
350
m.

C­
O
sole,
mottled
turbot
2,3,6
southern
Baja
California
2
Pleuronectidae
1,2
Pleuronichthys
verticalis
1,2
hornyhead
turbot
Oregon
to
at
least
Bahia
Magdelena,
southern
2
5­
201
m
sharpridged
turbot,
Baja
California
2
sharpridged
flounder,
sand
dab
2,3
Pleuronectidae
1,2
Pleuronichthys
ritteri
1,2
spotted
turbot
central
California
(
Morro
Bay)
to
Bahia
MagdaleNF,
2
1­
86
m
Ritter's
turbot
2,3
southern
Baja
California
2,3,4
Pleuronectidae
1,2
Pleuronichthys
decurrens
1,2
curlfin
turbot
Bering
Sea(?)
to
Isla
Cedros,
central
Baja
California
1,2
8­
532
m
California
turbot,
curlfinned
turbot,

curlfin
sole,
sanddab
2,
3
Pleuronectidae
1
Lyopsetta
exilis
slender
sole
110­
998
m
slender
flounder,
rough
sole
2
Cynoglossidae
2
Symphurus
atricauda
1,2
California
tonguefish
Puget
Sound,
Washington
to
Panama
2,3
1,2
1.5­
201
m
tongue
sole,
tonguefish
2,
3
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Kramer
et
al
1995
1­
Allen
1990
1­
Eschmeyer
and
Herald
1983
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
2­
Love
et
al
2000
2­
Love
et
al
2000
3­
Robert
Lea­
Cal.
Dept.
of
Fish
3­
Eschmeyer
and
Herald
1983
3­
Love
1996
3­
Ford
1965
Gulf
of
Alaska
(
east
of
Kodiak
Island­
north
Albatross
Bank)
to
southern
Baja
California1
Page
406
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
F
G
H
I
J
15Both:
W=
0.00000956L3.19554
(
L=
SL
(
mm))
NF
1,2,5,4
36
cmTL
NF
NF
NF
NF
1,2,4
37
cmTL
NF
NF
NF
NF
1
30
cmTL
NF
NF
NF
NF
1,2,4
37
cmTL
NF
NF
NF
NF
135
cm
TL
NF
NF
NF
NF
1,2
21
cmTL
NF
NF
1­
Haaker
1975
1­
Chamberlain
1979
1­
Love
et
al
2000
1­
Reed
and
Maccall
1988
2­
Chamberlain
1979
2­
Karpov
and
Kwiecien
1988
2­
Love
1996
2­
Smith
and
Nitsos
1969
3­
Arora
1951
3­
Lane
1975
3­
Rackowski
and
Pikitch
1989
3­
Van
Cleve
and
El
Sayad
1969
Page
407
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
K
L
M
N
O
P
Q
R
S
T
U
V
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Reed
and
Maccall
1988
2­
Smith
and
Nitsos
1969
3­
Van
Cleve
and
El
Sayad
1969
Page
408
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
W
X
Y
Z
AA
AB
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
NF
NF
dioecious
NF
NF
NF
220
yrs(
84
cm)
NF
dioecious
NF
NF
NF
NF
NF
dioecious
1­
Reed
and
Maccall
1988
1­
Reed
and
Maccall
1988
1­
Kramer
1991
1­
Kramer
and
Sunada
1992
1­
Sunada
et
al
1990
1­
Emmett
et
al
19
2­
Smith
and
Nitsos
1969
2­
Smith
and
Nitsos
1969
2­
Haaker
1975
2­
Love
1996
2­
Smith
1937
2­
Horton
1989
3­
Van
Cleve
and
El
Sayad
1969
3­
Van
Cleve
and
El
Sayad
1969
3­
Arora
1951
3­
Leos
1992
3­
Holland
1969
Page
409
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
AC
AD
AE
AF
AG
AH
AI
promiscuous
oviparous1,3,4,9
No
NF
No
NF
NF
promiscuous
oviparous1,3
No
NF
No
1
10
cm
1
17
cm
promiscuous
oviparous
1,3
No
NF
No
NF
NF
promiscuous
oviparous
1,3,4,10
No
NF
No
NF
NF
promiscuous
oviparous8,1,3
No
NF
NF
21NG
NG
12­
3
yrs
3­
5
yrs
promiscuous
oviparous
3
No
NF
No
NF
NF
91
1­
Moser
1996a
1­
Haaker
1975
1­
Orcutt
1950
1­
Love
1996
2­
Emmett
et
al
1991
2­
Reed
and
Maccall
1988
2­
Garrison
and
Miller
1982
2­
Emmett
et
al
1991
3­
Love
1996
3­
Love
1996
3­
C.
D.
F.
G
1982
Page
410
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
AJ
AK
AL
AM
AN
AO
AP
AQ
NF
NF
NF
NF
NF
NF
Puget
Sound
12
California7,36
San
Diego
24
CalCOFI
area
8
Pacific
coast
6
1
NG
1
NG
NG1
NG1
NG1
NF
Monterey
Bay
6,7
southern
California
6
NF
NF
NF
NF
NF
NF
CalCOFI
area
8
Pacific
coast
6
throughout
range39
NF
NF
NF
NF
NF
NF
Monterey
Bay
7
west
coast
9
14
cm
16
cm
NG
NG
NG
NF
Puget
Sound,
WA11
NG
NG
NG
Ng
Ng
northeast
Pacific30
CalCOFI
area
8
Pacific
coast6
NF
NF
NF
NF
NF
NF
southern
California35
California33
CalCOFI
area
8
Pacific
coast
6
1­
Hart
1973
1­
Moser
and
Watson
1990
2­
Tupen
1990
2­
Ford
1965
3­
Casillas
et
al
1998
3­
Kramer
and
Sunada
1992
Page
411
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
NF
*****
*****
*****
*****

*****
*****
*****
Batch
spawner4
*****
*****
*****
NF
*****
*****
*****
*****

*****
*****
*****
NF
NF
NF
*****
*****
1­
Caddell
et
al
1990
2­
Arora
1951
3­
Rackowski
and
Pikitch
1989
Page
412
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BD
BE
BF
BG
NF
NF
NF
5
11
days
6
12
days
NF
NF
NF
5
3­
4
days
16
approx.
4
days
NF
NF
NF
5,6,9
6­
7
days
16<
7
days
NF
NF
NF
NF
NF
NF
NF
NF
1­
Caddell
et
al
1990
1­
Emmett
et
al
1991
1­
Lassuy
1989
1­
Gadomski
et
al
1990
2­
Love
1996
2­
Ford
1965
2­
Peterman
et
al
1987
2­
Emmett
et
al
1991
3­
Ketchen
1956
3­
Love
1996
3­
Shvetsov
1979
3­
Love
1996
Page
413
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BH
BI
BJ
BK
BL
NF
16
8.2­<
17.0
mm
NF
1
1.2­
1.6
mm
pelagic
11,16
4
1.88
mm
3
1.20­
1.56
mm
121.28­
1.56
mm
(
CalCOFI)

NF
16
7.3­<
12.2
mm
NF
1
1.0­
1.2
mm
pelagic16,22,26
4
<
17
mm
(
coast)
3
1.00­
1.16
mm
5
19
mm
(
open
coast)
111.03­
1.11
mm
121.00­
1.16
mm
NF
16
6.0­<
12.7
mm
NF
1
0.9­
1.1
mm
pelagic
16
4
<
16
mm
(
coast)
8
3
0.94­
1.08
5
38
mm(
open
coast)
120.94­
1.08
mm
NF
16
>
21.0­<
29.4
NF
1
1.8­
2.1
mm
pelagic16,26
4
average:
1.44
mm
3,5
1.84­
2.08
mm
111.31­
1.50
mm
121.84­
2.08
mm
NF
NF
NF
101.47­
1.71
mm
pelagic21,16,11
11.5­
1.7
14approx.
2
months
4<
29
mm
(
coast
and
bays)
NF
1
0.74­
0.80
mm
pelagic
21
17,2119­
24.2
mm
SL(
coastal
waters)
3
0.71­
0.78
525­
51
mm(
open
coast)

1­
Allen
1988
1­
Allen
1990
1­
Reed
and
Maccall
1988
1­
Moser
1996a
1­
Valle
et
al
1999
2­
Gadomski
and
Peterson
1988
2­
Gadomski
and
Peterson
1988
2­
Emmett
et
al
1991
2­
Ahlstrom
et
al
1984
3­
Love
1996
3­
Emmett
et
al
1991
3­
Ahlstrom
et
al
1984
3­
Allen
and
Herbison
1990
121.20­
1.42
mm
(
King
Harbor)

Page
414
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
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109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BM
BN
BO
BP
BQ
primarily
offshore20
NF
1
3.7­
11.4
mm
pelagic
12
planktivore
NF
4
>
5.5
mm
3
3.9­>
11.4
NF
1
<
2.4­
11.0
mm
pelagic
12
planktivore
NF
3
2.4­>
11.0
coastal
waters2,24
93.16
mm
(
newly
hatched)
concentrated
in
midwater24
NF
1
2.1­
10.0
mm
pelagic
12
planktivore
NF
3
2.1­>
10.0
mm
NF
1
4.9­>
22mm
pelagic
12
planktivore
NF
4
>
3.9
mm
coastal
and
offshore
waters
2
3
4.9­>
21.0
mm
6
3.88­>
21.0
mm
93.88
mm
SL
(
at
hatching)

NF
75.6­
24.7
mm
SL
planktonic12
planktivore
NF
15.2­
24.7
mm
SL
found
in
bays
,
but
primarily
offshore20
mostly
inshore
up
to
250
miles
offshore11
NF
1
1.9­
10.8
mm
pelagic
19
planktivore
NF
3
1.9­
24.2
mm
1­
Casillas
et
al
1998
1­
Moser
1996a
1­
Valle
et
al
1999
1­
Allen
1990
1­
Gadomski
and
Boehlert
1984
2­
Emmett
et
al
1991
2­
Allen,
M.
J.
1982
2­
Fitch
and
Lavenberg
1971
3­
Ahlstrom
et
al
1984
3­
Emmett
et
al
1991
3­
Orcutt
1950
Page
415
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BR
BS
BT
BU
NF
NF
soft
or
rocky
bottom
7,22
shallow
sandy
areas
15
NF
NF
3
11.0
mm
to
maturity
soft
bottom
22
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
3
20.8
mm
to
maturity
sand
and
mud
of
bays
and
open
coast3,29
1­
Emmett
et
al
1991
1­
Fitch
and
Lavenberg
1971
1­
Kramer
1991
1­
Valle
et
al
1999
2­
Gadomski
and
Boehlert
1984
2­
Allen,
M.
J.
1982
2­
Kramer
and
Sunada
1992
2­
Allen
1988
3­
Watson
and
Davis
1989
3­
Emmett
et
al
1991
3­
Moser
1996a
3­
Love
1996
estuaries
used
as
nursery
and
feeding
areas,
most
leave
after
1st
year26
Page
416
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BV
BW
BX
BY
benthic
carnivore
(
polychaetes)
8,21
benthic
stalker
7
diurnal
and
nocturnal
8
blue
rockfish13
(
small
polychaetes,
molluscs,
and
crustaceans)
17
nocturnal
feeder
9
carnivore
bottom
feeder,
pulls
invertebrates
NF
California
halibut,
thornback,

(
polychaetes,
crustaceans,
anemones,
lamp
shells)
22
from
sand
8
and
other
fish
carnivores
lunge
feeder
8
NF
California
halibut,
thornback,

(
polychaetes,
anemones)
22
and
other
fish
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Emmett
et
al
1991
1­
Allen
1990
1­
Emmett
et
al
1991
1­
Allen
1990
2­
Allen
1990
2­
Hulberg
and
Oliver
1979
2­
Allen,
M.
J.
1982
2­
Leos
1992
3­
Wertz
and
Domier
1997
3­
Ford
1965
3­
Goldberg
and
Pham
1987
3­
Ford
1965
Page
417
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
BZ
CA
CB
CC
(
fish,
crustaceans,
worms,
molluscs)
5,19
(
adult
and
juvenile
geoduck
clams)
46
soft
or
rocky
bottom
4,17
benthic
carnivore
(
polychaetes)
2,34
benthic
stalker
2
diurnal
and
nocturnal
3,14
shallow
sandy
areas
12
(
small
polychaetes,
molluscs,
and
crustaceans)
21
digs
prey
out
of
sand
8
nocturnal
feeder
10
(
worms,
small
crustaceans,
and
small
fish)
5
(
annelids,
crabs,
shrimp,
clams)
18
(
polychaetes,
clams,
sm.
fish,
amphipods,
nudibranchs,

techibranchs)
35
soft
bottom
17
benthic
carnivore
(
polychaetes,
bivalve
siphons)
2
benthic
stalker
2
diurnal
3,4
abundant
in
shallow
water
areas,
buries
itself
(
polychaetes,
crustaceans,
anemones,
lamp
shells)
36
bottom
feeder,
pulls
invertebrates
from
sand
8,15
deeply
in
sediment
19
(
clams,
polychaetes)
35
(
worms,
clam
siphons)
5
found
on
soft
bottom
in
bays
and
along
benthic
carnivores
lunge
feeder
15
diurnal
3
semiprotected
coast
17,20
(
sessile
anemones
and
polychaetes)
36,37
locates
prey
by
sight
2
(
polychaetes,
clams,
amphipods)
35
sand
and
mud
bottom
21
carnivore
(
echiuroid
probosces
and
polychaetes)
2
benthic
stalker
2
most
likely
diurnal
and
(
polychaetes)
13
dig
in
bottom
for
prey
8
nocturnal
feeder
3
(
worms,
gammarid
amphipods,
worm­
like
peanut
worms)
5
demersal
on
shelf
to
mid­
slope17
NF
northern
elephant
seals4
sand
and
mud
bottom
5
benthic
carnivore
(
gammarid
amphipods)
2
benthic
searcher
2
nocturnal
3,4,15,18
remains
buried
in
the
bottom
during
the
day23
(
amphipods,
crabs,
worms)
5
prey
is
located
by
olfaction,
gustation,
and
(
microcrustaceans,
polychaetes,
and
molluscs)
38
mechanoreception
2
1­
Allen
1990
1­
Allen
1990
1­
Allen
1990
1­
Haaker
1975
2­
Haaker
1975
2­
Allen,
M.
J.
1982
2­
Allen,
M.
J.
1982
2­
Emmett
et
al
1991
3­
Kravitz
et
al
1977
3­
Emmett
et
al
1991
3­
Emmett
et
al
1991
3­
Allen,
M.
J.
1982
no
clear
diurnal
or
nocturnal
mode19
sandy
bottom
dwellers,
often
found
in
vicinity
of
rocks
and
algae5
rests
immobile
on
sandy
substrate
at
all
hours
of
day
and
night,
exposed
or
under
a
thin
layer
of
sediment26
found
in
sand­
bottom
kelp
holdfast
and
rockybottom
kelp­
bed
habitat29
(
specked
sanddab,
herring,
anchovies,
crustaceans,

worms,
and
mollusks)
29
(
squid,
and
smaller
fish,
including
their
own
species,

other
flatfishes,
sardines,
grunion,
jacksmelt,
queenfish,

corbina,
walleye
surfperch,
tubenose,
and
klipfish)
45
(
polychaetes,
nudibranchs,
crustacean
eggs,

tectibranchs,
brittle
stars)
35
deeper
water
(
148­
195
m),
mud­
bottom
association27
(
shrimps,
amphipods,
salps,
mysid
shrimps,
worms,
and
krill)
5
Page
418
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CD
CE
CF
CG
NF
NF
NF
NF
leopard
shark,
Pacific
angel
shark,
and
California
sea
lion
8
NF
NF
NF
leopard
shark,
Pacific
angel
shark,
and
California
sea
lion
8
NF
NF
NF
harbor
seals
2
NF
NF
NF
harbor
seal14
NF
NF
NF
California
sea
lion18
Arctic
loons,
pelagic,
Brandt's,
and
double­
breasted
cormorants
2NF
NF
NF
leopard
shark,
rockfish,
California
halibut,
angel
sharks,

cormorants,
and
Pacific
electric
rays
9
1­
Haaker
1975
1­
Hedgcock
and
Bartley
1988
2­
Love
1996
2­
Garrison
and
Miller
1982
3­
Emmett
et
al
1991
Page
419
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CH
CI
CJ
CK
CL
CM
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Hedgcock
and
Bartley
1988
1­
Hedgcock
and
Bartley
1988
1­
Hedgcock
and
Bartley
1988
1­
Hedgcock
and
Bartley
1988
Page
420
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CN
CO
CP
CQ
CR
CS
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Nf
NF
NF
NF
NF
NF
Nf
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Nf
NF
NF
NF
1­
Allen
1988
2­
Sunada
et
al
1990
3­
Castillo
et
al
1995
Page
421
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CT
CU
CV
CW
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Reed
and
Maccall
1988
2­
Ford
1965
3­
Peterman
et
al
1987
Page
422
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
CX
CY
CZ
DA
NF
NF
NF
NF
NF
NF
NF
Pleuronichthys
ritteri
5
NF
NF
NF
Pleuronichthys
verticalis
5
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
speckled
sanddab
2
1­
Allen
1990
2­
Ford
1965
3­
Casillas
et
al
1998
Page
423
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
DB
DC
parasites:
copepod
(
Lepeophtheirus
hospitalis
)
11
NF
parasites:
3
families,
4
spp.
(
see
reference)
14
parasite:
isopod
(
Lironeca
vulgaris
)
13
NF
parasites:
2
families,
2
spp.
(
see
reference)
14
NF
NF
parasite:
isopod
(
Lironeca
vulgaris
)
13
NF
parasites:
4
families,
4
spp.
(
see
reference)
14
parasites:
7
families,
9
spp.
(
see
reference)
14
NF
parasite:
isopod
(
Lironeca
vulgaris
)
15
NF
1­
Allen
1990
1­
Tupen
1990
2­
Chamberlain
1979
2­
Domier
and
Chun
1995
3­
Hubbs
and
Kuronama
1942
3­
Kramer
and
Sunada
1992
Page
424
of
1167
5­
2220e
­
Flatfish
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
DD
DE
DF
DG
NF
NF
No
NF
NF
NF
No
1
NF
NF
NF
No
NF
NF
NF
No
NF
NF
NF
No
NF
NF
NF
No
NF
1­
Domier
and
Chun
1995
1­
Tupen
1990
1­
Allen,
M.
J.
1982
1­
Day
1976
2­
Kramer
and
Sunada
1992
2­
Day
1976
3­
Allen
1990
3­
Alverson
1957
Page
425
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
A
B
C
D
E
and
Game
(
personal
communication)
4­
Love
1996
4­
Eschmeyer
and
Herald
1983
5­
Rackowski
and
Pikitch
1989
5­
Moser
1996a
6­
Hart
1973
6­
Miller
and
Lea
1972
7­
Kramer
1991
7­
Goldberg
1987
8­
Smith
1937
8­
Hart
1973
9­
Reilly
et
al
1994
Page
426
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
F
G
H
I
J
4­
Ford
1965
4­
Haaker
1975
4­
Eschmeyer
and
Herald
1983
4­
Best
1963
5­
Orcutt
1950
5­
Hart
1973
5­
Pedersen
1975
6­
Peterman
et
al
1987
6­
Pearson
and
Owen
1992
6­
Ketchen
and
Forrester
1966
7­
Smith
and
Nitsos
1969
7­
Lane
1975
7­
Lane
1975
8­
Yoklavich
1982a
8­
Forrester
and
Thomson
1969b
9­
Budd
1940
10­
Ketchen
and
Forrester
1966
11­
Lane
1975
12­
Forrester
and
Thomson
1969b
13­
Forrester
and
Thomson
1969a
14­
Yoklavich
1982b
15­
Quast
1968a
16­
Beardsley
1969
Page
427
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
K
L
M
N
O
P
Q
R
S
T
U
V
4­
Best
1963
5­
Pedersen
1975
6­
Ketchen
and
Forrester
1966
7­
Lane
1975
8­
Forrester
and
Thomson
1969b
Page
428
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
W
X
Y
Z
AA
AB
4­
Best
1963
4­
Best
1963
4­
Ford
1965
4­
Ford
1965
4­
Ketchen
and
Forrester
1966
5­
Pedersen
1975
5­
Pedersen
1975
5­
Smith
1937
5­
Forrester
1969(
a)
5­
Forrester
and
Thomson
1969b
6­
Ketchen
and
Forrester
1966
6­
Ketchen
and
Forrester
1966
6­
Campana
1984a
6­
Hart
1973
6­
Shubnikov
and
Lisovenko
1964
7­
Lane
1975
7­
Lane
1975
7­
Larouche
et
al
1982
7­
Ketchen
and
Forrester
1966
8­
Forrester
and
Thomson
1969b
8­
Forrester
and
Thomson
1969b
8­
Yoklavich
1982b
8­
Lane
1975
9­
Emmett
et
al
1991
9­
Reilly
et
al
1994
10­
Dygert
1990
11­
Shi
et
al
1996
12­
Larouche
1982
13­
Horton
1989
14­
Pattison
and
McAllister
1990
15­
Allen
1988
16­
Orcutt
1950
17­
Campana
1984c
18­
Campana
1983
19­
MBC
Appl.
Environ.
Sci.
1987
20­
Kramer
1991
21­
Campana
and
Neilson
1982
22­
Reilly
et
al
1994
23­
Beardsley
1969
Page
429
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
AC
AD
AE
AF
AG
AH
AI
4­
Hart
1973
4­
Arora
1951
4­
Arora
1951
5­
Casillas
et
al
1998
5­
Chamberlain
1979
5­
Ford
1965
6­
Pearson
and
Owen
1992
6­
Orcutt
1950
6­
Orcutt
1950
7­
Thomas
1991
7­
Ketchen
1956
7­
Castillo
1995
8­
Matarese
et
al
1989
8­
Thomas
1991
8­
Ketchen
1956
9­
Garrison
and
Miller
1982
9­
Casillas
et
al
1998
9­
Forrester
1969a
10­
Fitch
1963
10­
Baxter
1999d
10­
Harry
1959
11­
Young
1960
11­
Emmett
et
al
1991
12­
Heimann
1960
12­
Pearson
and
Owen
1992
13­
MBC
Appl.
Environ.
Sci.
1987
14­
Forrester
and
Thomson
1969b
15­
Smith
1937
16­
Love
and
Brookes
1990
17­
Baxter
1999d
18­
Garrison
and
Miller
1982
19­
Reilly
et
al
1994
20­
Heimann
1960
21­
Hart
1973
Page
430
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
AJ
AK
AL
AM
AN
AO
AP
AQ
4­
Horton
1989
4­
Lavenberg
et
al
1986
5­
Emmett
et
al
1991
6­
Love
1996
7­
Budd
1940
8­
Moser
1996a
9­
Casillas
et
al
1998
10­
Arora
1951
11­
Hart
1973
12­
Smith
1937
13­
Goldberg
and
Pham
1987
14­
Fitch
and
Lavenberg
1971
15­
Chamberlain
1979
16­
Orcutt
1950
17­
Castillo
1995
18­
Kruse
and
Tyler
1983
19­
Harry
1959
20­
Gregory
and
Jow
1976
21­
Alderdice
and
Forrester
1971
22­
Lane
1975
23­
Eldridge
1975
24­
Eigenman
1891
25­
Shvetsov
1981
26­
Horton
1989
27­
Manzer
1947
28­
Wang
1981
29­
Castillo
et
al
1993
30­
Matarese
et
al
1989
31­
Haugen
1992
32­
Pearson
and
Owen
1992
33­
Fitch
and
Lavenberg
1975
34­
Breder
and
Rosen
1966
35­
Baxter
1999d
36­
Garrison
and
Miller
1982
37­
Kramer
and
Smith
1973
38­
Hardy
et
al
1987
39­
Fitch
1963
40­
Reilly
et
al
1994
41­
Thornburgh
1978
42­
Young
1960
43­
Heimann
1960
44­
Best
1960
Page
431
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
4­
Love
1996
5­
Hart
1973
6­
Casillas
et
al
1998
7­
Ford
1965
8­
Kramer
and
Sunada
1992
9­
Baxter
1999d
10­
Garrison
and
Miller
1982
11­
Orcutt
1950
12­
Lassuy
1989
13­
Harry
1959
Page
432
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
BD
BE
BF
BG
4­
Wang
1981
4­
Hart
1973
4­
Garrison
and
Miller
1982
4­
Boehlert
and
Mundy
1987
5­
Forrester
1969a
5­
Budd
1940
6­
Harry
1959
6­
Hart
1973
7­
Casillas
et
al
1998
7­
Pearson
and
Owen
1992
8­
Eschmeyer
and
Herald
1983
8­
Alderdice
and
Forrester
1971
9­
Shvetsov
1979
9­
Casillas
et
al
1998
10­
Garrison
and
Miller
1982
10­
Hardy
et
al1987
11­
Thornburgh
1978
11­
Orsi
1968
12­
Heimann
1960
12­
Gadomski
and
Cadell
1995
13­
Thomson
1991
14­
Garrison
and
Miller
1982
15­
Hardy
et
al
1987
16­
Fitch
1963
17­
Heimann
1960
Page
433
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
BH
BI
BJ
BK
BL
4­
Emmett
et
al
1991
4­
Kramer
1991
4­
Hart
1973
4­
Emmett
et
al
1991
5­
Sakuma
and
Larson
1995
5­
Love
1996
5­
Casillas
et
al
1998
5­
Arora
1951
6­
Castillo
1995
6­
Sakuma
1992
6­
Thornburgh
1978
6­
Sakuma
1992
7­
Campana
1984d
7­
Thornburgh
1978
7­
Horton
1989
7­
Casillas
et
al
1998
8­
Thornburgh
1978
8­
Hart
1973
8­
Eldridge
1975
8­
Hart
1973
9­
Larouche
et
al
1982
9­
Gunderson
et
al
1990
9­
Wang
1981
9­
Kramer
and
Sunada
1992
10­
Rosenberg
1980
10­
Misitano
1976
10­
Matarese
et
al
1989
10­
Budd
1940
11­
Hart
1973
11­
Shi
et
al
1995
11­
Fitch
1963
11­
Love
1996
12­
Larouche
1982
12­
Krygier
and
Pearcy
1986
12­
Sumida
et
al
1979
12­
Alderdice
and
Forrester
1968
13­
Pearcy
et
al
1977
13­
Casillas
et
al
1998
13­
Gregory
and
Jow
1976
14­
Baxter
1999d
14­
Eldridge
1975
14­
Alderdice
and
Forrester
1971
15­
Heimann
1960
15­
Emmett
et
al
1991
15­
Eldridge
1975
16­
Sumida
et
al
1979
16­
Moser
1996a
17­
Moser
1996a
17­
Forrester
1969a
18­
Rackowski
and
Pikitch
1989
18­
Shubnikov
and
Lisovenko
1964
19­
Ahlstrom
et
al
1984
19­
Smith
1937
20­
Policansky
1982
20­
Hardy
et
al
1987
21­
Baxter
1999d
21­
Matarese
et
al
1989
22­
Breder
and
Rosen
1966
23­
Baxter
1999d
24­
Plummer
et
al
1983
25­
Kramer
and
Smith
1973
26­
Fitch
1963
27­
Heimann
1960
28­
Best
1960
Page
434
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
BM
BN
BO
BP
BQ
4­
Hart
1973
4­
Arora
1951
4­
Hart
1973
5­
Misitano
1976
5­
Sakuma
1992
5­
Emmett
et
al
1991
6­
Casillas
et
al
1998
6­
Casillas
et
al
1998
6­
Casillas
et
al
1998
7­
Matarese
et
al
1989
7­
Leos
1992
7­
Emmett
et
al
1991
8­
Baxter
1999d
8­
Orcutt
1950
8­
Kruse
and
Tyler
1989
9­
Fitch
1963
9­
Rosenberg
1980
9­
Gadomski
and
Boehlert
1984
10­
Allen
1990
10­
MBC
Appl.
Environ.
Sci.
1987
11­
Love
1996
11­
Lane
1975
12­
Moser
1996a
12­
Barraclough
1967c
13­
Eldridge
1975
13­
Robinson
et
al
1968
14­
Smith
1937
14­
Watson
and
Davis
1989
15­
Hardy
et
al
1987
16­
Allen
1988
17­
MBC
Appl.
Environ.
Sci.
1987
18­
Wang
1981
19­
Matarese
et
al
1989
20­
Pearcy
and
Myers1974
21­
Moser
and
Pommeranz
1998
22­
Garrison
and
Miller
1982
23­
Watson
and
Davis
1989
24­
Barnett
et
al
1984
25­
Moser
and
Pommeranz
1999
26­
Plummer
et
al
1983
Page
435
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
BR
BS
BT
BU
4­
Casillas
et
al
1998
4­
Emmett
et
al
1991
4­
Kramer
and
Sunada
1992
5­
Lassuy
1989
5­
Harry
1959
5­
Emmett
et
al
1991
6­
Casillas
et
al
1998
6­
Allen
1990
7­
Ahlstrom
et
al
1984
7­
Feder
et
al
1974
8­
Sakuma
1992
8­
Kravitz
et
al
1977
9­
Allen
1988
9­
Casillas
et
al
1998
10­
Gadomski
et
al
1990
10­
Toole
1980
11­
Hart
1973
12­
Rosenberg
1980
13­
Shi
et
al
1995
14­
Olson
and
Pratt
1973
15­
Becker
1988
16­
Eschmeyer
and
Herald
1983
17­
Fitch
and
Lavenberg
1971
18­
Lane
1975
19­
Norcross
et
al
1995
20­
Norcross
et
al
1999
21­
Miller
1965
22­
Moser
1996a
23­
MBC
Appl.
Environ.
Sci.
1987
24­
Krygier
and
Pearcy
1986
25­
Kramer
and
S.
W.
F.
Center
1991
26­
Baxter
1999d
27­
Pearcy
and
Myers
1974
28­
Westerheim
1955
29­
Fitch
and
Lavenberg
1975
30­
Limbaugh
1955
Page
436
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
BV
BW
BX
BY
4­
Love
1996
4­
Orcutt
1950
4­
Miller
1967
4­
Toole
1980
5­
Casillas
et
al
1998
5­
Toole
1980
5­
Lassuy
1989
5­
Rosenberg
1980
6­
Sakuma
1992
6­
Lane
1975
6­
Lane
1975
6­
Ambrose
1976
7­
Hulberg
and
Oliver
1979
7­
Allen,
M.
J.
1982
7­
Casillas
et
al
1998
7­
Allen,
M.
J.
1982
8­
Allen,
M.
J.
1982
8­
Luckinbill
1969
8­
Hobson
et
al
1981
8­
Emmett
et
al
1991
9­
Ford
1965
9­
Becker
1984
9­
Casillas
et
al
1998
10­
Orcutt
1950
10­
Fitch
and
Lavenberg
1971
11­
McCall
1992
12­
Haugen
1992
12­
Miller
1967
13­
Love
and
Ebeling
1978
13­
Miller
1965
14­
Thornburgh
1978
15­
Toole
1980
16­
Houge
and
Carey
1982
17­
Becker
1984
18­
Lane
1975
19­
Fitch
and
Lavenberg
1971
20­
Holladay
and
Norcross
1995
21­
Hobson
et
al
1981
22­
Luckinbill
1969
23­
Allen
1988
24­
Plummer
et
al
1983
Page
437
of
1167
5­
2220e
­
Flatfish
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169
170
171
172
173
174
175
176
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178
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180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
BZ
CA
CB
CC
4­
Feder
et
al
1974
4­
Wertz
and
Domier
1997
4­
Haaker
1975
4­
Love
1996
5­
Love
1996
5­
Love
1996
5­
Hulberg
and
Oliver
1979
5­
Ford
1965
6­
Casillas
et
al
1998
6­
Leos
1992
6­
Kravitz
et
al
1977
6­
Miller
1965
7­
Ambrose
1976
7­
Hulberg
and
Oliver
1979
7­
Pearcy
and
Handcock
1978
7­
Ambrose
1976
8­
Orcutt
1950
8­
Kravitz
et
al
1977
8­
Love
1996
8­
Miller
1967
9­
Ketchen
1956
9­
Valle
et
al
1999
9­
Ford
1965
9­
Lassuy
1989
10­
Yoklavich
1982a
10­
Thornburgh
1978
10­
Boothe
1967
10­
Becker
1984
11­
Hart
1973
11­
Ambrose
1976
11­
Ambrose
1976
11­
Lane
1975
12­
Becker
1988
12­
Ford
1965
12­
Lassuy
1989
12­
Lane
et
al
1979
13­
Emmett
et
al
1991
13­
Hilaski
1972
13­
Hilaski
1972
13­
Casillas
et
al
1998
14­
Eschmeyer
and
Herald
1983
14­
Barry
et
al
1996
14­
Lane
1975
14­
Best
1963
15­
Fitch
and
Lavenberg
1971
15­
Porter
1964
15­
Luckinbill
1969
15­
Telder
1981
16­
Lane
1975
16­
Allen
1988
16­
Onate
1991
16­
Onate
1991
17­
Moser
1996a
17­
Boothe
1967
17­
Skalkin
1968
18­
Miller
1965
18­
Fraser
1923
18­
Baxter
1999d
19­
Ford
1965
19­
Hart
1973
19­
Hobson
et
al
1981
20­
Allen
and
Herbison
1990
20­
Ketchen
1956
20­
Becker
and
Chew
1987
21­
C.
D.
F.
G.
1982
21­
Becker
1984
22­
MBC
Appl.
Environ.
Sci.
1987
22­
Kravitz
et
al
1977
23­
Baxter
1999d
23­
Lane
1975
24­
Garrison
and
Miller
1982
24­
Forrester
and
Thomson
1969b
25­
Radtke
1966
25­
Lang
et
al
1995
26­
Hobson
et
al
1981
26­
Shubnikov
and
Lisovenko
1964
27­
Pearcy
1978
27­
Skalkin
1968
28­
Grossman
1982
28­
Horton
1989
29­
Limbaugh
1955
29­
Casillas
et
al
1998
30­
Miller
1967
31­
Houge
and
Carey
1982
32­
Miller
1965
33­
Thornburgh
1978
34­
Hobson
et
al
1981
35­
Fitch
1963
36­
Luckinbill
1969
37­
Moser
1996a
38­
Telders
1981
39­
Skalkin
1968
40­
Plummer
et
al
1983
41­
Fitch
and
Lavenberg
1975
42­
Quast
1968d
43­
Becker
and
Chew
1987
44­
Reilly
et
al
1994
45­
Limbaugh
1955
46­
Goodwin
and
Pease
1989
47­
Birtwell
et
al
1984
48­
Heimann
1960
49­
Best
1960
Page
438
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169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
CD
CE
CF
CG
4­
Ford
1965
5­
Allen,
M.
J.
1982
6­
Casillas
et
al
1998
7­
Fitch
and
Lavenberg
1971
8­
Luckinbill
1969
9­
Telder
1981
10­
Olesiuk
et
al
1990
11­
Fitch
and
Lavenberg
1975
12­
Treacy
1985
13­
Harvey
1982
14­
Trumble
1995
15­
Torok
1994
16­
Parkin
1998
17­
Heimann
1960
18­
Hawes
1983
Page
439
of
1167
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­
Flatfish
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156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
CH
CI
CJ
CK
CL
CM
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165
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167
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169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
CN
CO
CP
CQ
CR
CS
4­
Ketchen
and
Forrester
1966
5­
Orcutt
1950
6­
Misitano
1976
7­
Forrester
and
Thomson
1969b
Page
441
of
1167
5­
2220e
­
Flatfish
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
CT
CU
CV
CW
4­
Van
Cleve
and
El
Sayad
1969
5­
Shi
et
al
1995
6­
Forrester
and
Thomson
1969b
7­
Tenera
Environmental
Services
2000
8­
Ketchen
and
Forrester
1966
9­
Kramer
1991
10­
Shi
et
al
1996
11­
Lassuy
1989
12­
Lane
1975
13­
Helvey
and
Witzig
1990
14­
Menasveta
1958
Page
442
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2220e
­
Flatfish
155
156
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159
160
161
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163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
CX
CY
CZ
DA
4­
Ketchen
1972
5­
Luckinbill
1969
Page
443
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1167
5­
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Flatfish
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156
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164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
DB
DC
4­
Love
1996
4­
Hart
1973
5­
Haugen
1992
5­
Pattie
1969
6­
Casillas
et
al
1998
6­
Jow
1969
7­
Kravitz
et
al
1977
7­
Forrester
1969b
8­
Lassuy
1989
8­
Alverson
1957
9­
Olson
and
Pratt
1973
9­
C.
D.
F.
G
1982
10­
Horton
1989
10­
Wertz
and
Domier
1997
11­
Fraser
1919
11­
Casillas
et
al
1998
12­
Hardy
et
al
1987
12­
Forrester
and
Thomson
1969b
13­
Keusink
1979
13­
Pearson
and
Owen
1992
14­
Love
and
Moser
1983
14­
Thomas
1991
15­
Bennett
1993
15­
Menasveta
1958
16­
Arai
1969
16­
Beardsley1969
Page
444
of
1167
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2220e
­
Flatfish
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156
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159
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162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
DD
DE
DF
DG
4­
Emmett
et
al
1991
4­
Beardsley
1969
5­
C.
D.
F.
G.
1982
6­
Rackowski
and
Pikitch
1989
7­
Chamberlain
1979
8­
Hart
1973
9­
Love
1996
10­
Jackson
1981
11­
Forrester
1969b
12­
Alverson
1957
13­
Lane
1975
14­
Forrester
and
Thomson
1969b
15­
Horton
1989
16­
Casillas
et
al
1998
17­
MBC
Appl.
Environ.
Sci.
1987
18­
Sakuma
et
al
1999
19­
Haugen
1992
20­
Pearson
and
Owen
1992
21­
Heimann
1960
Page
445
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Flatfish
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5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Gobiidae
1,2
Lythrpnus
dalli
1,2
bluebanded
goby
Morro
Bay,
central
California
to
Afuera
Is.,
Peru
and
1
intertidal­
76m
Catalina
goby
1,2
Islas
Galapagos
1
Gobiidae
1,2
Lythrypnus
zebra
1,2
zebra
goby
2
Carmel
Bay,
central
California
to
Isla
Clarion,
Mexico
1,2,3
1,2
intertidal­
97
m
Gobiidae
1,2
Gillichthys
mirabilis
1,2
longjaw
mudsucker
2
Tomales
Bay,
northern
California
to
Gulf
of
California
1,2,3
1
shallow
water
of
bays
and
coastal
sloughs
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Gates
and
Frey
1974
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Love
1996
2­
Eschmeyer
and
Herald
1983
2­
Eschmeyer
and
Herald
1983
3­
Love
1996
TAXONOMY
RANGE
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
F
G
H
I
J
K
L
M
N
O
Total
Length
(
cm)
Max.
reported
Geographic
area
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
Female
Both
Male
Female
Both
NF
NF
1,2
6.4
cm
TL
NF
NF
NF
NF
NF
NF
NF
NF
NF
1,2
5.7
cm
TL
NF
NF
NF
NF
NF
NF
NF
NF
NF
1,2
21
cm
TL
NF
NF
NF
NF
NF
NF
NF
1­
Love
et
al
2000
2­
Love
1996
Linf
in
cm
(
SE)
k
(
SE)

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9
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
P
Q
R
S
T
U
V
W
X
Y
r2
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.

NF
NF
NF
NF
NF
NF
NF
NF
NF
1
NF
NF
NF
NF
NF
NF
NF
NF
NF
1
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Behrents
1983
Growth
Parameters
to
(
SE)

AGE
and
GROWTH
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
Z
AA
AB
AC
AD
AE
AF
Age
composition
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
1
19
months
NF
can
be
simultaneous
hermophrodites
1
polygynous1,2
oviparous
1,2
NF
2
2
yrs.
simult.
hermaphrodites,
but
do
not
reproduce
as
males
and
females
simultaneously
2
323
months
NF
simultaneous
hermaphrodites
1
polygynous1
oviparous
1,2
yes,
male
guards
eggs
3
NF
simult.
hermaphrodites,
but
do
not
reproduce
as
males
and
females
simultaneously
2
1
2
yrs.
NF
NF
NF
oviparous
1
yes,
eggs
guarded
by
NF
42
yrs.
(
135­
140
mm
SL)
male
during
incubation
2,4
male
builds
nest
for
eggs7
1­
Love
1996
1­
Love
1996
1­
Behrents
1983
1­
Moser
1996a
1­
Feder
et
al
1974
2­
Steele
1997
2­
St.
Mary
1996
2­
Turner
et
al
1969
2­
Love
1996
2­
Eschmeyer
and
Herald
1983
3­
Behrents
1983
3­
Moser
1996a
4­
Walker
1961
4­
Moyle
1976
5­
Wiley
1976
6­
Turner
et
al
1969
7­
C.
D.
F.
G.
1982
8­
Limbaugh
1955
Breeding
Behavior
male
cleans
area
and
builds
nest
for
eggs6
yes,
male
protects
eggs
until
hatched1,2,5,6,8
Sexua
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13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AG
AH
AI
AJ
AK
AL
AM
AN
Total
length
(
age)
at
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
2NG
16.3­
20.9
mm
(
NG)
NG
NG
NG
NG
NG
319.8
mm
(
NG)
19.0
mm(
NG)
NG
NG
NG
NG
NG
males
larger2
2NG
19.4­
21.6
mm
(
NG)
NG
NG
NG
NG
NG
1
5­
8
cm
(
2
yrs)
5­
8
cm
(
2
yrs)
NG
NG
NG
NG
NG
1­
Behrents
1983
1­
Love
1996
2­
Wiley
1976
2­
Behrents
1983
3­
Walker
1961
3­
Wiley
1976
1st
maturity
50%
maturity
male
is
distiguished
by
genital
papilla3
male
genital
papilla
is
elongate
and
pointed,
female
genital
papilla
is
broad
and
truncated1,2
male
genital
papilla
is
elongate
and
pointed,
female
genital
papilla
is
broad
and
truncated1,2
al
dimorphism
REPRODUCTION
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
Fertilized
eggs
present
Both
Parturition
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.

NF
CalCOFI
area
1
Pacific
coast
2
*****
*****
*****
*****
*****

Catalina
Island,
CA6
Punta
Banda,
Mexico7
NF
CalCOFI
area
1
Pacific
coast
2
*****
*****
*****

Catalina
Island,
CA6
Punta
Banda,
Mexico7
1
CalCOFI
area
1
San
Francisco
Bay,
CA
Salton
Sea
,
CA5
California
4,5,8,9
Pacific
coast
2
*****
*****
*****

Pacific
coast
3
1­
Barlow
1963
1­
Moser
1996a
2­
Love
1996
3­
Eschmeyer
and
Herald
1983
4­
Moyle
1976
5­
Barlow
1963
6­
Behrents
1983
7­
Wiley
1976
8­
Breder
and
Rosen
1966
9­
C.
D.
F.
G.
1982
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BC
BD
BE
BF
Mean
Fecundity
Size
specific
eggs/
g
Number
of
eggs
fecundity
eq
Spawning
Frequency
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)

multiple
spawners4
3164­
1,884
eggs
NF
1Y=
131.2X­
2310.3;
r2=
0.81(
Bird
Rock,
Santa
Catalina
Is.)

392­
523
eggs/
nest
Y=
132.7X­
2592.7;
r2=
0.56
(
Arrow
Point,
Santa
Catalina
Is.)

51,058
eggs
(
avg.)

5100­
1,000
eggs
some
single
some
multiple,
1
170­
800
eggs
NF
1Y=
60.3X­
1098.8;
r2=
0.42
(
Arrow
Point,
Santa
Catalina
Is.)

most
males
and
females
3166­
800
eggs
die
after
spawning
5627
eggs
(
avg.)

and
nest
guarding
1
multiple
spawners4
multiplespawners
2,5,6
1,2,4,6
4,000­
9,000
eggs
per
spawninNF
NF
2­
3
times
per
season
3,6
1­
Love
1996
1­
Love
1996
1­
Behrents
1983
2­
Moyle
1976
2­
Barlow
1963
3­
Barlow
1963
3­
Behrents
1983
4­
Behrents
1983
4­
Fitch
and
Lavenberg
1975
5­
deVlaming
1972
5­
Waples
and
Rosenblatt
1987
6­
Fitch
and
Lavenberg
1975
6­
C.
D.
F.
G.
1982
Y=
62.07X­
1013.5;
r2=
0.60
(
Big
Fisherman's
Cove,
Santa
Catalina
Is.)

Y=
38.0X­
680.3;
r2=
0.44
(
Big
Fisherman's
Cove,
Santa
Catalina
Is.)

Fecundity
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51
52
53
BG
BH
BI
BJ
BK
BL
Eggs
Size
at
settlement
Size
Egg
duration
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
38­
10.5
days
1,42+
months
1
<
12.7
mm
(
reefs)
176
NF
20.4­
0.6
mm
spawned
in
empty
shells
1,2,9,10,1
2
2­
3
months
2.3
9­
11
mm
(
benthos)
demersal
and
adhesive
3
360
days
38.5­
11
days
1
2­
5
months
38­
11
mm
(
benthos)
NF
20.5
mm
(
avg.)
demersal
and
adhesive
3,4
360
days
1,2,4,5
10­
12
days
NF
NF
NF
1
2.8­
3.4
mm
demersal
and
adhesive
3,4
females
lay
eggs
in
nests
constructed
by
males
2,5,6
1­
Eschmeyer
and
Herald
1983
1­
Love
1996
1­
Love
1996
1­
Moser
1996a
1­
Feder
et
al
1974
2­
Love
1996
2­
Steele
1997
2­
Steele
1997
2­
Wiley
1976
2­
Eschmeyer
and
Herald
1983
3­
Behrents
1983
3­
Behrents
1983
3­
Behrents
1983
3­
Ahlstrom
et
al
1984
4­
Fitch
and
Lavenberg
1975
4­
Waples
and
Rosenblatt1987
4­
Moser
1996a
5­
C.
D.
F.
G.
1982
5­
Moyle
1976
6­
Love
1996
7­
White
1977
8­
Fitch
and
Lavenberg
1975
9­
Wiley
1976
10­
Turner
et
al
1969
DISPERSAL
&
RECRUITMENT
female
deposits
eggs
in
a
cleaned
out
depression
or
nest
8
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31
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BM
BN
BO
BP
BQ
BR
BS
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
NF
1
2.0mm
­?
planktonic
1
planktivore
NF
NF
NF
33.0­
11.0
mm
inshore5
NF
1
4mm­?
planktonic
1
planktivore
NF
NF
NF
33.0­
11.0
mm
NF
1
3.0
mm­
?
planktonic
1
planktivore
NF
NF
NF
2
mean:
3.5
mm
pelagic
in
bays
2,3
(
zooplankton)
1
shallow
embayments
and
estuaries4
1­
Moser
1996a
1­
Moser
1996a
1­
Walker
1961
2­
White
1977
2­
White
1977
3­
Behrents
1983
3­
Fitch
and
Lavenberg
1975
4­
Yoklavich
et
al
1992
5­
Waples
and
Rosenblatt
1987
Food
habits
Larvae
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BT
BU
BV
BW
BX
BY
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
high
in
tidal
zone
1
NF
NF
NF
(
Small
juveniles:
harpacticoids
and
punkyfly
larvae)
2
1­
Macdonald
1975
1­
Moyle
1996
2­
Yoklavich
et
al
1992
2­
Walker
1961
tidal
creeks
and
inland
slough
regions2
Food
habits
Life
Stage
(
Large
juveniles:
free­
living
nematodes,
brine
fly
larvae,
and
aquatic
Hemiptera)
2
(
Large
juveniles:
brinefly
larvae,
waterboatmen,

pileworms)
1
(
Small
juveniles:
copepods,
punkyfly
larvae,

freeliving
larvae)
1
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Juveniles
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BZ
CA
CB
CC
CD
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
omnivores
(
plankton,
small
crustaceans)
1
forages
from
exposed
ledges
1
diurnal
feeders
1
reef
dwelling
predators,
such
(
zooplankton)
2
crepuscular
feeders2
as
kelp
bass
1
fish,
invertebrates,
and
birds4
residents
in
reef
areas12
perch
horizontally13
microcarnivores
3
usu.
motionless,
feeding
involves
diurnal
feeders
1
fish,
invertebrates,
and
birds4
(
benthic
copepods,
isopods,
gammarid
and
swift
turn
of
head
and
a
rapid
nip
at
caprellid
amphipods)
1
the
substratum
1
(
crustaceans,
benthic
copepods,
isopods,
gammarid
picks
prey
from
the
benthic
turf2
amphipods,
ostracods)
3
residents
in
reef
areas12
high
in
tidal
zone
3
carnivore
NF
terns
2
(
ghost
shrimp,
yellow
shore
crabs)
4
(
invertebrates,
small
fish)
5
hide
in
holes
at
low
tide
6
(
polychaete
worms,
tiny
mollusks,
and
other
worms)
7
California
halibut
and
aquatic
birds
California
gull
chicks6
Caspian
terns
and
Fosters'
terns7
(
gammarid
amphipods
and
polychaete
worms)
10
1­
Eschmeyer
and
Herald
1983
1­
Feder
et
al
1974
1­
Hartney
1989
1­
Hartney
1989
1­
Steele
1997
2­
Feder
et
al
1974
2­
Love
1996
2­
Behrents
1983
2­
Behrents
1983
2­
Love
1996
3­
Macdonald
1975
3­
Hartney
1989
3­
Wiley
1976
3­
Walker
1961
3­
Haaker
1975
4­
Moyle
1976
4­
Macdonald
1975
4­
Behrents
1983
5­
Barlow
1963
5­
Moyle
1976
5­
C.
D.
F.
G.
1982
6­
Love
1996
6­
Behrents
1983
6­
Dierks
1990
7­
Behrents
1983
7­
Fitch
and
Lavenberg
1975
7­
Harvey
1982
8­
Fitch
and
Lavenberg
1975
8­
Wiley
1976
9­
Wiley
1976
9­
Walker
1961
10­
Wiley
1976
10­
Barry
et
al
1996
larger
fish,
such
as
California
halibut3
lives
under
rocks,
in
caves,
or
far
back
in
crevices12,15
swims
out
from
reef
and
grabs
food
item
in
midwater3
takes
most
of
food
from
within
the
home
shelter
or
directly
in
front
of
diurnal
feeding
heavier
than
3
rocky
areas
with
heavy
encrustation
of
shelled
animal
life,
usually
inside
crevices,
more
common
around
slands
than
off
mainland1,2,15
microcarnivores
(
crustaceans,
planktonic
copepods,

cyprids,
copepod
nauplii)
3
(
copepods,
amphipods,
ostracods,
isopods,

euphausiids,
decapods,
eggs,
gastropods,

pelecypods,
porifera,
bryozoa,
coelenterates)
8
tidal
flats,
bays,
costal
sloughs;
prefers
mud
bottom
in
shallow
water1,4,5,8,11,14
(
benthic
copepods,
isopods,
gammarid,
ostracods,

and
caprellid
amphipods)
6
found
oriented
vertically
or
upside­
down
on
sides
or
underhangings
of
crevices13
Paralabrax
clauthratus,

Paralabrax
nebulifer,

Heterostichus
rostratus,

Paralichthys
californicus,

Hyperprosopon
argenteum,
and
Scorpaena
guttata
4
Paralabrax
clauthratus,

Paralabrax
nebulifer,

Heterostichus
rostratus,

Paralichthys
californicus,

Hyperprosopon
argenteum,
and
Scorpaena
guttata
4
(
copepods,
amphipods,
ostracods,
isopods,

decapods,
pelecypods,
gastropods,
eggs,
and
coelenterates)
8
Food
habits
lives
crevices
on
the
sides
or
upper
surfaces
of
reefs10
feeds
on
zooplankton
in
the
water
column2
open
rocky
areas,
retreats
to
crevices
and
holes,
or
hides
in
spines
of
sea
urchins
when
threatened
1,2,7,9
(
planktonic
copepods,
cyprids,
early
copepod
larvae,

benthic
copepods,
isopods,
and
gammarid
amphipods)
6
(
pile
worms,
barnacles,
Desert
pupfish,
and
young
mudsuckers)
9
capable
of
aerial
respiration
and
can
stay
alive
out
of
water
for
several
days14
Adults
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31
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CE
CF
CG
CH
CI
CJ
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
5
1­
Waples
and
Rosenblatt
1987
1
No
substocks
found
between
La
Jolla,
CA,
Channel
Islands,
CA,

Punta
Eugenia,
CA,
and
Isla
de
Guadalupe,
CA
using
allozymes
Mean
heterozygo
Nucleotide
diversity
(#)
GENETICS
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CK
CL
CM
CN
CO
CP
CQ
CR
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Total
Length
(
age)
Fishery
/
Total
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
0.087
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
osity
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22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CS
CT
CU
CV
CW
CX
CY
CZ
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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30
31
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
DA
DB
DC
DD
DE
DF
Interactions
/
associations
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Lythrypnus
zebra
1
recruitment
is
enhanced
by
presence
of
older
NF
NF
NF
NF
resident
conspecifics,
recruitment
is
halved
by
the
presence
of
predators
1
parasites:
2
families,
2
spp.
(
see
reference)
5
Lythrpnus
dalli
1
NF
NF
NF
NF
NF
associated
with
Clevandia
ios
4
NF
NF
NF
NF
parasites:
15
families,
20
spp.
(
see
reference)
5
1­
Behrents
1983
1­
Steele
1997
2­
Behrents
1983
3­
Wiley
1976
4­
Allen,
L.
G.
1982
5­
Love
and
Moser
1983
parasites:
acanthocephalans,
trematodes,

cestodes,
nematodes2,3
parasites:
acanthocephalans,
trematodes,

cestodes,
nematodes2,3
BEHAVIOR
Movement
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31
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Territoriality
Yes1,2,5,6,7
Yes5,6,7
Yes
3,4
1­
Feder
et
al
1974
2­
Eschmeyer
and
Herald
1983
3­
Moyle
1976
4­
Love
1996
5­
Behrents
1983
6­
Wiley
1976
7­
Hartney
1989
Page
461
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66
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68
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70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
A
B
C
D
E
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98
99
100
101
102
103
104
105
106
107
108
F
G
H
I
J
K
L
M
N
O
Page
463
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64
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66
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68
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71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
P
Q
R
S
T
U
V
W
X
Y
Page
464
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1167
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2220e
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Gobiidae
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55
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
Z
AA
AB
AC
AD
AE
AF
Page
465
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Gobiidae
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55
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58
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62
63
64
65
66
67
68
69
70
71
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73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
AG
AH
AI
AJ
AK
AL
AM
AN
Page
466
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5­
2220e
­
Gobiidae
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
Page
467
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1167
5­
2220e
­
Gobiidae
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
BC
BD
BE
BF
Page
468
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Gobiidae
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55
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
BG
BH
BI
BJ
BK
BL
11­
Limbaugh
1955
Page
469
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1167
5­
2220e
­
Gobiidae
54
55
56
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
BM
BN
BO
BP
BQ
BR
BS
Page
470
of
1167
5­
2220e
­
Gobiidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
BT
BU
BV
BW
BX
BY
Page
471
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1167
5­
2220e
­
Gobiidae
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
BZ
CA
CB
CC
CD
11­
Yoklavich
et
al
1992
12­
Turner
et
al
1969
13­
Hartney
1989
14­
C.
D.
F.
G.
1982
15­
Limbaugh
1955
Page
472
of
1167
5­
2220e
­
Gobiidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
CE
CF
CG
CH
CI
CJ
Page
473
of
1167
5­
2220e
­
Gobiidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
CK
CL
CM
CN
CO
CP
CQ
CR
Page
474
of
1167
5­
2220e
­
Gobiidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
CS
CT
CU
CV
CW
CX
CY
CZ
Page
475
of
1167
5­
2220e
­
Gobiidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
DA
DB
DC
DD
DE
DF
Page
476
of
1167
5­
2220e
­
Gobiidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Page
477
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
A
B
C
D
E
Page
478
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1167
5­
2220e
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Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
F
G
H
I
J
K
L
M
N
O
Page
479
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
P
Q
R
S
T
U
V
W
X
Y
Page
480
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
Z
AA
AB
AC
AD
AE
AF
Page
481
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
AG
AH
AI
AJ
AK
AL
AM
AN
Page
482
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
Page
483
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
BC
BD
BE
BF
Page
484
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
BG
BH
BI
BJ
BK
BL
Page
485
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
BM
BN
BO
BP
BQ
BR
BS
Page
486
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
BT
BU
BV
BW
BX
BY
Page
487
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
BZ
CA
CB
CC
CD
Page
488
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
CE
CF
CG
CH
CI
CJ
Page
489
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
CK
CL
CM
CN
CO
CP
CQ
CR
Page
490
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
CS
CT
CU
CV
CW
CX
CY
CZ
Page
491
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
DA
DB
DC
DD
DE
DF
Page
492
of
1167
5­
2220e
­
Gobiidae
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Page
493
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5­
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Gobiidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
A
B
C
D
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Haemulidae
1,2
Anisotremus
davidsoni
1,2
sargo
1
Gulf
of
California
(
isolated
population),
and
from
Magdalena
Bay,

Baja
California
to
Santa
Cruz,
California
1
(
Anisotremus
davidsonii
2
)
perch,
blue
bass,
black
croaker,
china
croaker,
California
sargo,
sargo
raiado
1
Haemulidae
1,2
Xenistius
californiensis
1,2,3,4,5
salema
1
Peru
to
Monterey
Bay,
including
Gulf
of
California
1
bigeye
bass,
California
salema,
striped
bass
1
1
­
Robins
et
al.
1991
1
­
Robins
et
al.
1991
1
­
Gates
and
Frey
1974
1
­
Miller
and
Lea
1972
2
­
Eschmeyer
1998
2
­
Eschmeyer
1998
3
­
Eschmeyer
and
Herald
1983
4
­
Allen
and
Robertson
1994
5
­
McKay
and
Schneider
1995
TAXONOMY
RANGE
Page
494
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1167
5­
2220e
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Haemulidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
E
F
G
H
I
J
K
AGE
a
Length
(
mm)
to
Max.
recorded
Geographic
Area
Linf
in
mm
(
SE
Observed
depth
range
(
m)
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
1
surface
to
42.65
m
(
130
ft)
NF
NF
1
44.2
cm
(
17.4
in)
to
58.4
cm
(
23
in)
NF
NF
NF
2
intertidal
to
60
m
(
198
ft)
2,3
58
cm
(
23
in)

1
1.31­
11.48
m
(
4­
35
ft)
NF
NF
1
25.4
cm
(
10
in)
NF
NF
NF
4
30.3
cm
(
11.8
in)

1
­
Miller
and
Lea
1972
1
­
Miller
and
Lea
1972
2
­
Love
et
al.
2000
2
­
Love
et
al.
2000
3
­
McKay
and
Schneider
1995
4
­
Love
et
al.
2000
Page
495
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Haemulidae
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
L
M
N
O
P
Q
R
S
T
U
V
W
X
nd
GROWTH
E)
k
(
SE)
to
(
SE)

Both
Male
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
r2
Growth
Parameters
Page
496
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
Y
Z
AA
AB
AC
AD
AE
AF
Age
composition
Other
refs.
Longevity
(
Length)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
NF
15
yr
(<
43
cm)
1
NF
NF
NF
Oviparous
1
NF
NF
12
yr
(
33
cm)
2
NF
NF
NF
NF
Oviparous
1
NF
NF
1
­
Fitch
and
Lavenberg
1971
1
­
Watson
1996b
2
­
Love
1996
Breeding
Behavior
Sexual
Page
497
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2220e
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Haemulidae
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
AG
AH
AI
AJ
AK
AL
AM
AN
AO
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.

NF
17.8
cm
(
2yr)
1
17.8
cm
(
2yr)
1
NG
1
NG
1
NG
1
NG
1
NG
1
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1
­
Feder
et
al.
1974
1
­
Feder
et
al.
1974
1
­
Feder
et
al.
1974
1
­
Feder
et
al.
1974
1
­
Feder
et
al.
1974
1
­
Feder
et
al.
1974
1
­
Feder
et
al.
1974
50%
maturity
dimorphism
1st
maturity
Total
length
in
cm
(
age
in
yrs)
at
REPRODUCTION
Page
498
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
states
present
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
Page
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Haemulidae
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
BD
BE
BF
BG
BH
BI
BJ
BK
Fecundity
Mean
Fecundity
Size
specific
eggs/
g
Number
of
eggs
fecundity
eq
Size
at
settlement
Size
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)

NF
NF
NF
NF
NF
NF
NF
2
0.76
­
0.88
mm
4
NG
NF
NF
NF
NF
NF
NF
NF
2
NG
3
NG
4
NG
2
­
Watson
1996b
3
­
Fitch
and
Lavenberg
1975
4
­
Feder
et
al.
1974
DISPERSAL
&
RECRUITMENT
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32
33
34
35
36
37
38
39
40
41
42
43
44
BL
BM
BN
BO
BP
BQ
Habitat
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
planktonic
2
NF
1
2.2
­
12.6
mm
planktonic
1
1
NG
NF
pelagic
4
2
1.7
mm
NL
(
hatching)
to
22
mm
SL
(
transformation)
planktonic
2
2
NG
planktonic
2
NF
1
2.7
­
15.5
mm
planktonic
1
1
NG
NF
pelagic
3
2
<
2.2
mm
NL
(
hatching)
to
21
mm
SL
(
transformation)
planktonic
2
2
NG
pelagic
4
1
­
Watson
and
Walker
1992
1
­
Watson
and
Walker
1992
2
­
Watson
1996b
2
­
Watson
1996b
2
­
Watson
1996b
3
­
Fitch
and
Lavenberg
1975
4
­
Feder
et
al.
1974
Food
habits
Eggs
Larvae
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30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
BR
BS
BT
BU
When
/
Frequency
Predators
Fish
Size
Habitat
NF
NF
NG
(
juv
or
adult?)
1
rocky
reef
(
13
m),
Pendleton
Artificial
Reef
1
NG
(
juv
or
adult?)
2
kelp
beds,
shallow
rocky
or
rock/
sand
combo
2
NG
(
juvs
&
few
adults)
3
surfgrass
(
Phyllospadix
)
beds
3
100
­
314
mm
SL
(
juv
or
adult?)
4
rock­
and­
sand
4
NG
(
juv
or
adult?)
7
rock
or
rock­
sand
bottom
7
NF
NF
NG
(
juv
or
adult?)
1
cobble
bottom
kelp
forest
(
8­
18
m),
San
Onofre
Kelp
bed
1
NG
(
juv
&
adult)
3
surfgrass
(
Phyllospadix
)
beds
3
NG
(
juv
or
adult?)
4
kelp
bed
w/
mixed
sand­
and­
rock
substrate
4
163­
180
mm
SL
(
juv
or
adult?)
5
night:
upper
water
column
(
5­
15
m)
over
open
fields
of
Dictyopteris
(
brown
algae),
Santa
Catalina
Island
5
163­
170
mm
SL
(
juv
or
adult?)
6
night:
nearshore
water
column,
Santa
Catalina
Island
6
NG
(
juv
or
adult?)
7
over
&
among
rocks
in
shallow
water
from
1.3­
11.5
m
(
4­
35
ft);
often
in
kelp
beds
near
canopy
7
1
­
DeMartini
et
al.
1989
1
­
DeMartini
et
al.
1989
2
­
Fitch
and
Lavenberg
1971
2
­
Fitch
and
Lavenberg
1971
3
­
DeMartini
1981
3
­
DeMartini
1981
4
­
Quast
1968d
4
­
Quast
1968d
5
­
Hobson
and
Chess
1976
5
­
Hobson
and
Chess
1976
6
­
Hobson
et
al.
1981
6
­
Hobson
et
al.
1981
7
­
Feder
et
al.
1974
7
­
Feder
et
al.
1974
HABITAT
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31
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33
34
35
36
37
38
39
40
41
42
43
44
BV
BW
BX
BY
BZ
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Fish
Size
NG
1
NF
NF
least
terns
2
NG
(
juv
or
adult?)
1
crustaceans
(
amphipods,
isopods,
shrimp,
crabs,
barnacles),
mollusks
(
kelp
scallops,
clam
siphons,
small
snails),
polychaete
worms
2
NG
(
juv
or
adult?)
2
NG
3
NG
(
juvs
&
few
adults)
3
small
isopods,
amphipods,
shrimp,
kelp
scallops
(
Pecten
latiauratus
)
4
100
­
314
mm
SL
(
juv
or
adult?)
4
bryozoans,
small
crustaceans,
mollusks
(
chitons,
clams)
7
NG
1
NF
yellowtail
and
kelp
bass
1
NG
(
juv
or
adult?)
1
NG
3
NG
(
juv
&
adult)
3
ophiuroids,
perciform
fishes,
shrimp,
amphipods,
isopod
(
Idothea
resecata)
,
other
isopods,
copepods,
and
questionables
(
mysids,

polychaetes)
4
NG
(
juv
or
adult?)
4
gammaridean
amphipods,
mysids,
swimming
polychaetes,
cumaceans,

caprellid
amphipods,
ostracods,
nebaliaceans,
isopods,
fishes,
caridean
larvae,
caridean
adults
and
juvs,
reptantian
zoea,
calanoid
&
cyclopoid
copepods
5
nocturnal
5
163­
180
mm
SL
(
juv
or
adult?)
5
gammarids
(
Batea
transversa,
Ampelisca
cristata
),
caprelled
(
Caprella
pilidigita
),
cumacean
(
Cyclaspis
nubila
),
mysid
(
Siriella
pacifica),
epitokous
nereid
polychaetes
6
nocturnal
6
163­
170
mm
SL
(
juv
or
adult?)
6
amphipod
crustaceans
7
1
­
DeMartini
et
al.
1989
1
­
Sikkel
1986
1
­
DeMartini
et
al.
1989
2
­
Fitch
and
Lavenberg
1971
2
­
Love
1996
2
­
Fitch
and
Lavenberg
1971
3
­
DeMartini
1981
3
­
DeMartini
1981
4
­
Quast
1968d
4
­
Quast
1968d
5
­
Hobson
and
Chess
1976
5
­
Hobson
and
Chess
1976
6
­
Hobson
et
al.
1981
6
­
Hobson
et
al.
1981
7
­
Feder
et
al.
1974
Food
habits
Life
Stage
Juveniles
T
ASSOCIATION
&
TROPHIC
INTERACTIONS
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24
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
CA
CB
CC
CD
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
rocky
reef
(
13
m),
Pendleton
Artificial
Reef
1
NG
1
NF
NF
kelp
beds,
shallow
rocky
or
rock/
sand
combo
2
crustaceans
(
amphipods,
isopods,
shrimp,
crabs,
barnacles),
mollusks
(
kelp
scallops,
clam
siphons,
small
snails),
polychaete
worms
2
surfgrass
(
Phyllospadix
)
beds
3
NG3
rock­
and­
sand
4
small
isopods,
amphipods,
shrimp,
kelp
scallops
(
Pecten
latiauratus
)
4
cobble
bottom
kelp
forest
(
8­
18
m),
San
Onofre
Kelp
bed
1
NG1
nocturnal
1
surfgrass
(
Phyllospadix
)
beds
3
NG3
kelp
bed
w/
mixed
sand­
and­
rock
substrate
4
ophiuroids,
perciform
fishes,
shrimp,
amphipods,
isopod
(
Idothea
resecata)
,
other
isopods,
copepods,
and
questionables
(
mysids,

polychaetes)
4
night:
upper
water
column
(
5­
15
m)
over
open
fields
of
Dictyopteris
(
brown
algae),
Santa
Catalina
Island
5
gammaridean
amphipods,
mysids,
swimming
polychaetes,
cumaceans,

caprellid
amphipods,
ostracods,
nebaliaceans,
isopods,
fishes,
caridean
larvae,
caridean
adults
and
juvs,
reptantian
zoea,
calanoid
&
cyclopoid
copepods
5
nocturnal
5
night:
nearshore
water
column,
Santa
Catalina
Island
6
gammarids
(
Batea
transversa,
Ampelisca
cristata
),
caprelled
(
Caprella
pilidigita
),
cumacean
(
Cyclaspis
nubila
),
mysid
(
Siriella
pacifica),
epitokous
nereid
polychaetes
6
nocturnal
6
1
­
DeMartini
et
al.
1989
1
­
DeMartini
et
al.
1989
2
­
Fitch
and
Lavenberg
1971
2
­
Fitch
and
Lavenberg
1971
3
­
DeMartini
1981
3
­
DeMartini
1981
4
­
Quast
1968d
4
­
Quast
1968d
5
­
Hobson
and
Chess
1976
5
­
Hobson
and
Chess
1976
6
­
Hobson
et
al.
1981
6
­
Hobson
et
al.
1981
Food
habits
Adults
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24
25
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
CE
CF
CG
CH
CI
CJ
CK
CL
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)

least
terns
2
NF
NF
NF
NF
NF
NF
NF
yellowtail
and
kelp
bass
1
NF
NF
NF
NF
NF
NF
NF
1
­
Sikkel
1986
2
­
Love
1996
Nucleotide
diversity
(#)
Mean
heterozygosity
GENETICS
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
CM
CN
CO
CP
CQ
CR
CS
CT
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Approach
Effective
population
size
Fishery
/
Length
(
age)
Fishery
/
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
CU
CV
CW
CX
CY
CZ
DA
DB
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
DC
DD
DE
DF
Interactions
/
associations
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

Parasites:
2
Families;
2
Species;
See
Ref
4
NF
NF
NF
Parasite:
trematode
(
Brachadena
pyriformis
)
in
stomach
4,7
Parasite:
trematode
(
Lasiotocus
longicaecum
)
in
intestine
4,8
Juveniles
exhibit
intraspecific
cleaning
behavior
1
NF
NF
NF
Parasites:
copepod
(
Peniculus
fissipes
)
imbedded
in
caudal
peduncle
1
Juveniles
school
with
sargo,
juv
black
croaker
2
Parasite:
helminth
(
Polynemicola
californica
)
from
gills
3
Parasite:
trematode
(
Lecithophyllum
intermedium
)
in
intestine
4,5
Parasite:
trematode
(
Opecoelus
xenistii
)
in
intestine
4,6
Parasite:
trematode
(
Helicometrina
nimia
)
in
intestine
4,5
1
­
Sikkel
1986
2
­
Fitch
and
Lavenberg
1975
3
­
Bravo­
Hollis
1985
4
­
Love
and
Moser
1983
5
­
Arai
1962
6
­
Manter
1940
7
­
Manter
and
van
Cleave
1951
8
­
Bravo­
Hollis
1956
Movemen
BEHAVIOR
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22
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24
25
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
DG
DH
DI
DJ
DK
DL
DM
DN
DO
DP
Schooling
Territoriality
Yes,
near
kelp
&
rocky
reefs
1
NF
Yes,
over
rocky
or
rock­
sand
bottom
from
surface
to
42.7
m
(
130
ft),

concentrated
2.6­
8.2
m
(
8­
25
ft)
6
Yes,
over
rocky
or
rocky
sand
areas,
usually
shallow
depths
7
Juveniles
school
with
other
species
in
inshore
rocky
areas
7
Yes,
near
kelp
&
rocky
reefs
1
NF
Yes
2,3
Yes,
during
daylight
>
50
individuals
at
middepths
within
kelp
forests
4
Yes,
during
day
in
upper
water
column
within
kelp
forests
5
Yes,
0.3­
1.3
m
(
1­
4
ft)
off
sandy
bottoms
shoreward
of
rocks
of
pilings
6
1
­
Watson
1996b
2
­
Sikkel
1986
3
­
Fitch
and
Lavenberg
1975
4
­
Hobson
and
Chess
1976
5
­
Hobson
et
al.
1981
6
­
Feder
et
al.
1974
7
­
CDFG
1982
nt
Page
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47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
A
B
C
D
Page
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52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
E
F
G
H
I
J
K
Page
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54
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56
57
58
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
L
M
N
O
P
Q
R
S
T
U
V
W
X
Page
512
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54
55
56
57
58
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
Y
Z
AA
AB
AC
AD
AE
AF
Page
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
AG
AH
AI
AJ
AK
AL
AM
AN
AO
Page
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81
82
83
84
85
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89
90
91
92
93
94
95
96
97
98
99
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Page
515
of
1167
5­
2220e
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Haemulidae
45
46
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50
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52
53
54
55
56
57
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59
60
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62
63
64
65
66
67
68
69
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71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
BD
BE
BF
BG
BH
BI
BJ
BK
Page
516
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
BL
BM
BN
BO
BP
BQ
Page
517
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
BR
BS
BT
BU
Page
518
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
BV
BW
BX
BY
BZ
Page
519
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
CA
CB
CC
CD
Page
520
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
CE
CF
CG
CH
CI
CJ
CK
CL
Page
521
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
CM
CN
CO
CP
CQ
CR
CS
CT
Page
522
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
CU
CV
CW
CX
CY
CZ
DA
DB
Page
523
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
DC
DD
DE
DF
Page
524
of
1167
5­
2220e
­
Haemulidae
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
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72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
DG
DH
DI
DJ
DK
DL
DM
DN
DO
DP
Page
525
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
A
B
C
D
Page
526
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
E
F
G
H
I
J
K
Page
527
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
L
M
N
O
P
Q
R
S
T
U
V
W
X
Page
528
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
Y
Z
AA
AB
AC
AD
AE
AF
Page
529
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
AG
AH
AI
AJ
AK
AL
AM
AN
AO
Page
530
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Page
531
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
BD
BE
BF
BG
BH
BI
BJ
BK
Page
532
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
BL
BM
BN
BO
BP
BQ
Page
533
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
BR
BS
BT
BU
Page
534
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
BV
BW
BX
BY
BZ
Page
535
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
CA
CB
CC
CD
Page
536
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
CE
CF
CG
CH
CI
CJ
CK
CL
Page
537
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
CM
CN
CO
CP
CQ
CR
CS
CT
Page
538
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
CU
CV
CW
CX
CY
CZ
DA
DB
Page
539
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
DC
DD
DE
DF
Page
540
of
1167
5­
2220e
­
Haemulidae
100
101
102
103
DG
DH
DI
DJ
DK
DL
DM
DN
DO
DP
Page
541
of
1167
5­
2220e
­
Haemulidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
A
B
C
D
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Hexagrammidae
Hexagrammos
decagrammus
1,2,3
kelp
greenling
1,2,3
Aleutian
Isl.,
Alaska
to
La
Jolla,
California
1
[
Calfornia
seatrout,
greenling,
seatrout,
rock
trout,

kelp,
kelp
greenling,
rockfish,
bluefish,
spotted
rock
trout,
bodieron,
boregat,
greenling,
kelp
trout,

kelp
cod]
1
Attu
Isl,
Aleutian
Isl.
(?)
to
La
Jolla,
California
2,3
Hexagrammidae
Hexagrammos
superciliosus
2
rock
greenling
1,2,3
Bering
Sea
to
Pt.
Conception
1
(
Hexagrammos
lagocephalus)
1,3,4
[
fringed
greenling,
Pacific
red
rock
trout,
red
greenling,
sea
trout,
kelp
cod]
1
Japan,
Yellow
Sea,
Okhotsk
Sea
to
western
Bering
Sea,

northwest
Alaska
Peninsula
(
Izembek
Bay)
and
Aleutian
Isl.
(
Attu
Isl)
to
Point
Conception,
California
2,3
TAXONOMY
RANGE
Page
542
of
1167
5­
2220e
­
Hexagrammidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
E
F
G
H
Length
(
mm)
to
Max.
recorded
Observed
depth
range
(
m)
weight
(
g)
equation
Length
conversion
equations
length
(
cm)

Intertidal
to
49
m
(
150
ft)
1
10
W
=
0.00000356
TL
3.221
2
FL
=
1.146(
SL)
+
3.076
SL
=
0.871(
FL)
­
2.379
(
range
130­
345
mm
SL;
n=
17)
3
53
cm
(
21
in)

Intertidal
(?)
to
158
m
(
522
ft)
2
11
W
=
0.00000499
SL
3.16368
2
TL
=
1.027(
FL)
­
3.112
FL
=
0.973(
TL)
+
3.092
(
range
155­
404
mm
FL;
n=
17)

14
Both
(
53­
240
mm
SL):
W
=
0.00000490
SL
3.27104
Male
(>
240
mm
SL):
W
=
0.000220
SL
2.60736
Female
(>
240
mm
SL):
W
=
0.0000278
SL
2.96606
2
SL
=
0.848(
TL)
+
0.327
TL
=
1.177(
SL)
+
0.062
(
range
156­
415
mm
TL;
n=
17)

3
TL
=
1.0
+
1.16
(
SL)

(
r
=
0.997)

Intertidal
and
shallow
rocky
areas
1
NF
2
FL
=
1.180(
SL)
­
1.100
SL
=
0.842(
FL)
+
2.295
(
range
174­
297
mm
SL;
n=
9)
1,3
61
cm
(
2
ft)

Intertidal
to
80
m
(
264
ft)
2
2
TL
=
1.016(
FL)
­
2.541
FL
=
0.984(
TL)
+
2.590
(
range
205­
342
mm
FL;
n=
9)

max
596
m
(
1967
ft)
3
2
SL
=
0.828(
TL)
+
4.620
TL
=
1.199(
SL)
­
3.483
(
range
203­
344
mm
TL;
n=
9)

E
Page
543
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1167
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Hexagrammidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
I
J
K
L
M
N
O
P
Q
AGE
and
GROWTH
Geographic
Area
Linf
in
mm
(
SE)
k
(
SE)
to
(
SE)

(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
Female
11
northern
Puget
Sound
(
TL)
NG
NG
398.64
(
NG)
NG
NG
0.8807
(
NG)
NG
NG
19
northern
Puget
Sound
(
SL)
420
(
2.186)
470
(
3.237)
460
(
1.501)
0.25
(
0.0066)
0.22
(
0.0074)
0.21
(
0.0015)
­
0.49
(
0.074)
­
0.08
(
0.09)

20
central
California
(
Carmel
Bay;
TL)
395
(
NG)
430
(
NG)
NG
0.288
(
NG)
0.201
(
NG)
NG
­
1.4
(
NG)
­
2.5
(
NG)

24
northern,
central,
and
south­
central
California
(
SL)
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
33
Diabo
Cove,
CA
No
No
No
No
No
No
No
No
Shikotan
Island?
(?
L)
30
NG
NG
NG
NG
NG
NG
NG
NG
Simushir
and
Shikotan
Islands
(?
L)
31
NG
NG
NG
NG
NG
NG
NG
NG
east
coast
of
Kamchatka
(?
L)
32
NG
NG
NG
NG
NG
NG
NG
NG
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544
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Hexagrammidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
R
S
T
U
V
W
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
0.9195
(
NG)
NG
NG
NG
11
otoliths/
whole
&
cleared
No
­
0.42
(
0.0063)
NG
NG
NG
19
otoliths/
whole
&
cleared
No
NG
NG
NG
NG
20
otoliths
/
NG
(
n
=
120)
No
see
curve
see
curve
see
curve
see
curve
24
otoliths
/
whole
&
cleared
(
n=
631)
No
No
No
No
No
33
NG
No
NG
NG
NG
NG
hypurals
/
NG
30
No
NG
NG
NG
NG
scales
and
hypurals
/
NG
31
No
NG
NG
NG
NG
hypurals
/
NG
32
No
Growth
Parameters
r2
Page
545
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Hexagrammidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
X
Y
Z
AA
AB
Age
composition
Verification
Other
refs.
Longevity
(
Length)
information
Sexuality
11
intra­
reader
NF
11
Female:
13+
yr
(
370­
379?
mm
SL)
3
Dioecious
No
12
Male:
8
yr
(
380
mm
TL)
Female:
9
yr
(
405
mmTL)

No
18
9
yr
(
360
mm
TL)

24
intra­
and
inter­
reader
No
No
NF
15
Female:
4+
(
473
mm
?
L)
NF
Dioecious
No
16
7+
(
41­
44
cm
?
L)

No
17
Female:
11+
(
57.0
cm
?
L)
Male:

8+
(
47
cm
?
L)
Page
546
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1167
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Hexagrammidae
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10
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12
13
14
15
16
17
18
19
20
21
22
23
24
AC
AD
AE
AF
AG
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
NF
Oviparous
1
male:
guard
>
1
egg
mass
at
a
time
10
NF
dichromatic
1
male:
guard
1­
10
egg
masses
(
or
clutches
per
nest)
at
a
time
11
Male:
dorsal
profile
from
dorsal
fin
to
snout
forms
sloping
straight
line;
Female:
dorsal
and
ventral
profiles
of
head
and
body
symmetrical
4
male
(
avg
=
31
cm
SL):
guard
nests
with
1­

11
clutches
(
avg
4);
several
mothers
14,15
Male:
Ventral
profile
of
head
and
body
nearly
horizontal
4
Male:
Brownish­
black
with
pale
blue
spots
on
anterior
portion,
particularily
head,
light
below;
Female:
color
brownish
to
grayish
with
numerous
fine
brown
spots
on
head
and
body
4
Male:
dark
gray
with
bright
blue
spots
on
head
and
body;
Female:
gray­
brown
with
golden­
brown
spots
on
body
and
head
5,6
Male:
prominent
narrow
white
stripes
and
spots
bordered
with
contrasting
black
dots
or
lines
superimposed
on
the
coloration
of
the
head,
nape,
anterior
half
of
the
back,

and
pectoral
fins;
Female:
speckled
on
sides
with
black
spots,
each
covering
only
a
f
dichromatic
(
detailed
description)
9
NF
Oviparous
1
males
guard
nests
4
NF
Male:
sides
marked
with
light
oval
blotches
(
eye
size)
which
sometimes
fuse
to
form
broad
light
vermiculate
patterns
7
Female:
some
females,
less
distinct
coloration
pattern
of
males
7
Male:
dark
cherry­
red;
Female:

brownishgreen
with
paler
yellowish
spots
on
lower
part
of
head
8
dichromatic
9,10,11
Breeding
Behavior
Sexual
dimorphism
Page
547
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Hexagrammidae
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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
AH
AI
AJ
AK
Male
Female
Male
Female
9
NG
9
31.6
cm
TL
(
NG)
9
29.5
cm
(
3­
4
yr)
9
29.5
cm
(
3­
4
yr)

10
NG
10
NG
10
33%
mature
at
age
2yr
10
67%
mature
at
age
3yr
15
29­
35
cm
?
L
(
3­
4
yr)
15
29­
35
cm
?
L
(
3­
4
yr)
15
29­
35
cm
?
L
(
3­
4
yr)
15
29­
35
cm
?
L
(
3­
4
yr)

Total
length
in
cm
(
age
in
yrs)
at
1st
maturity
50%
maturity
REPRODUCTION
Page
548
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67
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
AL
AM
AN
AO
AP
AQ
AR
Fertilized
e
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.

otoliths/
whole
&
cleared
9
No
intra­
and
inter­
reader
NF
British
Columbia
14
otoliths/
whole
&
cleared
10
No
No
British
Columbia
4
British
Columbia
18
northern
and
south­
central
California
17
hypurals
15
No
No
NF
east
coast
of
Kamchatka
16
Kuril­
Kamchatka
19
Aleutian
Islands
15
Fertilization
and
Spawning
Period
(^^^^^=

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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Fecundity
Mean
Fecundity
eggs
present
Spawning
females
present
Both
reproductive
states
present
eggs/
g
Number
of
eggs
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
(
weight
range)
(
at
cm,
grams,
or
age)

multibatch
spawner
(
at
least
3
clutches
per
spawning
season)

2,3
2
4335
±
623
eggs
per
female
NF
multiple
spawner
4
intermittent
(
4
sizes
egg
groups
in
ovaries)

5
NF
7
100,000
(
50
cm
?
L)

intermittent
(
large
number
of
egg
batches)
6
8
17,000
(
465
g)
­
26,330
(
1,330
g)

peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BF
BG
BH
BI
BJ
Size
specific
fecundity
eq
Size
at
settlement
(
length
or
weight
range)
Egg
duration
Larval
duration
&
location
Annual
cohort
success
NF
NF
1
yr
(
pelagic)
2
NF
NF
NF
2,10
30
d
(
6­
10oC)
NF
NF
NF
DISPERSAL
&
RECRUITMENT
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BK
BL
BM
Size
(
diameter
in
mm)
Habitat
Predators
1
NG
demersal,
attached
1
NF
9
2.2­
2.5
mm
(
avg
2.3
mm);

(
1580­
9660
eggs/
mass;
mean
4340eggs)
3­
D
biological
substrates:
encrusting
epifauna,
empty
barnacle
valves,
branches
of
hydrocoral,
rock
substrate
9
14
NG
masses
on
rocks
14
15
NG
masses
on
rocks
in
the
kelp
zone
(
Bamfield
Inlet)
15
16
2.2
­
2.5
mm
rock;
rock
crevices;
rock
and
biological
substrate;
biological
substrate
16
19
2.3
mm
5­
17
m
depth;
rock,
rock
crevices,
or
rock
&
biological
substrate
19
21
NG
benthic
on
algae
21
1
NG
demersal,
adhesive,
attached
1
NF
14
NG
on
rocks
or
algal
holdfasts
in
areas
of
strong
currents
14
20
3
mm
adhesive,
algae
or
stones
20
21
2.0
­
2.6
mm
(
avg
2.5
mm)
3­
10
m
depth,
adhesive,
on
algae
21
Eggs
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BN
BO
BP
BQ
BR
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
1
7­
9
mm
SL
at
hatching
epipelagic
to
50
mm
SL
1
NG
1
NF
NF
2
19­
56
mm
FL
pelagic
2,9
copepods,
amphipods,
fish
(
Theragra
chalcogrammus
),

Brachyura
larvae,
eggs,
Oikopleura,
Bryozoa
larvae
2
3
15.5
­
53
mm
FL
pelagic,
surface
3
copepods,
amphipods,
euphausiids,
fish,
eggs
3
7
52
­
69
mm
FL
surface
waters
Strait
of
Georgia,
B.
C.
7
carnivore:
copepods,
amphipods,
euphausiids,
ostracods,

Cladocera,
Cirrepedia,
isopods,
fish,
decapods
(
see
ref
for
ID)
7
10
7
­
8
mm
TL
(
hatching,
December)
NG
10
13
8.6
­
60
mm
FL
(
larvae
and
juv)
surface
waters,
Strait
of
Georgia
13
copepods
(
Pseudocalanus
minutus,
Calanus
sp
,
nauplii,

Microcalanus
sp,
Metridia
sp.,
fragments),
amphipods
(
Parathemisto
pacifica,
Hyperoche
sp.,
Scina
borealis
),

larvacea
(
Oikopleura
sp.),
euphausiid
(
Thysanoessa
longipes
,

Calyptopsis
larva),
decapod
(
Br
15
8
mm
SL
at
hatching
pelagic
(
up
to
500­
600
miles
offshore)
15
NG
15
7­
9
mm
SL
at
hatching
1
pelagic
1
NG
1
NF
NF
9
mm
SL
at
hatching
15
pelagic
15
NG
15
Food
habits
Larvae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BS
BT
BU
BV
Predators
Fish
Size
Habitat
Foraging
guild
NF
3
NG
(
juvenile
or
adult)
rocky
nearshore
3
holothurians,
crabs,
shrimp,
chitons,
snails
3
7
NG
(
juvenile
or
adult)
bedrock
ledges
or
boulders
>
10
m
7
NG
7
9
53­
56
mm
FL
pelagic
surface,
Strait
of
Georgia,
B.
C.
9
copepods
9
11
52
­
69
mm
FL
surface
waters,
Strait
of
Georgia,
B.
C.
11
carnivore:
copepods,
amphipods,
euphausiids,
ostracods,

Cladocera,
Cirrepedia,
isopods,
fish,
decapods
(
see
ref
for
ID)
11
12
NG
(
juvenile
&
adult)
kelp
bed,
NW
Carmel
River
mouth,
CA
12
crabs
(
Pugettia
producta
,
P.
gracilis
,
P.
richi
),
shrimp
(
Crago
sp,

Cragon
sp,
Spirontocaris
sp),
fish
eggs,
gastropods,

Polyphacophora
(
Tonicella
lineata
),
octopus,
polychaetes,
algae
12
15
8.6
­
60
mm
FL
(
larvae
and
juv)
surface
waters,
Strait
of
Georgia
15
copepods
(
Pseudocalanus
minutus,
Calanus
sp
,
nauplii,

Microcalanus
sp,
Metridia
sp.,
fragments),
amphipods
(
Parathemisto
pacifica,
Hyperoche
sp.,
Scina
borealis
),
larvacea
(
Oikopleura
sp.),
euphausiid
(
Thysanoessa
longipes
,
Calyptopsis
larva),
decapod
(
Br
20
59
­
392
mm
TL
(
juv
&
adult)
Diablo
Cove,
CA
20
crustaceans,
Octopus
sp,
brittle
stars,
fish,
fish
eggs,
urchin
eggs,

chitons,
polychaetes,
algae,
snails,
abalones
20
NF
14
66
mm
FL
surface
waters,
Saanich
Inlet,
Vancouver
Isl.
14
Euphausiid
(
Euphausia
pacifica)
14
22
NG
benthic,
Northern
Kuril
Islands
&
Southeastern
Kamchatka
22
molluscs
(
octopus),
amphipods,
isopods,
mysids,
squid,
shrimps,

crabs,
worms,
snails,
clams,
other
22
Life
Stage
Food
habits
Juveniles
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
BW
BX
BY
BZ
CA
Foraging
method
When
/
Frequency
Predators
Fish
Size
Habitat
NF
NF
NF
11
NG
(
juvenile
or
adult)
bedrock
ledges
or
boulders
>
10
m
11
8
NG
Nanaimo,
B.
C.
8
13
24.4
cm
SL
90­
95
m,
Soquel
Canyon,
Monterey
Bay,
CA
13
14
NG
NG
14
20
59
­
392
mm
TL
(
juv
&
adult)
Diablo
Cove,
CA
20
NF
NF
NF
15
NG
benthic,
Kamchatka
15
16
NG
Shikotan
Island
16
17
NG
Simushir
&
Shikotan
Islands
17
22
NG
benthic,
Northern
Kuril
Islands
&
Southeastern
Kamchatka
22
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
CB
CC
CD
CE
CF
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
11
NG
NF
NF
NF
NF
carnivore:
fish
(
herring
eggs,
Anoplarchus
atropurpureus
),

crabs
8
Octopus
sp.
beaks,
crab
remains,
whole
brown
catshark
egg
case
(
Apristurus
brunneus
)
13
generalist
14
crustaceans,
Octopus
sp,
brittle
stars,
fish,
fish
eggs,
urchin
eggs,
chitons,
polychaetes,
algae,
snails,
abalones
20
benthophagous­
polyphagous:
polychaetes,
isopods,

amphipods,
decapods,
fish
eggs
15
NF
intensive
in
morning
&
evening,
also
high
tide
at
shoreline
2
NF
NF
fish
(
Hypomesus),
Pandalus
sp,
small
crabs,
hermit
crabs,

small
shrimp,
amphipods,
mysids,
polychaetes,
Nereis
sp,

Zostera,
Laminaria,
red
algae
16
polyphage
(
wide
range):
amphipods,
isopods,
polychaetes,

molluscs,
algae,
fish
eggs
17
molluscs
(
octopus),
amphipods,
isopods,
mysids,
squid,

shrimps,
crabs,
worms,
snails,
clams,
other
22
Adults
Food
habits
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
CG
CH
CI
CJ
CK
CL
CM
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Mean
heterozygosity
Nucleotide
diversity
(#)
GENETICS
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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
CN
CO
CP
CQ
CR
CS
CT
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Effective
population
size
Fishery
/
Length
(
age)
Fishery
/
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
NF
Sport
/
319.5
mm
SL
(
5
yr)
2
Sport
/
363
mm
SL
(
7
yr)
2
hook
&
line
2
otoliths/
whole
&
cleared
2
NG
2
NG
2
NF
NF
NF
NF
NF
NF
NF
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
CU
CV
CW
Mortality
Estimates
Natural
(
M)
Fishing
(
F)
Total
(
Z)

8
NG
8
NG
8
0.3386
(
Sexes
combined)

NF
NF
NF
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10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
CX
CY
CZ
Rate
of
Ex
Method
of
Age
range
used
Rate
of
mortality
estimation
to
calculate
mortality
exploitation
(
u)

age­
frequency
analysis
&
otolith
age
data
8
8
5­
14
yr
8
NG
NF
NF
NF
MORTALITY
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
DA
DB
DC
xploitation
Exploitation
Interactions
/
associations
ratio
(
F/
Z)
Competition
with
other
organisms
8
NG
NF
Parasite:
copepod
(
Chondracanthus
deltoideus
)
in
gill
cavity
6
Parasite:
copepod
(
Chondracanthus
pinguis
)
6
Parasites:
9
Families;
~
12
Species;
see
Refs
17
Parasite:
copepod
(
Haemobaphes
sp
)
11
NF
NF
Parasite:
endoparasitic
worms
13
Parasite:
mongenetic
fluke
(
Microcotyle
gotoi
)
on
gills
14
Parasites:
18
Families;
~
36
Species;
see
Ref
17
Parasites:
Digenea
(
6
sp),
Monogenea
(
1
sp),
Cestoda
(
2
sp),
Nematoda
(
3
sp)
11
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
DD
DE
DF
DG
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
NF
NF
NF
NF
NF
NF
NF
NF
Movement
BEHAVIOR
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
DH
Territoriality
Male:
interspecifically
and
intraspecifically
of
nests
1,2
Male:
during
spawning
season
5
NF
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25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
A
B
C
D
Hexagrammidae
Ophiodon
elongatus
1,
2,3
lingcod
1,2,3
Kodiak
Isl.,
Alaska
to
Pt.
San
Carlos,
central
Baja
California
1
[
cultus,
skilfish,
buffalo]
1
Shumagin
Isl.,
southwestern
Gulf
of
Alaska
to
Pt.
San
Carlos,

central
Baja
California
3
[
Pacific
cultus,
cultus
cod,
ling]
1,5
center
of
abundance
off
coast
of
British
Columbia
4
[
bluefish,
cod,
card,
testoni]
1,6
[
leopard
cod,
codfish,
blue
cod,
green
cod,

bocalao,
greenling,
white
cod]
1,5,6
[
buffalo
cod]
5,6
[
ling,
dragonfish,
greenlinger]
4
[
steamer
cod]
6
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26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
E
F
G
H
post
larvae
to
7.6
cm
(
3
in)
pelagic
nearshore
and
offshore
1
1
curves
drawn,
no
equations
1
FL
=
0.981(
TL)
­
0.521
TL
=
1.019(
FL)
+
0.562
(
range
23­
1046
mm;
n=
619)
1,4
114.3
cm
TL
(
45
in)

Juvs
in
shallow
bays
and
on
sand
&
mud
bottoms
from
beach
area
to
several
hundred
ft
1
2
Ripening
Female:
W
=
3.277
x
10­
10
FL3.6558
(
W
=
lbs;
FL
=
76­
106
cm)
1
SL
=
0.873(
TL)
+
0.308
TL
=
1.145(
SL)
­
0.286
(
range
23­
933
mm;
n=
578)
2,3,5,6,8
152
cm
TL
(
5
ft)

Adults
from
surface
to
459
m
(
1400
ft)
1
3
Male
(
suspect):
W
=
0.00004905
TL3.24
Female
(
suspect):
W
=
0.00002299
TL
3.12
1
SL
=
0.889(
FL)
+
0.884
FL
=
1.124(
SL)
­
0.942
(
range
23­
933
mm;
n=
555)
5,7
Male:
92
cm
TL
(
36.25
in)

Barely
subtidal
to
491
m
(
1620
ft)
2
4
W
=
0.000000564
L
3.498
2
FL
=
1.135(
SL)
­
6.085
SL
=
0.880(
FL)
+
5.937
(
range
381­
852
mm
SL;
n=
25)
5
Female:
132
cm
TL
(
52
in)

4
3­
400
m
5
W
=
0.000007177
TL
3.0687
(
W
=
kg;
TL
=
cm)
2
TL
=
1.013(
FL)
+
4.090
FL
=
0.987(
TL)
­
3.942
(
range
430­
967
mm
FL;
n=
25)

6
W
=
0.01145
TL
2.99533
(
W
=
g;
TL
=
51­
101
cm)
2
SL
=
0.869(
TL)
+
2.407
TL
=
1.150(
SL)
­
2.146
(
range
440
­
983
mm
TL;
n=
25)

7
Male:
W
=
0.003953
TL
3.214900
Female:
W
=
0.001760
TL
3.397800
(
W
=
g;
TL
=
cm)
4
FL
=
­
1.8
+
1.258
DL
8,9
Male:
W
=
0.00000241
TL
3.2149
Female:
W
=
0.0000007042
TL
3.3978
5
TL
=
­
18
+
1.258
DL
(
length
mm;
550­
1070
mm
TL)

12
W
=
0.00000333
FL
3.097
6
TL
=
0.0558
+
1.2280
DL
(
60­
115
cm
TL)

13
Male:
W
=
0.003953
TL
3.2149
Female:
W
=
0.001760
TL
3.3978
15
W
=
0.000003488
FL
3.246
(
note:
W
=
kg
and
FL
=
cm)

16
W
=
0.00001050
FL
2.991
(
note:
W
=
kg
and
FL
=
cm)

17
SW
Vancouver
Island
Male:
W
=
­
13.86
FL
3.37
Female:
W
=
­
13.32
FL
3.27
17
Queen
Charlotte
Sound
Male:

W
=
­
13.29
FL
3.28
Female:
W
=
­
11.83
FL
3.05
Page
565
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
I
J
K
L
M
N
O
P
Q
1
SE
Alaska
(
FL)
No
No
No
No
No
No
No
No
2
Alaska
(
NG)
No
No
No
No
No
No
No
No
3
Vancouver
Island,
Strait
of
Georgia,
B.
C.
(
TL)
No
No
No
No
No
No
No
No
3
Vancouver
Island,
Strait
of
Georgia,
B.
C.
(
TL)
No
No
No
No
No
No
No
No
4
Vancouver
Island,
Strait
of
Georgia,
B.
C.
(
TL)
No
No
No
No
No
No
No
No
5
Strait
of
Georgia,
B.
C.
Canada
(
FL)
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
6
Strait
of
Georgia,
B.
C.
[
FL
(
95%
CI)]
770
(
39)
1040
(
40)
No
2.5
(
0.5)
1.8
(
0.3)
No
­
11.9
(
3.5)
­
11.1
(
2.3)

28
Strait
of
Georgia,
B.
C.
(
FL)
No
No
No
No
No
No
No
No
29
Strait
of
Georgia,
B.
C.
(
TL?,
cm)
91
115
No
0.26
0.16
No
­
0.6
­
1.4
6
west
coast
Vancouver
Island
[
FL
(
95%
CI)]
861
(
21)
1128
(
41)
No
2.7
(
0.3)
1.8
(
0.2)
No
­
8.0(
3.7)
­
9.0
(
3.4)

27
west
coast
Vancouver
Island
(
FL)
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
25
Southwest
Vancouver
Island
(
FL)
NG
NG
NG
NG
NG
NG
NG
NG
26
Gulf
of
Georgia
(?
L)
NG
NG
NG
NG
NG
NG
NG
NG
7
British
Columbia
No
No
No
No
No
No
No
No
Page
566
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
R
S
T
U
V
W
No
No
No
No
1
fin
rays/
thin
sections
&
otoliths/
whole
No
No
No
No
No
2
otoliths/
whole
&
cleared;
break
&
burn
No
No
No
No
No
3
none/
Length
Frequency;
age
0y,
1y
(
27cm
TL),
2y
(
47cm
TL)
No
No
No
No
No
3
Vertebrae/
Whole;
age
1y
(
35cm
TL),
2y
(
47cm
TL)
No
No
No
No
No
4
Vertebrae/
Whole;
growth
curves
plotted,
no
parameters
No
see
curve
see
curve
see
curve
see
curve
5
fin
rays/
thin
sections?
OTC:
Male
(
1­
11yr);
Female
(
2­
13
yr)

No
No
No
No
6
length­
frequency
&
length­
increment
data
(
tag­
recapture)
No
No
No
No
No
28
fin
rays/
sections
&
tag­
recapture
for
growth
rates
No
No
No
No
No
29
vertebrae/?
and
length­
frequency
No
No
No
No
No
6
length­
frequency
&
length­
increment
data
(
tag­
recapture)
No
see
curve
see
curve
see
curve
see
curve
27
fin
rays
/
sections
No
NG
NG
NG
NG
25
fin
rays/
cross
sections
Tagging
and
OTC
attempted
NG
NG
NG
NG
26
No
/
Tagging
studies
to
determine
growth
rates
No
No
No
No
No
7
unknown
unknown
Page
567
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
X
Y
Z
AA
AB
between
structures
1
1,2
Male:
14
yr
(
91
cm
TL);

Female:
16
yr
(
110
cm
TL)
1,2,4
Dioecious
intra­
reader
precision
3
Male:
12
yr
(
NG);
Female:

15
yr
(
NG)

No
4
16­
20
yr
(
40
lb)

No
5
Male:
11
yr
(
89
cm
TL);
Female:

12
yr
(
110
cm
TL)

No
6
Male:
11yr
(~
80
cm
FL);

Female:
13yr
(~
96
cm
FL)

unknown
7
Male:
14
yr
(~
85
cm
TL);

Female:
20
yr
(
123
cm
TL)

No
8
Male:
15
yr
(
812
mm
TL);

Female:
19
yr
(
1122
mm
TL)

No
9
Male
(
north):
16
yr
(
88
cm
TL);
Female
(
north):
20
yr
(
118
cm
TL);
Male
(
south):
15
yr
(
74
cm
TL);
Female
(
south):
19
yr
(
110
cm
TL)

No
10
Male:
13yr
(
855
mm
FL);
Female:
19yr
(
1150
mm
FL)

No
13
Male:
13
yr
(
75
cm
FL);
Female:
14
yr
(
111
cm
FL)

No
14
Male:
6
yr
(
68.5
cm
FL);
Female:
12
yr
(
105.7
cm
FL)

NG
No
unknown
Page
568
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
AC
AD
AE
AF
AG
Male
may
fertilize
ova
from
more
than
one
female
1,2,3
Oviparous
1
males
guard
nest
1,2,3,5,6,7,8,9,12,13
NF
Male:
large
dark
greenish
brown
areas
on
back
and
sides
outlined
with
tracings
of
pale
blue
or
orange;
Female:
similar
to
males
but
usually
lighter,
with
orange
tracings
rather
than
blue,
small
orange
spots
scattered
amonst
larger
areas
1
Males
monogamous
or
polygamous
4
external
fertilization
2
usually
one
nest
per
male
7
Male:
anal
papilla
2,3
males
guard
 
1
nest
3,8,9,13
Coloration
variable
(
detailed
description)
7
dichromatic
(
detailed
description)
9
Page
569
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
AH
AI
AJ
AK
1
46
cm
TL
(
2yr)
1
70
cm
TL
(
5yr)
1
NG
1
NG
2
66
cm
TL
(
3yr)
2
66
cm
TL
(
3yr)
2
NG
2
NG
3,6
58.4
cm
TL
(
3yr)
3,6
58.4
cm
TL
(
3yr)
3,6
NG
3,6
NG
4
50
cm
TL
(
2yr)
4
80
cm
TL
(
7yr)
4
NG
4
61­
75
cm
TL
(
3­
5
yr)

5
39
cm
TL
(
2yr)
5
51
cm
TL
(
3yr)
5
NG
(
2yr)
5
NG
(~
5yr)

7
52
cm
TL
(
NG)
7
77
cm
TL
(
NG)
7
NG
7
NG
8
39.0
cm
FL
(
2yr)
8
43.7
cm
FL
(
2yr)
8
46.1
cm
FL
(
3.2
yr)
8
55.7
cm
FL
(
3.8
yr)

11
NG
11
NG
11
NG
11
83
cm
TL
(
NG)

12
NG
12
NG
12
57.1
cm
TL
(
3.5
yr)
12
63.6
cm
TL
(
3.9
yr)

12
NG
12
NG
12
62.8
cm
TL
(
5
yr)
12
67.5
cm
TL
(
4.5
yr)

13
NG
13
NG
13
51.3
(
3.4
yr)
13
63.4
(
4.6
yr)

14
60
cm
FL
(
3
yr)
14
68
cm
FL
(
3
yr)
14
60
cm
FL
(
4
yr)
14
76
cm
FL
(
5
yr)

16
<
51
cm
FL
(
NG)
16
~
60
cm
FL
(
NG)
16
<
51
cm
FL
(
NG)
16
70
cm
FL
(
NG)

17
all
males
examined
were
mature
and
resting
17
SW
Vancouver
Island:
60
cm
FL
(
NG)

Queen
Charlotte
Sound:
63
cm
FL
(
NG)
17
all
males
examined
were
mature
and
resting
17
SW
Vancouver
Island:
65
cm
FL
(
NG)

Queen
Charlotte
Sound:
69
cm
FL
(
NG)

Page
570
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
AL
AM
AN
AO
AP
AQ
AR
NG
1
NG
1
NG
1
NF
southeast
Alaska
1
*****

British
Columbia
2
Canadian
waters
3
*****
*****

Departure
Bay,
B.
C.,
Canada
4
unknown:
(
otoliths
and/
or
catch
data)
5
No
5
No
5
Dodd
Narrows,
B.
C.,
Canada
5
*****
*****

northern
Puget
Sound
6
*****

northern
Puget
Sound
7
Washington
(
Shilshole
breakwater,
Seattle)
8
*****
*****

Pacific
Northwest
9
northern
and
central
California
10
See
reference
for
sea
central
California
(
Point
Reyes
&
Gulf
of
Farallones)
11
California
12
*****

none
No
No
Generic
13
*****
*****

Page
571
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
night
1
1
26
eggs/
g
1
500,000
(
118
cm
TL)

1,2
60,000
(
70
cm
TL)

2
518,000
(
104
cm
TL)

3
280,000
(
98
cm
TL)

3
436,000
(
112
cm
TL)

*****
4
97,000
(
70
cm
TL)

4
490,000
(
120
cm
TL)

*****
5,6
170,000
(
86
cm
TL)

5
476,000
(
118
cm
TL)

asonal
gonad
maturity
stage
*****
*****

Page
572
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
BF
BG
BH
BI
BJ
1
N
=
0.0002824
x
FL
3.001
(
N
=
#
eggs;
FL
=
mm)
1
unknown
1
yolk
sac
absorbed
in
10
days
80
mm
TL
(
3
mo),
shallow
(<
20
m),

nearshore
bottom,
sand
or
sparse
rock
1
NF
1
N
=
16210
W
0.8498
(
N
=
#
eggs;
W
=
lbs)
3
6
wk
3
1­
2
months
(
Strait
of
Georgia,

March­
May);
in
May,
pelagic
postlarvae
move
inshore
and
become
demersal
after
a
few
weeks
juveniles
(
3
mo)
settle
on
sandy
bottom
near
eelgrass
or
kelp
beds
2
1,2
N
=
0.0003026
x
TL
3.000
4
5­
11
wk
(
7
avg)

5
7
wk
6
>
5
wk,
estimated
7­
8
wk,
one
nest
11
wk
7
5­
7
wk
8
35
days
at
10
oC;
48
days
at
8
oC
9
21
days
Page
573
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
BK
BL
BM
1
2.24­
3.23
mm
demersal,
adhesive,
attached
1
crabs
(
Pagurus
sp)
3
2
2.8­
3.5
mm
attached
to
substrate
in
large
masses
below
low­
tide
mark
2
sharp­
nose
crab
(
Scyra
acutifrons)
3,12,16
3
3.5
mm
adhesive
masses
in
crevices,
caves,
under
or
between
boulders;
2.4­
19.8
m
below
MLLW
(
not
investigated
below
24.3
m)
3
crab
(
Phyllolithodes
papillosus
)
16
4
2.8­
3.5
mm
adhesive
and
stick
in
large
masses
to
rocky
crevasses
4
unid
shrimp
3
5
2.8
­
3.5
mm;
nest:
>
76
cm
diam,
5.6
kg
3­
10
m
below
low
tide,
crevices
in
boulder
area,
where
little
marine
growth
5
red
sea
urchin
(
Strongylocentrotus
franciscanus
)
3,6,10,12,16
6
2.8
­
3.5
mm
NG
6
purple
sea
urchin
(
Strongylocentrotus
purpuratus
)
3
7
nest:
5
Liters
rocky
crevices,
current­
swept
narrows,
shallow
water,

exposed
coastal
reefs;
­
1.8
to
­
21.9
m
MLLW
(
San
Juan
Islands,
WA);
+
1.0
to
­
10.4
m
MLLW,
cobble
&
oyster
shells
(
Dabob
Bay,
WA)
7
green
sea
urchin
(
Stongylocentrotus
drobachiensis
)
11,16
8
nests:
0.5­
2
liter
volume
33­
97
m;
in
vertical
cracks
between
boulders,
over
boulder
overhangs,
or
in
V­
shaped
cuts
within
lava
flows;
Sitka
Sound,
AK
8
gastropod
(
Amphissa
columbiana)
3,6,12,16
10
NG
crevices
or
rock
ledges
&
some
water
movement;
to
33
m
(
100
ft)
from
high
tide
level
10
gastropod
(
Calliostoma
ligatum
)
3,6,12,16
11
NG
1.6
­
8.2
m
of
the
plus
2
tide
level,
crevices
or
rock
ledges
11
sunflower
star
(
Pycnopoda
helianthoides
)
10
12
NG
(
nest
volume:
2­
3
liter)
nest
in
rocky
areas
with
high
current
velocites;
spawning
on
cobble­
oyster
shell
substrate
in
slow
current;
­
1.8
m
to
22
m
below
MLLW;
shallowest
in
slow
current
area
12
starfish
(
Pisaster
ochraceus
)
12
13
NG
nests
in
crevices,
beneath
overhanging
rock,
extend
lower
intertidal
to
18.9m
13
anemone
(
Tealia
crassicornis
)
10
16
2.8
mm;
nest
45.7
x
33.0
x
43.3
cm
adhesive,
on
jagged
rock
16
Scalyhead
sculpin
(
Artedius
harringtoni
)
3,16
17
2.8
mm
adhesive,
under
rock,
0.5
m
below
surface
(
intertidal)
17
Puget
Sound
rockfish
(
Sebastes
emphaeus
)
3,12
Page
574
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
BN
BO
BP
BQ
BR
1
7­
10
mm
SL
(
avg
9.3
mm
SL)
at
hatching
pelagic
1
NG
1
NF
NF
2
15­
51
mm
FL
pelagic
2
copepods,
eggs,
euphausiids,
phytoplankton
2
4
10
mm
TL
at
hatching
NG
4
NG
4
3
18­
49
mm
FL
pelagic,
surface
3
copepods,
amphipods,
Brachyura
larvae,
eggs,
euphausiids,
fish
larvae
3
5
7
mm
SL
at
hatching
NG
5
NG
5
6
6­
10
mm
TL
at
hatching
pelagic,
surface
6
NG
6
6
10­
20
mm
TL
pelagic,
surface
6
small
eggs,
copepod
nauplii,
copepodite,
small­
medium
calanoids,
larvacean
6
6
21­
30
mm
TL
pelagic,
surface
6
small/
med/
lg
calanoids,
small
eggs,
juv
euphausiid,
copepod
nauplii,
med
eggs,
copepodite
6
6
31­
50
mm
TL
pelagic,
surface
6
med/
sm/
lg
calanoids,
small
eggs,
decapod
larva,
harpacticoid,

amphipod,
herring
(
7­
15mm)
6
6
51­
70
mm
TL
pelagic,
surface
6
herring
(
7­
30
mm),
med/
lg/
sm
calanoids,
decapod
larva,
small
eggs,
harpacticoid,
amphipod
6
6
>
70
mm
TL
mainly
juv
herring;
lessor
(
lg/
med/
sm
calanoid,

decapod
larva)
6
NG
6
7
60
mm
FL
surface
waters
Strait
of
Georgia,
B.
C.
7
copepod
(
Calanus
pacificus
)
7
8
10­
20
mm
FL
pelagic,
surface
8
small
calanoids,
crustacean
eggs,
copepodites,
medium
calanoids,
larvacean,
copepod
nauplii
8
8
21­
30
mm
FL
pelagic,
surface
8
small
calanoids,
crustacean
eggs,
medium
calanoids,
juv
euphausiids
8
Page
575
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
BS
BT
BU
BV
coho
salmon
1
1
1
to
<
46
cm
TL
males
1
NG
piscivore
1
1
1
to
<
70
cm
TL
females
1
NG
piscivore
1
2
NG
(
small
juv)
British
Columbia
2
shrimp
(
Neomysis
macrops
),
prawn
(
Pandalus
danae
)
2
2
NG
(
large
juv)
British
Columbia
2
young
herring,
other
small
fishes
2
3
NG
(
juv
or
adult)
rocky
nearshore
3
fish,
prosobranch
gastropod
3
4
<
70
mm
TL
(
juv)
epipelagic
4
NG
4
2,4,5
>
70
mm
TL
(
juv)
various
(
eelgrass
beds,
bays,
estuaries)
2,4,5
NG
2,4,5
4
350
­
500
mm
TL
(
juv)
shallow
rocky
reefs
4
NG
4
6
>
70
mm
TL
demersal;
kelp
beds;
eelgrass
beds;
flat
bottom
6
juvenile
herring,
Pacific
sand
lance,
flatfish
(
Pleuronectidae),

shiner
perch,
juv
walleye
pollack;
shrimp,
prawns
6
6
NG
(
juv
age­
0)
shallow
water
near
kelp
&
eelgrass
beds
6
NG
6
6
NG
(
juv
age­
1)
nearshore
areas,
segregated
from
older
age
classes
6
NG
6
6
NG
(
juv
age­
2)
move
to
adult
habitat
6
NG
6
7
<
40
cm
FL
bedrock
ledges
or
boulders
>
10
m
7
NG
7
8
NG
(
juvenile)
NG
8
herring,
Pacific
sand
lance
8
Page
576
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
BW
BX
BY
BZ
CA
NF
NF
size
NG
(
juv
or
adult):
harbor
seal
(
Phoca
vitulina
richardsi
)
1
1
Male:
 
46
cm
TL;

Female:
 
70
cm
TL
1
NG
2
NG
(
adult)
British
Columbia
2
3
NG
(
juv
or
adult)
rocky
nearshore
3
4
NG
(
adults)
nearly
all
bottom
habitats
from
intertidal
zone
to
at
least
427
m
4
5
NG
5
NG
6
60
­
114
cm
TL
offshore,
midwater
and
bottom
tows
in
Hecate
Strait
and
Queen
Charlotte
Sound,
B.
C.
6
7
NG
(
adult)
7
NG
8
NG
Nanaimo,
B.
C.
8
9
NG
(
adult)
Kelp
bed
bottom
9
10
470­
740
mm
SL
(
juv
&
adult)
S
Cal
rocky
inshore
and
kep
beds
10
12
NG
demersal;
west
coast
Vancouver
Island,
B.
C.
12
18
>
45
cm
FL
(
adult
estimate)
various
bottom
(
broken
rock/
boulder
or
bedrock
wall)
18
19
see
data
(
juv
and
adult)
SW
Vancouver
Island
19
19
see
data
(
juv
and
adult)
Queen
Charlotte
Sound
19
Page
577
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
CB
CC
CD
CE
CF
piscivore:
(
sandlance,
herring,
flounders,
Pacific
cod,
whiting,

small
lingcod,
crabs,
shrimp,
squid)
1
NF
early
am
to
afternoon
(
shallow):
sandlances,

flatfishes;
afternoon
(
deeper):
herring,
dogfish
1
sea
lions,
harbour
seals
1
NF
sand
lances,
herring,
flounders,
young
lingcod,
dogfish,
gray
cod,
whiting,
crabs,
shrimp,
squid
2
harbour
seals
2
fish,
prosobranch
gastropod
3
size
NG
(
juv
or
adult):
harbor
seal
(
Phoca
vitulina
richardsi
)
3
carnivore:
fishes
(
juv
rockfishes,
Sebastes
jordani
,
other),

cephalopod,
gastropods,
decapods
(
see
ref
for
ID)
4
opportunistic
carnivores:
fish
(
Pacific
herring,
Pacific
sand
lance,
flatfish,
rockfish,
spiny
dogfish,
young
lingcod,
Pacific
cod,
Pacific
hake,
sablefish,
Pacific
tomcod,
salmon);
inverts
(
crabs,
shrimps,
squid,
octopus)
5
carnivore:
sandlance,
small
lingcod,
sole,
blackcod,
herring,

euphausiids
6
omnivore:
rockfishes,
herring,
squid,
small
sharks,
small
lingcod
7
carnivore:
fish
(
herring,
perch),
crustacea
(
Cancer
productus
,

spider
crab,
hermit
crab)
8
mesocarnivore
9
fish
(
halfmoon,
topsmelt),
squid,
algae
10
herring,
rex
sole,
unid
rockfish,
unid
fish
remains,
sablefish,

squid,
octopus,
tomcod,
unid
flatfish,
unid
digested
matter
12
NG
18
Pacific
hake,
butter
sole,
octopus,
shrimp
unid
fish
19
Pacific
hake,
Pacific
sand
lance,
arrowtooth
flounder,
flathead
sole,
Pacific
lamprey,
squid,
unid
fish
19
Page
578
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
CG
CH
CI
CJ
CK
CL
CM
NF
1
NG
1
1.4
(
0.1)
­
1.6
(
0.1)
1
31.7
­
51.2
1
0.046
(
0.017)
­
0.052
(
0.018)
1
0.044
(
0.016)
­
0.057
(
0.021);
mean:
0.050
(
0.001)
allozymes
1
Page
579
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
CN
CO
CP
CQ
CR
CS
CT
1
NG
1
NG
Commercial
/
58
cm
FL?
(
6yr)
1
trawl
and
handline
1
NG
1
NG
1
NG
1
Sport
/
27.5
cm
SL
(
1
yr)
2
Sport
/
46
cm
SL
(
2
yr)
2
creel
census
and
hook
&
line
2
age
frequency
analysis
2
NG
2
NG
2
Page
580
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
CU
CV
CW
1
0.20
1
0.20
1
0.40
2
0.42­
0.64
(
Male)
2
0.30­
0.61
(
Male)
2
0.72­
1.25
(
Male)

2
0.24­
0.44
(
Female)
2
0.05­
0.33
(
Female)
2
0.29­
0.77
(
Female)

3
NG
3
NG
3
0.12­
0.42
4
0.724
4
1.450
(
year
1:
1960­
1961)
4
2.174
(
year
1:
1960­
1961)

4
0.724
4
0.898
(
year
2:
1961­
1962)
4
1.622
(
year
2:
1961­
1962)

5
0.32
(
Male)
5
0.255
(
Male,
Sport,
1991)
5
NG
5
0.32
(
Male)
5
0.312
(
Male,
Sport,
1992)
5
NG
5
0.32
(
Male)
5
0.372
(
Male,
Sport,
1993)
5
NG
5
0.32
(
Male)
5
0.848
(
Male,
Sport,
1994)
5
NG
5
0.32
(
Male)
5
0.603
(
Male,
Trawl,
1991)
5
NG
5
0.32
(
Male)
5
0.589
(
Male,
Trawl,
1992)
5
NG
5
0.32
(
Male)
5
0.617
(
Male,
Trawl,
1993)
5
NG
5
0.32
(
Male)
5
0.283
(
Male,
Trawl,
1994)
5
NG
Page
581
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
CX
CY
CZ
tag­
recapture;
length
distributions
&
vertebral
age
estimates
1
1
6­
7
yr
1
NG
mark­
recapture
2
2
none
2
NG
mark­
recapture
2
2
none
2
NG
catch
and
CPUE
3
3
none
3
NG
tag­
recapture
&
fishing
effort
4
4
none
4
0.483
(
year
1960)

tag­
recapture
&
fishing
effort
4
4
none
4
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
Page
582
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
DA
DB
DC
1
NG
NF
Parasites:
trematodes
(
Derogenes
varicus,
Genolinea
robusta,
Hemiurus
levinseni,

Lecithochirium
exodicum
)
in
stomach
1
2
NG
Parasite:
nematode
(
Cucullanus
elongatus
)
in
intestine
2
2
NG
Parasite:
copepod
(
Lepeophtheirus
trifidus
)
on
body
surface
3
3
NG
Parasite:
copepod
(
Lepeophtheirus
breviventris
)
in
buccal
cavity
4
4
NG
spine
penetration:
diatom
(
Chaetoceros
convolutus
)
with
siliceous
spines
embedded
in
gill
tissue;
possibly
lethal
5
4
NG
Parasite:
copepod
(
Lepeophtheirus
breviventris
)
in
mouth
6
5
NG
Parasite:
copepod
(
Lepeophtheirus
pravipes
)
6,8
5
NG
Parasite:
isopod
(
Livonica
vulgaris
)
in
branchial
chamber
7
5
NG
Parasite:
copepod
(
Homoiotes
palliata
)
9
5
NG
Parasite:
copepod
(
Chondracanthus
narium
)
in
nasal
cavity
10
5
NG
Parasites:
trematodes
(
Lecithaster
gibbosus
,
Parahemiuris
merus,
Rhipidocotyle
elongata,
Stephanostomum
tristephanum,
Tubulovesicula
lindbergi
)
11
5
NG
Parasites:
Digenea
(
5
sp)
11
5
NG
Parasites:
cestodes
(
Nybelinia
surmenicola,
Phyllobothrium
sp.)
11
5
NG
Parasites:
nematodes
(
Anisakis
sp.,
Porrocaecum
sp)
11
Page
583
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
DD
DE
DF
DG
>
5
naut
mi
(
9%
of
419
recaptures);
80
naut
mi
(
one
case
of
419
recaptures)
1
1
several
resident
on
reef
in
Depoe
Bay,
Oregon
(
mature;
n=
6)
point
of
release
1
juveniles
(
y­
o­
y)
school
on
open
sand
bottom
areas
from
June­

September
1,2
no
movement
(
42.2%
of
206
recaptures);
 
1
naut
mi
(
21.4%
of
208
recaptures)
2
return
to
home
site
(
60
h)
after
displacement
up
to
2.8
km
(
n=
5)
2
>
1
naut
mi
(
36.4%
of
208
recaptures);
80
naut
mi
(
one
case
of
208
recaptures)
2
one
individual
transplanted
&
recaputured
at
original
site
(
6
miles)
3
>
1
naut
mi
(
21.3%
of
342
recaptures);
>
5
naut
mi
(
9.4%
of
342
recaptures)
3
transplants
and
returns
suggest
homing
4
No
movement
4
1
naut
mi
(
1
of
15
recaptures);
3
naut
mi
(
1
of
15
recaptures)
5
no
movement
(
74%
of
284
recaptures);
movement
(
5%
of
284
recaptures)
6
80
miles
(
129
km)
7
max
movement
99
km
(
male
65.5
cm
FL);
max
movement
71
km
(
female
53.5
cm
FL)
8
female
mean
linear
displacement
rate
1040
m/
d;
male
mean
linear
displacement
rate
500
m/
d
8
non­
migratory
(<
8.1
km;
80.7%
of
363
recaptures);

migratory
( 
8.1
km;
19.3%
of
363
recaptures);

migratory
(
8.1­
50
km;
12.7%
of
363
recaptures
);

migratory
(>
50
km;
6.6%
of
363
recap
non­
migratory
(<
8.1
km;
50.3%
of
149
recaptures);

migratory
( 
8.1
km;
49.7%
of
149
recaptures);

migratory
(
8.1­
50;
40.9%
of
149
recaptures);
migratory
(>
50
km;
8.7%
of
149
recaptures)
10
 
10
km;
93.7%
of
2476
recaptures;
11­

50
km;
5.2%
of
2476
recaptures;
>
50
km;

1.2%
of
2476
recaptures;
therefore,

nonmigratory
11
 
10
km;
95.4%
of
722
recaptures;
11­

50
km;
4.3%
of
722
recaptures;
>
50
km;
0.003%
of
722
recaptures;
therefore,

non­
migratory
11
Page
584
of
1167
5­
2220e
­
Hexagrammidae
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
DH
Males:
spawn
and
spawning
sites
3,6
Males:
nests
4
Page
585
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
A
B
C
D
1
­
Eschmeyer
1998
1
­
Gates
and
Frey
1974
1
­
Miller
and
Lea
1972
2
­
Miller
and
Lea
1972
2
­
Miller
and
Lea
1972
2
­
Eschmeyer
and
Herald
1983
3
­
Robins
et
al.
1991
3
­
Robins
et
al.
1991
3
­
Love
et
al.
2000
Page
586
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
E
F
G
H
1
­
Miller
and
Lea
1972
1
­
Chatwin
1956a
1
­
Laidig
et
al.
1997
1
­
Miller
and
Lea
1972
2
­
Love
et
al.
2000
2
­
Hart
1967
2
­
Karpov
and
Kwiecien
1988
2
­
Hart
1973
3
­
Orlov
1998a
3
­
Wendler
1953
(
Eqs
suspect)
3
­
Rothrock
1973
3
­
Eschmeyer
and
Herald
1983
Page
587
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
I
J
K
L
M
N
O
P
Q
8
west
coast
Canada
(
FL)
No
No
No
No
No
No
No
No
9
British
Columbia
(
NG)
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
10
British
Columbia
(
TL)
1062
(
NG)
1171
(
NG)
No
0.079
(
NG)
0.149
(
NG)
No
­
8.06
(
NG)
­
2.15
(
NG)

10
British
Columbia
(
TL)
1436
(
NG)
1096
(
NG)
No
0.042
(
NG)
0.192
(
NG)
No
­
10.77
(
NG)
­
1.94
(
NG)

11
northern
Puget
Sound
(
TL)
No
No
1149.99
No
No
0.1280
No
No
12
Puget
Sound
(
TL)
No
No
No
No
No
No
No
No
21
PSMC
US­
Vancouver
and
Columbia
areas
(
cm
TL)
166.2
(
14.838)
146.04
(
2.438)
No
0.0449
(
0.0122)
0.0897
(
0.0034)
No
­
4.84
(
0.6241)
­
1.61
(
0.1095)

21
PSMC
US­
Vancouver
and
Columbia
areas
(
TL)
93.21
(
5.35)
131.05
(
3.84)
No
0.1694
(
0.0283)
0.1137
(
0.0086)
No
NG
NG
13
PSMC
US­
Vancouver
and
Columbia
areas
(
LCN,
Lincod­
north;
TL)
917.48
(
NG)
1289.77
(
NG)
No
0.170
(
NG)
0.110
(
NG)
No
L1=
388.69
(
NG)
L1=
422.24
(
NG)

23
Columbia
INPFC
Area
(
Oregon;
TL)
816
(
NG)
1319
(
NG)
No
0.205
(
NG)
0.097
(
NG)
No
­
1.63
(
NG)
­
2.10
(
NG)

13
PSMC
Eureka,
Monterey,
Conception
areas
(
LCS,
Lingcod­
south;
TL)
762.51
(
NG)
1279.04
(
NG)
No
0.224
(
NG)
0.091
(
NG)
No
L1=
325.40
(
NG)
L1=
336.63
(
NG)

14
PSMC
Eureka,
Monterey,
Conception
areas
(
LCS,
Lingcod­
south;
TL)
853
(
NG)
1266
(
NG)
No
0.19
(
NG)
0.11
(
NG)
No
­
0.95
(
NG)
­
0.79
(
NG)

15
northern
&
central
California
(
Crescent
City
to
Morro
Bay;
FL)
No
No
No
No
No
No
No
No
16
central
California
(
TL)
No
No
No
No
No
No
No
No
17
central
California
(
Point
Reyes
&
Gulf
of
Farallones;
SL);
Gompertz
No
No
No
No
No
No
No
No
18
central
California
(
TL)
No
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
see
curve
22
central
California
(
TL)
No
No
741
No
No
NG
No
No
23
Monterey
INPFC
Area
(
TL)
817
(
NG)
1256
(
NG)
No
0.156
(
NG)
0.089
(
NG)
No
­
2.49
(
MG)
­
2.43
(
NG)

1
­
Rosenthal
et
al.
1981
2
­
Munk
2000
3
­
Chatwin
1954
Page
588
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
R
S
T
U
V
W
No
No
No
No
8
fin
rays/
sections
&
OTC
OTC;
 
14
yr
see
curve
see
curve
see
curve
see
curve
9
fin
rays/
thin
section
NG
No
No
No
No
10
fin
rays/
thin
sections
No
No
No
No
No
10
scales/
stained
No
­
2.1097
No
No
No
11otoliths/
whole
&
cleared
No
No
No
No
No
12
fin
rays/
thin
sections
No
No
0.52
0.71
No
21
Average
length
at
age
data;
unconstrained
VBGF
No
No
0.96
0.99
No
21
Average
length
at
age
data;
constrained
VBGF,
L(
age
1)
=
30
cm
No
No
No
No
No
13
fin
rays/
thin
sections;
L
=
Linf
+
(
L1­
Linf)*
exp(
K*(
1­
Age))
unknown
No
No
No
No
23
fin
rays/
thin
sections
No
No
No
No
No
13
fin
rays/
thin
sections;
length
composition;
L
=
Linf
+
(
L1­
Linf)*
exp(
K*(
1­
Age))
unknown
No
No
No
No
14
see
Jagielo
et
al.
1997
14
see
Jagielo
et
al.
1997
No
No
No
No
15
fin
rays/
cross
section
No
No
No
No
No
16
fin
rays/
sections
&
captive
w/
OTC
( 
2y);
growth
curves,
no
parameters
captive/
OTC
No
No
No
No
17
otoliths/
whole:
daily
increments,
Gompertz
growth,
size
at
hatching
No
see
curve
see
curve
see
curve
see
curve
18
otoliths/
whole
&
cleared
No
NG
No
No
No
22
none
/
length
frequency;
Ford
Walford
plot
analysis:
l
t+
1
=
Linf
(
1­
e­
K)
+
e
­
K
lt
No
No
No
No
No
23
fin
rays/
thin
sections
No
1
­
Rosenthal
et
al.
1981
2
­
Munk
2000
3
­
Chatwin
1954
Page
589
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
X
Y
Z
AA
AB
No
NG
between
structures
between
structures
intra­
reader
intra­
reader
intra­
reader
intra­
reader
between
readers
unknown
ageing
error
modeled
using
LCN
error
14
see
Jagielo
et
al.
1997
unknown
No
No
No
No
unknown
1
­
Richards
and
Hand
1988
1
­
Chatwin
1956a
1
­
Bargmann
1982
2
­
Phillips
1959
2
­
Cass,
McFarlane,
Rutherford,
Barber
1984
3
­
Forrester
1969a
3
­
Barker
1979
Page
590
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
AC
AD
AE
AF
AG
1
­
Jewell
1968
1
­
Matarese
et
al.
1989
1
­
Feder
et
al.
1974
1
­
Clemens
and
Wilby
1961
2
­
Low
and
Beamish
1978
2
­
Phillips
1959
2
­
Forrester
1969a
2
­
Wilby
1937
3
­
LaRiviere
et
al.
1981
3
­
LaRiviere
et
al.
1981
3
­
Shaw
and
Hassler
1989
Page
591
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
AH
AI
AJ
AK
1
­
Forrester
1969a
1
­
Forrester
1969a
1
­
Forrester
1969a
1
­
Forrester
1969a
2
­
Fitch
and
Lavenberg
1971
2
­
Fitch
and
Lavenberg
1971
2
­
Fitch
and
Lavenberg
1971
2
­
Fitch
and
Lavenberg
1971
3
­
Phillips
1959
3
­
Phillips
1959
3
­
Phillips
1959
3
­
Phillips
1959
Page
592
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
AL
AM
AN
AO
AP
AQ
AR
1
­
Forrester
1969a
1
­
Forrester
1969a
1
­
Forrester
1969a
1
­
Gordon
1994
2
­
Fitch
and
Lavenberg
1971
2
­
Fitch
and
Lavenberg
1971
2
­
Fitch
and
Lavenberg
1971
2
­
Forrester
1969a
3
­
Phillips
1959
3
­
Phillips
1959
3
­
Phillips
1959
3
­
Cass
et
al.
1990
Page
593
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
1
­
Cass
et
al.
1990
1
­
Cass
et
al.
1990
1
­
Forrester
1969a
2
­
Crow
1995
2
­
Rothrock
1973
2
­
Wilby
1937
3
­
Crow
et
al.
1997
3
­
Phillips
1959
Page
594
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
BF
BG
BH
BI
BJ
1
­
Hart
1967
1
­
Forrester
1969a
1
­
Forrester
1969a
1
­
Grosse
1982
2
­
Adams
et
al.
1999
2
­
Matarese
et
al.
1989
2
­
Gorbunova
1970
2
­
Jagielo
et
al.
2000
3
­
Wilby
1937
3
­
Hand
and
Richards
1991
Page
595
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
BK
BL
BM
18
NG
3
m,
intertidal,
large
rocks
covered
with
oysters,
brown
alga
18
quillback
rockfish
(
Sebastes
maliger
)
6,15
yelloweye
rockfish
(
Sebastes
ruberrimus
)
6
copper
rockfish
(
Sebastes
caurinus
)
6,17
kelp
greenling
(
Hexagrammos
decagrammus
)
10
striped
surfperch
(
Embiotoca
lateralis
)
10
pile
perch
(
Rhacochilus
vacca
)
11
longfin
sculpin
(
Jordania
zonope
)
10
Puget
Sound
sculpin
(
Artedius
meanyi
)
10
cabezon
(
Scorpaenichthys
marmoratus
)
11
dusky
perch
11
1
­
Matarese
et
al.
1989
1
­
Matarese
et
al.
1989
3
­
LaRiviere
et
al.
1981
2
­
Forrester
1969a
2
­
Forrester
1969a
6
­
Cass
et
al.
1990
3
­
LaRiviere
et
al.
1981
3
­
LaRiviere
et
al.
1981
10
­
Low
and
Beamish
1978
Page
596
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
BN
BO
BP
BQ
BR
8
31­
50
mm
FL
pelagic,
surface
8
medium
calanoids,
small
calanoids,
decapod
larvae,
crustacean
eggs,
large
calanoids,
herring
larvae
8
8
51­
70
mm
FL
pelagic,
surface
8
medium
calanoids,
juvenile
herring,
large
calanoids,
small
calanoids,
decapod
larvae
8
8
>
70
mm
FL
(
larvae
&
juv)
pelagic,
surface,
inshore
8
piscivore:
juvenile
herring
8
10
9.5
­
11
mm
TL
(
hatching)
laboratory
rearing
10
NG
10
11
11­
12
mm
TL
(
hatching)
laboratory
rearing
11
NG
11
12
10
­
65
mm
FL
surface
waters,
Saanich
Inlet,
Vancouver
Isl.
12
copepods
(
eggs,
Pseudocalanus
minutus
,
Acartia
longiremis,

Calanus
pacificus,
Calanus
sp,
nauplii,
copepodites,
Calanus
plumchrus,
Scolecithricella
sp,
Trotanus
discaudatus
,

Megalopa),
decapods
(
Caridean
zoea,
Brachyura
zoea),

amphipod
(
Parathemisto
pacif
13
19
­
55
mm
FL
surface
waters,
Strait
of
Georgia
13
copepods
(
eggs,
Pseudocalanus
minutus
,
Acartia
longiremis,

Calanus
sp,
nauplii,
Metridia
sp.,
Epilabidocera
amphitrites
,

fragments),
trematode
(
Lecithaster
gibbosus
),
fish
(
Stichaeidae,

Ammodytes
hexapterus
)
13
14
11
­
16.5
mm
TL
(
6
­
24
days
old)
in
captivity,
fed
wild
plankton
from
surface
waters
14
Calanoid
copepods/
copepodites,
crustaceans
(
larvae,
parts),

barnacle
cypris,
gammarid
amphipod,
Mysid
or
Natantia
14
16
6­
12
mm
FL
(
hatching)
surface
waters,
Strait
of
Georgia
16
NG
16
1
­
Matarese
et
al.
1989
1
­
Matarese
et
al.
1989
1
­
Matarese
et
al.
1989
2
­
Barraclough
1967a
2
­
Barraclough
1967a
2
­
Barraclough
1967a
3
­
Barraclough
1967b
3
­
Barraclough
1967b
3
­
Barraclough
1967b
Page
597
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
BS
BT
BU
BV
9
65
mm
FL
pelagic
surface
(
Strait
of
Georgia,
B.
C.)
9
carnivore:
copepod
(
Calanus
plumchrus
),
amphipod
(
Parathemisto
pacifica
)
9
9
72
mm
FL
pelagic
surface
(
Strait
of
Georgia,
B.
C.)
9
carnivore:
copepod
(
Calanus
plumchrus
)
9
10
470­
740
mm
SL
(
juv
&
adult)
S
Cal
rocky
inshore
and
kep
beds
10
fish
(
halfmoon,
topsmelt),
squid,
algae
10
11
77
mm
FL
surface
waters
Strait
of
Georgia,
B.
C.
11
copepod
(
Calanus
plumchrus
)
11
13
>
70
mm
FL
(
larvae
&
juv)
pelagic,
surface,
inshore
13
piscivore:
juvenile
herring
13
16
NG
(
juvenile)
nearshore,
subtidal,
Pysht
River
mouth
16
piscivore:
fish
(
Unid.
Teleosts,
Ammodytes
hexapterus
,
Clupea
harengus
pallasi
,
Sebastes
sp),
crustaceans
(
Mysideacea,
Unid
Caridea,
Crangonidae)
16
17
12­
27
cm
FL
(
YOY)
benthic
(
73­
146
m),
Strait
of
Georgia
17
piscivore:
fish
(
herring,
sandlance
flatfish,
shiner
perch)
17
18
21­
45
cm
FL
(
juv
estimate)
various
bottom
(
fine
sediment,
cobble,
rock/
boulder,
bedrock
slope,
walls,
ledges)
18
NG
18
19
see
data
(
juv
and
adult)
SW
Vancouver
Island
19
Pacific
hake,
butter
sole,
octopus,
shrimp
unid
fish
19
19
see
data
(
juv
and
adult)
Queen
Charlotte
Sound
19
Pacific
hake,
Pacific
sand
lance,
arrowtooth
flounder,
flathead
sole,
Pacific
lamprey,
squid,
unid
fish
19
1
­
Cass
et
al.
1990
1
­
Forrester
1969a
1
­
Forrester
1969a
1
­
Forrester
1969a
2
­
Wilby
1937
2
­
Wilby
1937
2
­
Wilby
1937
3
­
Moulton
1977
3
­
Moulton
1977
3
­
Moulton
1977
Page
598
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
BW
BX
BY
BZ
CA
20
420
­
875
mm
TL
Diablo
Cove,
CA
20
1
­
Trumble
1995
1
­
Forrester
1969a
1
­
Forrester
1969a
2
­
Wilby
1937
2
­
Wilby
1937
3
­
Moulton
1977
3
­
Moulton
1977
Page
599
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
CB
CC
CD
CE
CF
fish,
octopus,
juv
yellowtail
rockfish,
blue
rockfish,
bonehead
sculpin
(
Artedius
notospilotus
)
20
1
­
Forrester
1969a
1
­
Wilby
1937
1
­
Cass
et
al.
1990
2
­
Wilby
1937
2
­
Klyashtorin
1970
2
­
Olesiuk
et
al.
1990
3
­
Moulton
1977
3
­
Trumble
1995
Page
600
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
CG
CH
CI
CJ
CK
CL
CM
1
­
Jagielo
et
al.
1996
1
­
Jagielo
et
al.
1996
1
­
Jagielo
et
al.
1996
1
­
Jagielo
et
al.
1996
1
­
Jagielo
et
al.
1996
1
­
Jagielo
et
al.
1996
Page
601
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
CN
CO
CP
CQ
CR
CS
CT
1
­
Cass
et
al.
1990
1
­
Cass
et
al.
1990
1
­
Cass
et
al.
1990
1
­
Cass
et
al.
1990
1
­
Cass
et
al.
1990
1
­
Cass
et
al.
1990
1
­
Cass
et
al.
1990
2
­
Barker
1979
2
­
Barker
1979
2
­
Barker
1979
2
­
Barker
1979
2
­
Barker
1979
2
­
Barker
1979
Page
602
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
CU
CV
CW
5
0.18
(
Female)
5
0.196
(
Female,
Sport,
1991)
5
NG
5
0.18
(
Female)
5
0.182
(
Female,
Sport,
1992)
5
NG
5
0.18
(
Female)
5
0.356
(
Female,
Sport,
1993)
5
NG
5
0.18
(
Female)
5
0.510
(
Female,
Sport,
1994)
5
NG
5
0.18
(
Female)
5
1.025
(
Female,
Trawl,
1991)
5
NG
5
0.18
(
Female)
5
0.740
(
Female,
Trawl,
1992)
5
NG
5
0.18
(
Female)
5
0.697
(
Female,
Trawl,
1993)
5
NG
5
0.18
(
Female)
5
0.385
(
Female,
Trawl,
1994)
5
NG
6
0.77
(
calc
from
annual
survival)
6
0.32
(
calc
from
fishing
survival)
61.11
(
calc
from
overall
survival)

7
NG
7
NG
7
0.11
(
Female)

7
NG
7
NG
7
0.47
(
Male)

7
NG
7
NG
7
0.35
(
Sexes
combined)

7
0.26
(
Sexes
combined)
7
0.09
(
Sexes
combined)
7
0.35
(
Sexes
combined)

8
NG
8
NG
8
0.3999
(
Sexes
combined)

8
0.3060
8
0.0939
8
0.3999
9,11
0.56
9
sport:
0.075­
0.128
(
mean
0.099)
9
0.208­
1.091
(
mean
0.663)

10
Male:
0.32
10
NG
10
NG
10
Female:
0.18
10
NG
10
NG
11
0.20­
0.80
11
NG
11
NG
11
0.28
11
NG
11
NG
11
0.25
11
NG
11
NG
12
0.36­
0.80
12
NG
12
NG
13
Male:
0.23­
0.39
(
avg
0.32)
13
NG
13
NG
13
Female:
0.16­
0.19
(
avg
0.18)
13
NG
13
NG
14
0.26
(
0.2
­
0.4)
14
NG
14
NG
1
­
Chatwin
1958
1
­
Chatwin
1958
1
­
Chatwin
1958
2
­
Smith
et
al.
1990
2
­
Smith
et
al.
1990
2
­
Smith
et
al.
1990
3
­
Richards
and
Hand
1989
3
­
Richards
and
Hand
1989
3
­
Richards
and
Hand
1989
Page
603
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
CX
CY
CZ
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
tag­
recapture
5
5
NG
(
see
Jagielo
1994)
5
NG
catch
&
effort,
growth
rates
&
recruitment,
tag­
recapture
6
6
6­
12
yr
6
0.33
(
1984,
B.
C.
Canada)
0.52
(
1985,
B.
C.
Canada)

catch
curves
of
age
composition
(
Ricker
1975)
7
7
4­
10
yr
7
NG
catch
curves
of
age
composition
(
Ricker
1975)
7
7
4­
10
yr
7
NG
catch
curves
of
age
composition
(
Ricker
1975)
7
7
4­
10
yr
7
NG
Tagging
and
Seber
model
(
1970)
7
7
none
7
0.08
(
sexes
combined)

age­
length
formulation
8
8
0­
20
yr
8
NG
tag­
recapture
&
non­
tag
return
estimate
8
8
0­
20
yr
8
0.0774
tag­
recapture
&
fishery
data
9
9
none
9
0.048­
0.089
(
mean
0.075)

avg
values
from
Jagielo
1994
10
10
see
Jagielo
1994
10
NG
avg
values
from
Jagielo
1994
10
10
see
Jagielo
1994
10
NG
tagging
11
11
see
Jagielo
1994
11
NG
Age
composition
11
11
NG
11
NG
empirical
average
11
11
see
Jagielo
1994
11
NG
tag­
recapture
12
12
NG
12
0.119
­
0.305
3
empirical
models
13
13
NG
13
NG
3
empirical
models
13
13
NG
13
NG
catch
curves
and
tagging
(
Cass,
Cameron,
and
Barber
1983)
14
14
NG
14
0.03
­
0.21
(
3
areas)

1
­
Chatwin
1958
1
­
Chatwin
1958
1
­
Chatwin
1958
2
­
Smith
et
al.
1990
2
­
Smith
et
al.
1990
2
­
Smith
et
al.
1990
3
­
Richards
and
Hand
1989
3
­
Richards
and
Hand
1989
3
­
Richards
and
Hand
1989
Page
604
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
DA
DB
DC
5
NG
Parasite:
tricodinid
ciliates
on
gills
12
5
NG
Parasite:
nematodes
(
Phocanema
&
Anisakis)
in
muscle
and
viscera
15
5
NG
Parasite:
isopod
(
Lironeca
vulgaris)
in
gill
or
buccal
cavities
16
5
NG
Parasites:
15
Families;
~
32
Species;
See
Refs
17
5
NG
5
NG
5
NG
5
NG
6
NG
7
NG
7
NG
7
NG
7
NG
8
NG
8
0.2348
9
NG
10
NG
10
NG
11
NG
11
NG
11
NG
12
NG
13
NG
13
NG
14
NG
1
­
Chatwin
1958
1
­
Lloyd
1938
2
­
Smith
et
al.
1990
2
­
Berland
1983
3
­
Richards
and
Hand
1989
3
­
Shiino
1965
Page
605
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
DD
DE
DF
DG
no
movement;
66.7
%
of
9
recaptures;
39
km;

11.1
%
of
9
recaptures;
228
km;
11.1
%
of
9
recaptures;
unknown
distance;
11.1%
of
9
recaptures
12
no
movement;
100%
of
1
tag­
recapture
13
no
movement
(
81%
of
36
recaptures);
moved
good
distance;
19%
of
36
recaptures
14
growth
info
15
growth
info
16
Male:
possible
directed
inshore
spawning
migration
during
late
fall
and
winter;
Female:
possible
brief
inshore
spawning
migration
17,18,19
no
movement
(
61%
of
383
recaptures);
 
5
km
(
92%
of
383
recaptures);
7­
19
km
(
5%
of
383
recaptures);
 
20
km
(
3%

of
383
recaptures)
20
no
movement
inshore/
offshore
21
0­
10
naut
mi
(
95.3%
of
213
recaptures);
>
10
naut
mi
(
4.7%
of
213
recaptures);
greatest
distance
160
nautical
miles
22
no
movement
(
97%
of
1442
recaptures);
mean
movement
44
km
(
3%
of
1442
recaptures)
23
 
5
naut
mi
(
11%
of
624
recaptures);
>
25
naut
mi
(
5%
of
624
recaptures)
24
 
5
km
(
96%
of
99
recaptures);
6­
40
km
(
4%
of
99
recaptures)
25
movement
(
13
­
298
km)
26
no
substantial
migration
27
no
movement
(
100%
of
12
recaptures)
28
<
1
km
(
43%
of
21
recaptures)
>
20
km
(
19%

of
21
recaptures)
29
no
movement
(
57%
of
14
recaptures)
11­
442
km
(
43%
of
14
recaptures)
30
1
­
Phillips
1959
1
­
Coombs
1979
1
­
Reeves
1966
1
­
Buckley
and
Hueckel
1984a
2
­
Hart
1943b
2
­
Matthews
1992
2
­
Buckley
and
Hueckel
1984b
3
­
Chatwin
1956b
3
­
Chatwin
1956b
Page
606
of
1167
5­
2220e
­
Hexagrammidae
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
DH
1
­
DeMartini
1986
2
­
Barker
1979
3
­
LaRiviere
et
al.
1981
Page
607
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
A
B
C
D
4
­
Quast
1960
4
­
Reilly
et
al.
1994
4
­
Cass
et
al.
1990
5
­
Phillips
1959
6
­
Wilby
1937
Page
608
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
E
F
G
H
4
­
Cass
et
al.
1990
4
­
Grosse
1982
4
­
Cass
et
al.
1988
4
­
Fitch
and
Lavenberg
1971
5
­
Bargmann
1982
5
­
Wendler
1953
5
­
Wilby
1937
6
­
Forrester
and
Thomson
1969a
6
­
Forrester
1973
6
­
Clemens
and
Wilby
1961
7
­
Jagielo
et
al.
2000
7
­
Phillips
1960b
8
­
Adams
et
al.
1999
8
­
Shaw
and
Hassler
1989
9
­
Jagielo
et
al.
1997
10
­
Moulton
1977
11
­
Barker
1979
12
­
Phillips
and
Barraclough
1977
13
­
Jagielo
1994
14
­
Rothrock
1973
15
­
Cass
et
al.
1988
16
­
Richards
and
Hand
1988
17
­
Cass,
Selsby,
and
Richards
1986
Page
609
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
I
J
K
L
M
N
O
P
Q
4
­
Chatwin
1956a
5
­
Cass,
McFarlane,
Rutherford,
Barber
1984
6
­
Smith
and
McFarlane
1990
7
­
Forrester
1969a
8
­
Cass
and
Beamish
1983
9
­
Cass
et
al.
1990
10
­
Beamish
and
Chilton
1977
11
­
Moulton
1977
12
­
Bargmann
1982
13
­
Jagielo
et
al.
2000
14
­
Adams
et
al.
1999
15
­
Silberberg
et
al.
2000
16
­
Adams
et
al.
1994
17
­
Adams
et
al.
1993
18
­
Miller
and
Geibel
1973
19
­
Barker
1979
20
­
Bryant
1978
21
­
Jagielo
1994
22
­
Karpov
et
al.
1995
23
­
Adams
1994
24
­
Rothrock
1973
25
­
Cass,
Cameron,
and
Barber
1983
26
­
Davis
1986
27
­
Cass,
Beamish,
Smith
1984
28
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
29
­
Forrester
1973
30
­
Rutenberg
1970
31
­
Klyashtorin
1970
32
­
Gorbunova
1970
33
­
Burge
and
Schultz
1973
Page
610
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
R
S
T
U
V
W
4
­
Chatwin
1956a
5
­
Cass,
McFarlane,
Rutherford,
Barber
1984
6
­
Smith
and
McFarlane
1990
7
­
Forrester
1969a
8
­
Cass
and
Beamish
1983
9
­
Cass
et
al.
1990
10
­
Beamish
and
Chilton
1977
11
­
Moulton
1977
12
­
Bargmann
1982
13
­
Jagielo
et
al.
2000
14
­
Adams
et
al.
1999
15
­
Silberberg
et
al.
2000
16
­
Adams
et
al.
1994
17
­
Adams
et
al.
1993
18
­
Miller
and
Geibel
1973
19
­
Barker
1979
20
­
Bryant
1978
21
­
Jagielo
1994
22
­
Karpov
et
al.
1995
23
­
Adams
1994
24
­
Rothrock
1973
25
­
Cass,
Cameron,
and
Barber
1983
26
­
Davis
1986
27
­
Cass,
Beamish,
Smith
1984
28
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
29
­
Forrester
1973
30
­
Rutenberg
1970
31
­
Klyashtorin
1970
32
­
Gorbunova
1970
Page
611
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
X
Y
Z
AA
AB
4
­
Fitch
and
Lavenberg
1971
4
­
Beamish
et
al.
1978
5
­
Miller
and
Geibel
1973
6
­
Cass,
McFarlane,
Rutherford,
Barber
1984
7
­
Cass
et
al.
1990
8
­
Adams
et
al.
1999
9
­
Jagielo
et
al.
2000
10
­
Silberberg
et
al.
2000
11
­
Rothrock
1973
12
­
Bryant
1978
13
­
Cass,
Cameron,
and
Barber
1983
14
­
Cass,
Beamish,
Smith
1984
15
­
Rutenberg
1970
16
­
Klyashtorin
1970
17
­
Gorbunova
1970
18
­
Burge
and
Schultz
1973
Page
612
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
AC
AD
AE
AF
AG
4
­
Garrison
and
Miller
1982
4
­
Kendall
and
Vinter
1984
4
­
Feder
et
al.
1974
5
­
Phillips
1959
5
­
Reilly
et
al.
1994
6
­
Moulton
1977
6
­
DeMartini
1986
7
­
Wilby
1937
7
­
Quast
1960
8
­
Jewell
1968
8
­
Zolotov
and
Tokranov
1989
9
­
Low
and
Beamish
1978
9
­
Rutenberg
1970
10
­
Howard
1992
10
­
Klyashtorin
1970
11
­
DeMartini
1986
11
­
Gorbunova
1970
12
­
Clemens
and
Wilby
1961
13
­
O'Connell
1993
14
­
Crow
1995
15
­
Crow
et
al.
1997
Page
613
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
AH
AI
AJ
AK
4
­
Cass
et
al.
1990
4
­
Cass
et
al.
1990
4
­
Cass
et
al.
1990
4
­
Cass
et
al.
1990
5
­
Miller
and
Geibel
1973
5
­
Miller
and
Geibel
1973
5
­
Miller
and
Geibel
1973
5
­
Miller
and
Geibel
1973
6
­
Phillips
1960b
6
­
Phillips
1960b
6
­
Phillips
1960b
6
­
Phillips
1960b
7
­
Wilby
1937
7
­
Wilby
1937
7
­
Wilby
1937
7
­
Wilby
1937
8
­
Silberberg
et
al.
2000
8
­
Silberberg
et
al.
2000
8
­
Silberberg
et
al.
2000
8
­
Silberberg
et
al.
2000
9
­
Rothrock
1973
9
­
Rothrock
1973
9
­
Rothrock
1973
9
­
Rothrock
1973
10
­
Barker
1979
10
­
Barker
1979
10
­
Barker
1979
10
­
Barker
1979
11
­
Gordon
1994
11
­
Gordon
1994
11
­
Gordon
1994
11
­
Gordon
1994
12
­
Richards
et
al.
1990
12
­
Richards
et
al.
1990
12
­
Richards
et
al.
1990
12
­
Richards
et
al.
1990
13
­
Jagielo
1994
13
­
Jagielo
1994
13
­
Jagielo
1994
13
­
Jagielo
1994
14
­
Cass,
Beamish,
Smith
1984
14
­
Cass,
Beamish,
Smith
1984
14
­
Cass,
Beamish,
Smith
1984
14
­
Cass,
Beamish,
Smith
1984
15
­
Gorbunova
1970
15
­
Gorbunova
1970
15
­
Gorbunova
1970
15
­
Gorbunova
1970
16
­
Barss
et
al.
1982
16
­
Barss
et
al.
1982
16
­
Barss
et
al.
1982
16
­
Barss
et
al.
1982
17
­
Cass,
Selsby,
and
Richards
1986
17
­
Cass,
Selsby,
and
Richards
1986
17
­
Cass,
Selsby,
and
Richards
1986
17
­
Cass,
Selsby,
and
Richards
1986
Page
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Hexagrammidae
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86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
AL
AM
AN
AO
AP
AQ
AR
4
­
Cass
et
al.
1990
4
­
Cass
et
al.
1990
4
­
Cass
et
al.
1990
4
­
Fraser
1923
5
­
Miller
and
Geibel
1973
5
­
Miller
and
Geibel
1973
5
­
Miller
and
Geibel
1973
5
­
Low
and
Beamish
1978
6
­
Phillips
1960b
6
­
Phillips
1960b
6
­
Phillips
1960b
6
­
LaRiviere
et
al.
1981
7
­
Wilby
1937
7
­
Wilby
1937
7
­
Wilby
1937
7
­
Moulton
1977
8
­
Silberberg
et
al.
2000
8
­
Silberberg
et
al.
2000
8
­
Silberberg
et
al.
2000
8
­
Jewell
1968
9
­
Rothrock
1973
9
­
Rothrock
1973
9
­
Rothrock
1973
9
­
Wilby
1937
10
­
Barker
1979
10
­
Barker
1979
10
­
Barker
1979
10
­
Silberberg
et
al.
2000
11
­
Gordon
1994
11
­
Gordon
1994
11
­
Gordon
1994
11
­
Adams
et
al.
1993
12
­
Richards
et
al.
1990
12
­
Richards
et
al.
1990
12
­
Richards
et
al.
1990
12
­
Adams
and
Hardwick
1992
13
­
Jagielo
1994
13
­
Jagielo
1994
13
­
Jagielo
1994
13
­
Miller
and
Geibel
1973
14
­
Cass,
Beamish,
Smith
1984
14
­
Cass,
Beamish,
Smith
1984
14
­
Cass,
Beamish,
Smith
1984
14
­
Marliave
1975a
15
­
Gorbunova
1970
15
­
Gorbunova
1970
15
­
Gorbunova
1970
15
­
Simenstad
1971
16
­
Barss
et
al.
1982
16
­
Barss
et
al.
1982
16
­
Barss
et
al.
1982
16
­
Gorbunova
1970
17
­
Rothrock
1973
18
­
Clemens
and
Wilby
1961
19
­
Zolotov
1985
Page
615
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Hexagrammidae
74
75
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80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
4
­
Rothrock
1973
4
­
Cass
et
al.
1990
5
­
Klyashtorin
1970
5
­
Clemens
and
Wilby
1961
6
­
Gorbunova
1970
6
­
Fraser
1923
7
­
Gorbunova
1970
8
­
Klyashtorin
1970
Page
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Hexagrammidae
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80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
BF
BG
BH
BI
BJ
4
­
Cass
et
al.
1990
5
­
Jewell
1968
6
­
Low
and
Beamish
1978
7
­
LaRiviere
et
al.
1981
8
­
Appelbaum
et
al.
1995
9
­
Dempster
and
Rohrs
1983
10
­
Gorbunova
1970
Page
617
of
1167
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2220e
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Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
BK
BL
BM
4
­
Phillips
1959
4
­
Phillips
1959
11
­
Jewell
1968
5
­
Wilby
1937
5
­
Wilby
1937
12
­
Giorgi
1981
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
15
­
Murie
1995
7
­
Giorgi
and
Congleton
1984
7
­
Giorgi
and
Congleton
1984
16
­
Moulton
1977
8
­
O'Connell
1993
8
­
O'Connell
1993
17
­
Buckley
and
Hueckel
1985
9
­
DeMartini
1986
9
­
DeMartini
1986
10
­
Low
and
Beamish
1978
10
­
Low
and
Beamish
1978
11
­
Jewell
1968
11
­
Jewell
1968
12
­
Giorgi
1981
12
­
Giorgi
1981
13
­
Miller
and
Geibel
1973
13
­
Miller
and
Geibel
1973
14
­
Simenstad
1971
14
­
Simenstad
1971
15
­
Marliave
1975a
15
­
Marliave
1975a
16
­
Dempster
and
Rohrs
1983
16
­
Dempster
and
Rohrs
1983
17
­
Fraser
1915
17
­
Fraser
1915
18
­
Canfield
and
Buckley
1984
18
­
Canfield
and
Buckley
1984
19
­
Crow
et
al.
1997
19
­
Crow
et
al.
1997
20
­
Klyashtorin
1970
20
­
Klyashtorin
1970
21
­
Gorbunova
1970
21
­
Gorbunova
1970
Page
618
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1167
5­
2220e
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Hexagrammidae
74
75
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77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
BN
BO
BP
BQ
BR
4
­
Forrester
1969a
4
­
Forrester
1969a
4
­
Forrester
1969a
5
­
Adams
et
al.
1993
5
­
Adams
et
al.
1993
5
­
Adams
et
al.
1993
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
7
­
Robinson
et
al.
1968b
7
­
Robinson
et
al.
1968b
7
­
Robinson
et
al.
1968b
8
­
Phillips
and
Barraclough
1977
8
­
Phillips
and
Barraclough
1977
8
­
Phillips
and
Barraclough
1977
9
­
Richardson
and
Washington
1980
9
­
Richardson
and
Washington
1980
9
­
Richardson
and
Washington
1980
10
­
Marliave
1975a
10
­
Marliave
1975a
10
­
Marliave
1975a
11
­
Fraser
1915
11
­
Fraser
1915
11
­
Fraser
1915
12
­
Barraclough
et
al.
1968
12
­
Barraclough
et
al.
1968
12
­
Barraclough
et
al.
1968
13
­
Robinson
et
al.
1968a
13
­
Robinson
et
al.
1968a
13
­
Robinson
et
al.
1968a
14
­
Benson
1984
14
­
Benson
1984
14
­
Benson
1984
15
­
Gorbunova
1970
15
­
Gorbunova
1970
15
­
Gorbunova
1970
16
­
Cass
and
Scarsbrook
1984
16
­
Cass
and
Scarsbrook
1984
16
­
Cass
and
Scarsbrook
1984
Page
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Hexagrammidae
74
75
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78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
BS
BT
BU
BV
4
­
Miller
and
Geibel
1973
4
­
Miller
and
Geibel
1973
4
­
Miller
and
Geibel
1973
5
­
Chatwin
1954
5
­
Chatwin
1954
5
­
Chatwin
1954
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
7
­
Richards
et
al.
1985
7
­
Richards
et
al.
1985
7
­
Richards
et
al.
1985
8
­
Grosse
1982
8
­
Grosse
1982
8
­
Grosse
1982
9
­
Barraclough
1967c
9
­
Barraclough
1967c
9
­
Barraclough
1967c
10
­
Quast
1968d
10
­
Quast
1968d
10
­
Quast
1968d
11
­
Robinson
et
al.
1968b
11
­
Robinson
et
al.
1968b
11
­
Robinson
et
al.
1968b
12
­
Bryant
1978
12
­
Bryant
1978
12
­
Bryant
1978
13
­
Phillips
and
Barraclough
1977
13
­
Phillips
and
Barraclough
1977
13
­
Phillips
and
Barraclough
1977
14
­
Barraclough
et
al.
1968
14
­
Barraclough
et
al.
1968
14
­
Barraclough
et
al.
1968
15
­
Robinson
et
al.
1968a
15
­
Robinson
et
al.
1968a
15
­
Robinson
et
al.
1968a
16
­
Buckley
and
Hueckel
1984a
16
­
Buckley
and
Hueckel
1984a
16
­
Buckley
and
Hueckel
1984a
17
­
Cass,
Smith,
Barber,
and
Rinhofer
1983
17
­
Cass,
Smith,
Barber,
and
Rinhofer
1983
17
­
Cass,
Smith,
Barber,
and
Rinhofer
1983
18
­
Richards
and
Cass
1985
18
­
Richards
and
Cass
1985
18
­
Richards
and
Cass
1985
19
­
Cass,
Selsby,
and
Richards
1986
19
­
Cass,
Selsby,
and
Richards
1986
19
­
Cass,
Selsby,
and
Richards
1986
20
­
Burge
and
Schultz
1973
20
­
Burge
and
Schultz
1973
20
­
Burge
and
Schultz
1973
22
­
Orlov
1998b
22
­
Orlov
1998b
22
­
Orlov
1998b
Page
620
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Hexagrammidae
74
75
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77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
BW
BX
BY
BZ
CA
4
­
Miller
and
Geibel
1973
4
­
Miller
and
Geibel
1973
5
­
Cass
et
al.
1990
5
­
Cass
et
al.
1990
6
­
Taylor
1970
6
­
Taylor
1970
7
­
Phillips
1960b
7
­
Phillips
1960b
8
­
Fraser
1923
8
­
Fraser
1923
9
­
Quast
1968b
9
­
Quast
1968b
10
­
Quast
1968d
10
­
Quast
1968d
11
­
Richards
et
al.
1985
11
­
Richards
et
al.
1985
12
­
Cass,
Beamish,
Smith
1984
12
­
Cass,
Beamish,
Smith
1984
13
­
Long
1996
13
­
Long
1996
14
­
Henriques
1991
14
­
Henriques
1991
15
­
Zolomov
and
Tokranov
1991
15
­
Zolomov
and
Tokranov
1991
16
­
Rutenberg
1970
16
­
Rutenberg
1970
17
­
Klyashtorin
1970
17
­
Klyashtorin
1970
18
­
Richards
and
Cass
1985
18
­
Richards
and
Cass
1985
19
­
Cass,
Selsby,
and
Richards
1986
19
­
Cass,
Selsby,
and
Richards
1986
20
­
Burge
and
Schultz
1973
20
­
Burge
and
Schultz
1973
21
­
Trumble
1995
21
­
Trumble
1995
22
­
Orlov
1998b
22
­
Orlov
1998b
Page
621
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74
75
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77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
CB
CC
CD
CE
CF
4
­
Miller
and
Geibel
1973
5
­
Cass
et
al.
1990
6
­
Taylor
1970
7
­
Phillips
1960b
8
­
Fraser
1923
9
­
Quast
1968b
10
­
Quast
1968d
11
­
Richards
et
al.
1985
12
­
Cass,
Beamish,
Smith
1984
13
­
Long
1996
14
­
Henriques
1991
15
­
Zolomov
and
Tokranov
1991
16
­
Rutenberg
1970
17
­
Klyashtorin
1970
18
­
Richards
and
Cass
1985
19
­
Cass,
Selsby,
and
Richards
1986
20
­
Burge
and
Schultz
1973
21
­
Trumble
1995
22
­
Orlov
1998b
Page
622
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Hexagrammidae
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75
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77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
CG
CH
CI
CJ
CK
CL
CM
Page
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Hexagrammidae
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75
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79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
CN
CO
CP
CQ
CR
CS
CT
Page
624
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Hexagrammidae
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
CU
CV
CW
4
­
Reeves
1966
4
­
Reeves
1966
4
­
Reeves
1966
5
­
Jagielo
1999
5
­
Jagielo
1999
5
­
Jagielo
1999
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
7
­
Bargmann
1982
7
­
Bargmann
1982
7
­
Bargmann
1982
8
­
Barker
1979
8
­
Barker
1979
8
­
Barker
1979
9
­
Jagielo
1995
9
­
Jagielo
1995
9
­
Jagielo
1995
10
­
Jagielo
et
al.
2000
10
­
Jagielo
et
al.
2000
10
­
Jagielo
et
al.
2000
11
­
Adams
et
al.
1999
11
­
Adams
et
al.
1999
11
­
Adams
et
al.
1999
12
­
Forrester
1973
12
­
Forrester
1973
12
­
Forrester
1973
13
­
Jagielo
1994
13
­
Jagielo
1994
13
­
Jagielo
1994
14
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
14
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
14
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
Page
625
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1167
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Hexagrammidae
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75
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77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
CX
CY
CZ
4
­
Reeves
1966
4
­
Reeves
1966
4
­
Reeves
1966
5
­
Jagielo
1999
5
­
Jagielo
1999
5
­
Jagielo
1999
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
6
­
Cass
et
al.
1990
7
­
Bargmann
1982
7
­
Bargmann
1982
7
­
Bargmann
1982
8
­
Barker
1979
8
­
Barker
1979
8
­
Barker
1979
9
­
Jagielo
1995
9
­
Jagielo
1995
9
­
Jagielo
1995
10
­
Jagielo
et
al.
2000
10
­
Jagielo
et
al.
2000
10
­
Jagielo
et
al.
2000
11
­
Adams
et
al.
1999
11
­
Adams
et
al.
1999
11
­
Adams
et
al.
1999
12
­
Forrester
1973
12
­
Forrester
1973
12
­
Forrester
1973
13
­
Jagielo
1994
13
­
Jagielo
1994
13
­
Jagielo
1994
14
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
14
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
14
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
Page
626
of
1167
5­
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­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
DA
DB
DC
4
­
Reeves
1966
4
­
Kabata
1973
5
­
Jagielo
1999
5
­
Bell
1961
6
­
Cass
et
al.
1990
6
­
Fraser
1919
7
­
Bargmann
1982
7
­
Turner
et
al.
1969
8
­
Barker
1979
8
­
Wilson
1912
9
­
Jagielo
1995
9
­
Wilson
1905
10
­
Jagielo
et
al.
2000
10
­
Kabata
1969
11
­
Adams
et
al.
1999
11
­
Arai
1969
12
­
Forrester
1973
12
­
Bell
1962
13
­
Jagielo
1994
13
­
Zhukov
1960
14
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
14
­
Bykhovskii
1957
15
­
Dailey
et
al.
1981
16
17
­
Love
and
Moser
1983
Page
627
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
DD
DE
DF
DG
4
­
Anonymous
1954
4
­
Hueckel
and
Buckley
1984
5
­
Miller
and
Geibel
1973
6
­
Reeves
1966
7
­
Manzer
1946
8
­
Smith
et
al.
1990
9
­
Jagielo
1990
10
­
Mathews
and
LaRiviere
1987
11
­
Cass
et
al.
1990
12
­
Coombs
1979
13
­
Turner
et
al.
1969
14
­
Hart
1939
15
­
Hart
1940
16
­
Hart
1943a
17
­
Jagielo
1995
18
­
Jagielo
1999
19
­
Jagielo
et
al.
2000
20
­
Cass,
McFarlane,
Rutherford,
Barber
1984
21
­
Barss
and
Demory
1989
22
­
Davis
1986
23
­
Cass,
Cameron,
and
Barber
1983
24
­
Barss
and
Demory
1989
25
­
Cass,
McFarlane,
Smith,
Barber,
and
Rutherford
1986
26
­
Hueckel
and
Buckley
1984
27
­
Forrester
1973
28
­
Mathews
et
al.
1979
29
­
Cass,
Smith,
Barber,
and
Rinhofer
1983
30
­
DeMott
1983
Page
628
of
1167
5­
2220e
­
Hexagrammidae
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
DH
4
­
O'Connell
1993
5
­
Bryant
1978
6
­
Low
and
Beamish
1978
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629
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Hexagrammidae
1234
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Kyphosidae1,2
Girella
nigricans
1,2
1intertidal
to
30
(
Girellidae)
3
Kyphosidae1,2
Hermosilla
azurea
1,2
zebra
perch;
convict
fish;
perch1,2
1intertidal
to
27
(
Kyphosidae)
3
Kyphosidae1,2
Medialuna
californiensis
1,2
1surface
to
40
(
Scorpididae)
3
TAXONOMY
RANGE
opaleye;
green
perch;
black
perch;
blue­
eyed
perch;
bluefish;
Catalina
perch;
button
perch;

greenfish;
blue
bass;
blue­
eye;
blue
perch;

Jack
Benny;
California
opaleye
1,2,3,4
halfmoon;
blue
perch;
blue
bass;
medialuna;

Catalina
perch;
California
halfmoon;
Catalina
blue
perch;
balck
perch
1,2,3
northern
California
(
Klamath
River
estuary)
to
Gulf
of
California1,2
British
Columbia
(
Vancouver
I.)
to
Gulf
of
California1
Oregon
(
Otter
Rock)
to
southern
Baja
California
(
Cabo
San
Lucas)

1,3
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
F
G
H
I
J
K
LMNOP
Length
weight
equation
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
1Both:
W=
0.0000528SL2.93
1TL=
5.47SL1.21
(
SE=
7.66)
1
66
NF
NF
NF
NF
NF
NF
NF
NF
1TL=
­
0.44FL1.03
(
SE=
0.2.36)

1SL=
­
3.97TL0.827
(
SE=
6.34)

1SL=
­
14.96FL0.902
(
SE=
2.0)

1FL=
0.489TL0.972
(
SE=
2.3)

1FL=
5.82SL1.172
(
SE=
0.794)

NF
NF
1
44
NF
NF
NF
NF
NF
NF
NF
NF
1Both:
W=
0.0000083SL3.26
NF
1
48
NF
NF
NF
NF
NF
NF
NF
NF
AGE
&
GR
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
Page
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Kyphosidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
Q
R
S
T
U
V
W
X
Y
Z
AA
AB
to
(
SE)
Age
composition
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
NF
NF
NF
NF
NF
NF
NF
NF
1­
juvenile
growth
in
tidepools
1
>
10
yrs.
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
ROWTH
r2
Page
632
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Kyphosidae
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AC
AD
AE
AF
AG
AH
AI
AJ
AK
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
Female
NF
Oviparous1,2
NF
NF
NF
2,420
to
23
(
2
to
3)­
sex
unspecified
NG
(
NG)
NG
(
NG)

NF
Oviparous1
NF
NF
NF
NF
NF
NF
NF
NF
Oviparous1,2
NF
NF
NF
1,3,5
19
(
2)­
sex
unspecified
NG
(
NG)
NG
(
NG)

TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
1st
maturity
50%
maturity
Breeding
Behavior
Sexual
dimorphism
Page
633
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Kyphosidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AL
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
Fertilized
eggs
present
Spawning
females
present
Both
reprod
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.

NG
NG
NG
NF
California1,2,5,6
*****
*****

NF
NF
NF
NF
California1
NG
NG
NG
NF
California1,3,4,6
*****

REPRODUCTION
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
Page
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Kyphosidae
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BA
BB
BC
BD
BE
BF
BG
BH
BI
ductive
states
present
Spawning
Frequency
Size
specific
fecundity
equation
Size
at
settlement
Nov.
Dec.
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
Larval
duration
(
SL
in
mm)

NF
NF
NF
NF
NF
NF
1,3
25
225
to
30
4
21
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
DISPERSAL
&
RECRUITMEN
Mean
(
range)
fecundity
in
eggs/
g
female
wt.
Fecundity
Fecundity
range
in
#
eggs
(@

cm,
grams,
or
age)
Page
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BJ
BK
BL
BM
BN
BO
BP
BQ
BR
BS
Size
Fish
Size
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
(
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
NF
11.0
to
1.1
planktonic1,3,4
NF
1,23.0
to
17.0
NF
NF
NF
NF
NF
11.0
to
1.1
planktonic1
NF
1,2<
2.5
to
17.7
NF
NF
NF
NF
NF
11.0
to
1.1
planktonic1,2
NF
12.6
to
pelagic;
neuston1,2,3,4
NF
NF
NF
NF
2
3.0
to
13.8
pelagic;
nearshore
shallow
1,2,4
pelagic1,2,4,6/
warm
inshore
waters5
Eggs
Larvae
NT
Food
habits
Page
636
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BT
BU
BV
BW
BX
BY
BZ
Fish
Size
Food
habits
(
SL
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Habitat
2
17.0
to
maturity
omnivore6
scavenger1
NF
NF
at
night
in
crevices
and
holes14
117.7
to
omnivore
NF
NF
NF
invertebrates8
2
13.8
to
maturity
pelagic2,10,20/
kelp
1,6,8,20
NF
NF
NF
NF
(
red
and
green
algae;
small
crustaceans)
3,4
(
fleshy
bits
from
animals;
small
invertebrates;
algae)
1,2,4,7
pelagic2,10,17/
tide
pools;
shallow
rocky
intertidal;
surfgrass
beds
1,3,4,5,7,8,9,10,12,13,14,15,16,17,18,19,21
kelp
forest
inner
edges,
near
Egregia
laevigata,
and
kelp
canopy
;
nearshore
near
rocks;
shallow
reefs;
midwater;

artificial
reefs
1,2,3,4,6,7,8,10,11,12,13,14,16,17
,18,19,20,21,22,23,24,25
kelp
forest
outer
edges
and
canopy,

midkelp
rocky
areas;
surfgrass
beds;

artificial
reefs1,4,5,7,8,10,12,13,14,15,22,23,24,25
pelagic2/
inshore
and
in
tide
pools1,12,20
kelp
forest;
shallow
subtidal;
surfgrass
beds;
tide
pools
1,9,10,15,19,24
<
33
mm
(
more
carnivorous
than
herbivorous);
pelagic
juvenile
mostly
carnivorous;
become
more
herbivorous
as
they
grow5
Juveniles
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CA
CB
CC
CD
CE
CF
CG
Adults
Food
habits
Haplotype
diversity
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
(
Geographic
Area)

herbivore6,12,16,19,24,26/
omnivore8,10,17
browser1
diurnal1
seals
and
sea
lions1,2,3
NF
NF
NF
browser/
grazer2
avoids
corraline
algae10
microcarnivore20
herbivore10,16,19,26/
omnivore17
browser/
grazer2
NF
NF
NF
NF
NF
(
small
invertebrates)
17,21
herbivore:
red,
green,
brown
algae
25
herbivore16,19,26/
omnivore8,17
browser1
diurnal1,4
NF
NF
NF
browser/
grazer2
(
invertebrates)
17
meso/
microcarnivore20
(
small
crustaceans;
ophiuroids;
small
fish)
21,23
herbivore
25
(
Macrocystis,
coralline
reds,
and
other
seaweeds;
bryozoans;
polychaetes;
mussels;

gammerids)
1,2,3,5,10,11,13,14,15,16,17,18,21,22,23,24
bald
eagles,
least
terns;
and
Brandt's
comorants;
sea
lions4
(
red,
green
and
brown
seaweeds;
bryozoans;

sponges)
1,4,5,7,11,13,16,18,21,22,23
(
sheet­
like
red
[
Polysiphonia
spp.
and
others],

green
(
Ulva
),
and
brown
algae)
9,16,17,21
diurnal,
with
reduced
feedinga
t
night2,4
diurnal,
possibly
at
night2,3
(
small
crustaceans,
such
as
copepods,

caprellids,
and
amphipods;
bryozoans;

ophiuroids;
polychaetes)
21,23
sea
lions
and
fur
seals;
artic
loons,

rhinoceros
auklets,
least
terns,

Brandt's
cormorants,
and
bald
eagles4
Nucleotide
diversity
(#)

Page
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CH
CI
CJ
CK
CL
CM
CN
CO
CP
RECRUITMENT
INTO
Genetic
variance
Mean
number
of
%
Polymorphic
First
Recruitment
Full
Recruitment
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Total
length
(
age)
Total
length
(
age)
Sampling
gear
NF
NF
1,2NG
0.025
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1,2NG
0.025
allozymes
NF
NF
NF
NF
GENETICS
Mean
heterozygosity
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
FISHERY
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
DA
DB
DC
DD
DE
Interactions
/
associations
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

competes
with
Hypsypops
rubicundus
for
food1
schools
with
M.
californiensis
1
Residential1,2,3
Yes1
cleaned
by
Oxyjulis
califonica
12
NF
NF
Transient3
NF
competes
with
Hypsypops
rubicundus
for
food1
schools
with
G.
nigricans
1
NF
Residential2,3
NF
juveniles
clean
ectoparasites
from
fish
3
infected
by
trematodes
from
the
family
Capsalidae;
infestation
occurs
at
~
200
mm
SL
(
5
yrs.)
4,5
juveniles
move
towards
warmer
water
temperatures,
inshore
after
upwelling;
also
show
migrations
from
intertidal
pool
to
pool
until
finally
abandoning
the
intertidal
for
subtidal2
juveniles
demostrate
air­
breathing
capabilities
and
survival
out
of
water6
a
reservoir
host
for
an
infectious
sea
lion
calicivirus
7,9,15
juveniles
infected
and
killed
by
the
coccidium
Goussia
girellae
10
juveniles
sometimes
found
schooling
under
jellyfish14
infested
by
trematodes
from
the
family
Hemiuridae,
Lepocreadiinae,
Opecoelidae,

Haploporidae,
and
Haplosplanchnidae11
infested
by
parasites
from
the
families
Caligidae,
Pennellidae,
Haploporidae,

Haplosplanchnidae,
Hemiuridae,

Lepcreadiidae,
Opecoelidae,
Capsalidae,

Camallanidae,
Pseudaliidae,

andOphryoglenidae16
school
with
both
G.
nigricans
and
M.

californiensis
1,13,17
infected
by
ectoparasites
from
the
families
Caligidae
and
Lerneopodidae2
infected
by
ectoparasites
from
the
family
Lernaeidae;
also
monogenetic
trematodes2
tidally
controlled
vertical
movements1
BEHAVIOR
infested
by
parasites
from
the
families
Caligidae,
Pennellidae,
Hemiuridae,

Lepcreadiidae,
and
Waretrematidae16
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants8
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants;
juveniles
most
susceptible8
Movement
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20
21
22
23
24
25
26
27
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
DF
DG
Schooling
Territoriality
Yes,
especially
pelagic
juveniles1,5,12,13
NF
Yes4,6,9,10
Yes1,9,11,13
NF
Yes­
juveniles8
NF
Mostly
solitary2
loose
schools3,7,13
schooling
juveniles5
aggregate
in
shallow
portion
of
kelp­
bed
durng
mating
season1
Mostly
solitary,
but
sometimes
form
schools1,9
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61
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63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
A
B
C
D
E
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Love
et
al.
2000
1­
Love
et
al.
2000
2­
Gosline
1985
2­
Eschmeyer
1998
2­
Gates
&
Frey
1974
2­
Fritzsche
et
al.
1991
3­
Waugh
1991
3­
Roedel
1948
3­
Bond
1985
4­
Sumner
&
Fox
1933
Page
643
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­
Kyphosidae
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53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
F
G
H
I
J
K
LMNOP
1­
Quast
1968b
1­
Love
et
al.
2000
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­
Kyphosidae
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
Q
R
S
T
U
V
W
X
Y
Z
AA
AB
1­
Norris
1963
1­
Smith,
Skilling,
&
Brown
1980
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­
Kyphosidae
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53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
AC
AD
AE
AF
AG
AH
AI
AJ
AK
1­
Watson
1996c
1­
Limbaugh
1955
2­
Love
1996
2­
Fitch
&
Lavenberg
1971
3­
Fitch
&
Lavenberg
1975
4­
Feder
et
al.
1974
5­
Love
1996
Page
646
of
1167
5­
2220e
­
Kyphosidae
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59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
AL
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
1­
Watson
1996c
2­
Fitch
&
Lavenberg
1971
3­
Fitch
&
Lavenberg
1975
4­
Kramer
&
Smith
1973
5­
Norris
1963
6­
Love
1996
Page
647
of
1167
5­
2220e
­
Kyphosidae
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53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
BA
BB
BC
BD
BE
BF
BG
BH
BI
1­
Limbaugh
1955
2­
Williams
1957
3­
Stevens
et
al.
1989
4­
Norris
1963
Page
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53
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57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
BJ
BK
BL
BM
BN
BO
BP
BQ
BR
BS
1­
Watson
1996c
1­
Limbaugh
1955
1­
Watson
1996c
1­
Limbaugh
1955
2­
Fitch
&
Lavenberg
1975
2­
Stevens
et
al.
1989
2­
Watson
1996c
3­
Norris
1963
3­
Fitch
&
Lavenberg
1975
4­
Feder
et
al.
1974
4­
Stevens
et
al.
1989
5­
Gibson
1982
6­
Feder
et
al.
1974
Page
649
of
1167
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­
Kyphosidae
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53
54
55
56
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
BT
BU
BV
BW
BX
BY
BZ
1­
Watson
1996c
1­
Limbaugh
1955
1­
Limbaugh
1955
1­
Limbaugh
1955
1­
Limbaugh
1955
2­
Stevens
et
al.
1989
2­
Watson
1996c
2­
Stephens
1992
2­
Fitch
&
Lavenberg
1971
3­
Fitch
&
Lavenberg
1971
3­
Sturm
&
Horn
1998
3­
Larson
&
DeMartini
1984
4­
Martin
1995
4­
Barry
&
Ehret
1993
4­
Stephens
et
al.
1984
5­
Pondella
&
Stephens
1994
5­
Williams
&
Williams
1954
5­
Fitch
&
Lavenberg
1975
6­
Fitch
&
Lavenberg
1975
6­
Norris
1963
6­
CDF
&
G
1982
7­
CDF
&
G
1982
7­
Mitchell
1953
7­
Carlisle
et
al.
1964
8­
Stephens
1992
8­
Fidopiastis
1995
8­
Turner
et
al.
1969
9­
Williams
1957
9­
Sturm
&
Horn
1998
10­
Stevens
et
al.
1989
10­
Stevens
et
al.
1989
12­
Barry
&
Ehret
1993
11­
Orton
et
al.
1987
13­
Martin
1993
12­
Allen
1985
14­
DeMartini
1981
13­
Johnson
et
al.
1994
15­
Smith
et
al.
1981
14­
Ebeling
&
Bray
1976
16­
Williams
&
Williams
1954
15­
DeMartini
1981
17­
Norris
1963
16­
Williams
&
Williams
1954
18­
Barry
1988
17­
Hose
et
al.
1984
19­
Yoshiyama
et
al.
1986
18­
Goldberg
et
al.
1991
20­
Feder
et
al.
1974
19­
Norris
1963
21­
Love
1996
20­
Winget
1968
21­
Barry
1988
22­
Ebeling,
Larson,
&
Alevizon
1980
23­
Quast
1968a
24­
Feder
et
al.
1974
25­
Love
1996
Page
650
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1167
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­
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
CA
CB
CC
CD
CE
CF
CG
1­
Limbaugh
1955
1­
Stephens
1992
1­
Sikkel
1995
1­
Smith,
Skilling,
&
Brown
1980
2­
Robles
1987
2­
Horn
1989
2­
Hobson
et
al.
1981
2­
Smith
et
al
1981
3­
Fitch
&
Lavenberg
1971
3­
Ebeling
&
Bray
1976
3­
Hawes
1983
4­
Fitch
&
Lavenberg
1975
4­
Love
1996
4­
Love
1996
5­
Stephens
1992
6­
Sigalit
&
Rainboth
1992
7­
Carlisle
et
al.
1964
8­
Sikkel
1995a
9­
Sturm
&
Horn
1998
10­
Barry
&
Ehret
1993
11­
Hobson
et
al.
1981
12­
Sousa
1979
13­
Irelan
&
Horn
1991
14­
Norris
1963
15­
Holbrook
et
al.
1997
16­
Horn
1989
17­
Barry
1988
18­
Harris
et
al.
1984
19­
Hobson
1994
20­
Quast
1968a
21­
Quast
1968d
22­
Feder
et
al.
1974
23­
Love
1996
24­
Mitchell
1953
25­
Fidopiastis
1995
26­
Waugh
1991
Page
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53
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56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
CH
CI
CJ
CK
CL
CM
CN
CO
CP
1­
Waples
&
Rosenblatt
1987
2­
Waples
1987
Page
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59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
Page
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
DA
DB
DC
DD
DE
found
schooling
with
Racochilus
vacca
17
1­
Sikkel
1995
1­
Limbaugh
1955
1­
Williams
1957
1­
Fitch
&
Lavenberg
1971
1­
Williams
1957
2­
Hobson
1971
2­
Norris
1963
2­
Larson
&
DeMartini
1984
3­
Stephens
1992
3­
Stephens
et
al.
1994
4­
Hargis
1953
5­
Goldberg
et
al.
1991
6­
Martin
1993
7­
Smith
et
al.
1981
8­
Hose
et
al.
1983
9­
Smith,
Skilling,
&
Brown
1980
10­
Kent
et
al.
1988
11­
Martin
1978
12­
Limbaugh
19??

13­
Lockley
1952
14­
Norris
1963
15­
Smith,
Skilling,
Dardiri,
&
Latham
1980
16­
Love
&
Moser
1983
17­
Feder
et
al.
1974
Page
654
of
1167
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­
Kyphosidae
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53
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56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
DF
DG
1­
Limbaugh
1955
2­
Stephens
et
al.
1984
3­
Fitch
&
Lavenberg
1975
4­
CDF
&
G
1982
5­
Stephens
1992
6­
Williams
1957
7­
Carlisle
et
al
.1964
8­
Stevens
et
al.
1989
9­
Barry
&
Ehret
1993
10­
Williams
&
Williams
1954
11­
Lockley
1952
12­
Norris
1963
13­
Feder
et
al.
1974
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655
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­
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5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Labridae1
Halichoeres
semicinctus
1
Oxyjulis
californica
1,2
Semicossyphus
pulcher
1,2
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Love
et
al.
2000
1­
Love
et
al.
2000
2­
Eschmeyer
1998
2­
Gates
&
Frey
1974
3­
Roedel
1948
California
sheephead;
sheephead;
California
redfish;
fathead;
humpy;
redfish1,2,3
central
California
(
Diable
Cove)
to
southern
Baja
California
(
Isla
Asuncion)
1
northern
California
(
Salt
Point)
to
southern
Baja
California
(
Bahia
Magdalena)
1
central
California
(
Monterey
Bay)
to
lower
Gulf
of
California1
rock
wrasse;
California
wrasse;
parrot
fish;

wrasse1,2
señorita;
kelpfish;
pescerey;
kelp
wrasse;

butterfish
1,2,3
1intertidal
to
40
1intertidal
to
97
1intertidal
to
85
TAXONOMY
RANGE
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7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
F
G
H
I
J
K
L
MNO
P
Length
weight
equation
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
1*
Both:
W=
0.003412TL3.412
(
r2=
0.96)
1
38
NF
NF
NF
NF
NF
NF
NF
NF
2Both:
W=
0.00000128SL3.5
1
29.9
NF
NF
NF
NF
NF
NF
NF
NF
1*
Both:
W=
0.2148TL2.723
(
r2=
0.92)
1
91
2Both:
W=
0.0000131SL3.16
1­
DeMartini
et
al.
1994
1­
Love
et
al.
2000
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
AGE
&
GROWTH
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6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
Q
R
S
T
U
V
W
X
Y
Z
AA
to
(
SE)
Age
composition
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
NF
NF
NF
NF
NF
NF
NF
NF
NF
4
9
NF
6
14
NF
NF
NF
NF
NF
NF
NF
NF
NF
4
4
NF
1,2,4
1
29
3
(
juveniles)
2,5,6
53
320
to
25
at
San
Nicolas
I.

1­
Cowen
1990
1­
Limbaugh
1955
2­
Warner
1975
2­
Fitch
&
Lavenberg
1971
3­
Cowen
1991
3­
Cowen
1985
4­
DeMartini
et
al.
1994
4­
Fitch
&
Lavenberg
1975
5­
CDF
&
G
1982
6­
Love
1996
r2
Page
658
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1167
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14
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20
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22
23
24
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26
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30
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AB
AC
AD
AE
AF
AG
AH
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
vs.
Female
length
Secondary
Characteristics
Male
protogynous
hermaphrodite3,5
NF
Oviparous1
NF
NF
NF
1,6
NG
(
NG)

no
evidence
for
hermaphroditism4
4
NG
(
5)

some
individuals
are
hermaphrodites
6
NF
NF
Oviparous1,2
NF
NF
NF
1,4,6,714
(
1)­
unspecified
sex
protogynous
hermaphrodite1,2,5
Oviparous1
Males>
Females1,2
1,5
NG
(
NG)

2NG^
(
6)

2NG^
(
4)

2NG^
(
5
to
6)

2NG^
(
5)

3
25^
(
5)

317^
(
5)

6
30
(
7
to
8)

1­
Warner
1975
1­
Watson
1996d
1­
Warner
1975
1­
Warner
1975
1­
Limbaugh
1955
2­
Cowen
1990
2­
Love
1996
2­
Cowen
1990
2­
Crozier
1966
2­
Cowen
1990
3­
Fitch
&
Lavenberg
1975
3­
Warner
1975
4­
Stephens
et
al.
1994
4­
Fitch
&
Lavenberg
1975
5­
Feder
et
al.
1974
5­
Crozier
1966
6­
Love
1996
6­
Feder
et
al.
1974
7­
Love
1996
Sexual
dimorphism
males­
bump
on
forehead,
back
head
and
caudal
region;
females­
uniformaly
red2
local
population
demography
determines
when
sex
change
occurs2
1st
maturity
Females
salmon­
colored
with
white
chin;
most
males
have
black
anterior
and
posterior,
with
mid­
region
red
and
chin
white;
males
also
with
large
nuchal
hump
and
filamentous
extensions
of
median
fins1
Breeding
Behavior
TL,
FL*

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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AI
AJ
AK
AL
AM
AN
AO
AP
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
NG
(
2
to
3)
NG
(
NG)
NG
(
NG)
NG
NG
NG
California2,9
NG
(
2)
NG
(
NG)
NG
(
NG)
NG
NG
NG
NG
(
NG)
NG
(
NG)
NG
NG
NG
California1,2,3,5,6,7,8,9
25
to
28
(
NG)
NG
(
NG)
NG
(
NG)
NA
NA
NA
s.
California2,3,4,9
28
to
30
(
5)
48
(
13
to
14)
NG
(
6)
dorsal
spines
Marginal
increment
analysis
interreader
comparisons;
multiple
spine
comparison
NG
(
3
to
4)
20
to
22
(
5
to
6)
NG
(
3
to
4)
dorsal
spines
Marginal
increment
analysis
interreader
comparisons;
multiple
spine
comparison
NG
(
3)
26
to
28
(
6
to
7)
NG
(
4
to
5)
dorsal
spines
Marginal
increment
analysis
interreader
comparisons;
multiple
spine
comparison
NG
(
3
to
4)
26
to
28
(
6
to
7)
NG
(
3
to
4)
dorsal
spines
Marginal
increment
analysis
interreader
comparisons;
multiple
spine
comparison
19
(
4)
33
(
7)
21
(
4)
dorsal
spines
Marginal
increment
analysis
interreader
comparisons;
multiple
spine
comparison
14
(
4)
21
(
7)
15
(
4)
dorsal
spines
Marginal
increment
analysis
interreader
comparisons;
multiple
spine
comparison
20
(
4
to
5)
NG
(
NG)
NG
(
NG)
dorsal
spines
Marginal
increment
analysis
interreader
comparisons;
multiple
spine
comparison
1­
Limbaugh
1955
2­
Warner
1975
3­
Watson
1996d
4­
Cowen
1985
5­
Bolin
1930
6­
Breder
&
Rosen
1966
7­
Fitch
&
Lavenberg
1975
8­
Wang
1981
9­
Love
1996
50%
maturity
,
or
SL^
in
cm
(
age
in
yrs)
at
Fer
REPRODUCTION
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15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
states
present
Spawning
Frequency
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
(
per
year)

*****
NF
*****
*****
*****
*****
NF
*****
*****
*****
multiple1,2
1­
Warner
1975
2­
DeMartini
et
al.
1994
every
1.3
days
during
breeding
season;
86
batches
per
year
2
rtilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
Page
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BD
BE
BF
BG
BH
BI
BJ
Size
specific
fecundity
equation
Size
at
settlement
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
Larval
duration
(
SL
in
mm)
Annual
cohort
success
NF
NF
NF
NF
129.9
±
3.6
days
NF
NF
NF
NF
NF
1
48
hours
139.4
±
2.6
days
NF
NF
1ovary
wt.=
0.00131SL2.95
NF
137.4
±
5.1
days
112
to
13
NF
266.5
days
213.5
(
12.7
to
16)

215
±
8.7
(
NG)
fecundity
increases
with
size1
360
(
37
to
78)
days
2
5,755
±
3,577
(
NG)

1­
Warner
1975
1­
Cowen
1990
1­
Warner
1975
1­
Bolin
1930
1­
Victor
1986
1­
Cowen
1985
2­
DeMartini
et
al.
1994
2­
Warner
1975
2­
Cowen
1985
2­
Cowen
1991
3­
Cowen
1991
Fecundity
Fecundity
range
in
#
eggs
(@

cm,
grams,
or
age)

Mean
(
range)
fecundity
in
eggs/
g
female
wt.
36,068
(
20.9
SL)
to
258,000
(
35.9
SL)
yolky
oocytes2
1
NG
(
4,450
to
6,200)
yolky
oocytes
per
g
ovary
wt.
DISPERSAL
&
RECRUITMENT
Page
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BK
BL
BM
BN
BO
BP
BQ
BR
BS
BT
Size
Fish
Size
Fish
Size
(
diameter
in
mm)
Habitat
Predators
(
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
(
SL
mm)

NF
planktonic1,2
11.5
to
2.0
to
21.9
planktonic1
1,2,30.74
to
0.79
planktonic1,2,4,5,6
12.0
to
18.6
planktonic1
4
0.75
2
2.1
to
2.6
to
(
TL)

1
0.8
planktonic1,2,3,5
12.5
to
15.6
planktonic1
planktonic,
midwater2
1­
Watson
1996d
1­
Limbaugh
1955
1­
Watson
1996d
1­
Watson
1996d
2­
Bolin
1930
2­
Watson
1996d
2­
Wang
1981
2­
Cowen
1985
3­
Breder
&
Rosen
1966
3­
Cowen
1985
4­
Morris
1956a
4­
Fitch
&
Laveneberg
1975
5­
Feder
et
al.
1974
6­
Love
1996
Food
habits
Eggs
Larvae
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BU
BV
BW
BX
BY
BZ
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Habitat
NF
NF
NF
NF
surfgrass
beds5
carnivore/
planktivorous3
NF
diurnal1
NF
at
night
they
burrow
into
sand
or
mud1,7
surf
and
eelgrass
beds;
bays5,7
gorgonian
beds1,6,7
NF
NF
NF
moray
eels1
surfgrass
beds5
sleeps
in
rock
crevices
and
holes
at
night
2,8,10,18,22,27
1­
Limbaugh
1955
1­
Tegner
&
Dayton
1981
1­
Tegner
&
Dayton
1981
1­
Wiley
1974
1­
Limbaugh
1955
2­
Coyer
1979
2­
Hartney
1996
2­
Wiley
1974
3­
Turner
et
al.
1969
3­
Hobson
&
Chess
1976
3­
Larson
&
DeMartini
1984
4­
Hobson
&
Chess
1976
4­
Love
1996
4­
Hobson
1971
5­
DeMartini
1981
5­
Stephens
et
al.
1984
6­
Crozier
1966
6­
Hobson
&
Chess
1986
7­
Feder
et
al.
1974
7­
Fitch
&
Lavenberg
1975
8­
Love
1996
8­
CA
Dept
of
Fish
&
Game
1982
9­
Carlisle
et
al.
1964
10­
Turner
et
al.
1969
burrow
into
the
sand
at
night
and
sleep
1,10,12,13,16,18,20,27,28
(
larval
sea
urchins;
calanoid
and
cyclopoid
copepods;

bryzoan
larvae;
mollusk
larvae;
fish
eggs)
1,2,3,4
kelp
forest;
surfgrass
beds;
rocky
bottoms;
forage
over
sandy
bottoms;
high­
and
low­
relief;
artificial
reefs
1,3,5,6,9,10,14,15,17,18,22,24,26,27,36
flat
sandy­
rocky
areas;
rocky
reefs;

disappear
into
sand
when
frightened
1,3,7
kelp
forest,
throughout
water
column;
rocky
areas;

surfgrass
beds;
artificial
reefs;
soft
substrata
1,3,4,5,9,10,14,15,16,18,19,20,22,23,24,25,26,27,28
kelp
forest;
rocky
areas;
disappear
into
bottom
cover
when
frightened1,2,4,7,8
rocky
and
sandy
bottom
of
kelp
forest;
burrow
between
rocks
at
night;
surf
grass
beds;
artificial
reefs;
tide
pools1,3,5,6,7,9,10,11,13,14,17,21,24,26,27,28
Juveniles
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Page
664
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11
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13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CA
CB
CC
CD
CE
CF
Adults
Food
habits
Haplotype
diversity
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
(
Geographic
Area)

carnivore/
microcarnivore19
picker1,4
diurnal1,6,8,9,10,15
NF
NF
NF
grazer5
scavange
sea
urchins
form
sheephead
attacks
3
omnivore/
microcarnivore18,23/
planktivore20/
mesocarnivore32
picker2,3,6,7
diurnal5,10,11,12,13,14,15
NF
NF
Caspian
terns7
sea
lions9,10
scavange
sea
urchins
form
sheephead
attacks
3
carnivore/
microcarnivore19,23
crusher1
diurnal1,6,7,10,12,15
giant
sea
bass2,8
NF
NF
crusher/
winnower4
strictly
diurnal2,4
1­
Limbaugh
1955
1­
Limbaugh
1955
1­
Limbaugh
1955
1­
Limbaugh
1955
2­
Diaz
&
Hammann
1987
2­
Tegner
&
Dayton
1981
2­
Nelson
&
Vance
1979
2­
Fitch
&
Lavenberg
1971
3­
Nelson
&
Vance
1979
3­
Hobson
1971
3­
Cowen
1983
3­
Hobson
1971
4­
Robles
&
Robb
1993
4­
Hobson
&
Chess
1986
4­
Vance
&
Schmitt
1979
4­
Fitch
&
Lavenberg
1975
5­
Tegner
&
Dayton
1981
5­
Stephens
et
al.
1994
5­
Hobson
1971
5­
Burge
&
Schultz
1973
6­
Robles
1987
6­
Hartney
1996
6­
Hobson
&
Chess
1986
6­
Hubbs
et
al.
1970
7­
Cowen
1983
7­
Bray
&
Ebeling
1975
7­
CA
Dept
of
Fish
&
Game
1
7­
Harvey
1982
8­
Fitch
&
Lavenberg
1971
8­
Stephens
et
al.
1994
8­
Fitch
1960
9­
Turner
&
Ebert
1963
9­
Sikkel
1995
9­
Hawes
1983
10­
Vance
&
Schmitt
1979
10­
Hobson
et
al.
1981
10­
Love
1996
individuals
concentrate
on
one
type
of
prey,
but
populations
show
wide
variety
of
prey
taken31
(
small
crustaceans,
like
crabs
and
gammerids;
bryozoans;

mussels
and
gastropods;
sea
urchins;

polychaetes)
1,2,4,5,6,14,15,17,22,27,28,33,34,36
birds,
such
as
cormorants1,4,5,6,10
(
larval
fish;
plant
debris;
external
fish
parasites,
such
as
caligids;
squid,
mussels,
and
gastropods;
bryozoans;

crustanceans,
like
crabs,
caprellids,
and
gammerids;
small
sea
urchins;
fish
eggs,
such
as
blacksmith
eggs)
1,2,4,5,9,13,15,18,21,24,25,28,31,33,34,36
diurnal,
with
peak
in
the
morning3
bocaccio
and
starry
rockfish3
(
sea
urchins,
ophiuroids
and
other
echinoderms;
mussels
and
gastropods;
crabs,
such
as
decorator
crabs,
spiny
lobster,
and
barnacles;
squid;
bryozoans;
tubiculous
polychaetes;
fish
eggs,
such
as
blacksmith
eggs)

1,2,3,4,5,6,7,8,9,10,11,12,14,16,22,25,26,29,30,33,34,35,36,37
Page
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CG
CH
CI
CJ
CK
CL
CM
CN
CO
Genetic
variance
Mean
number
of
%
Polymorphic
First
Recruitment
Full
Recruitment
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Total
length
(
age)
Total
length
(
age)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1,2NG
0.033
allozymes
NF
NF
NF
1­
Waples
&
Rosenblatt
1987
2­
Wales
1987
GENETICS
Nucleotide
diversity
(#)
Mean
heterozygosity
Page
666
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CP
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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46
47
48
49
50
51
52
53
DA
DB
DC
DD
DE
Interactions
/
associations
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

competes
with
Hypsypops
rubicundus
for
food1
No
parasites
found
on
this
species4
NF
Residential1,4,5
NF
NF
NF
Residential1,2,4,6
Yes1
competes
with
Hypsypops
rubicundus
for
food1
NF
Residential1,3,4,6
NF
1­
Sikkel
1995
1­
Limbaugh
1955
1­
Larson
&
DeMartini
1984
1­
Hartney
1996
2­
Cowen
1983
2­
Hobson
1971
3­
Wiley
1974
3­
CDF
&
G
1982
4­
Hobson
1971
4­
Stephens
et
al.
1994
5­
Sigalit
&
Rainboth
1992
5­
Thomson
&
Lehner
1976
6­
Carlisle
et
al.
1964
6­
Helvey
1985
7­
Turner
et
al.
1969
8­
Hartney
1996
9­
Bray
&
Ebeling
1975
10­
Gotshall
1966
BEHAVIOR
Movement
infested
by
parasites
from
the
families
Caligidae
and
Ceratomyxidae14
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants11
infested
by
parasites
from
the
families
Caligidae,

Gorgoderidae,
Lepcreadiidae,
Gnathiid
larvae,
and
Capsalidae14
Cleans
external
parasites
from
pomacentrids
like
blacksmiths
and
garabaldis,
sea
basses,

embiotocids,
topsmelt,
bat
rays,
surfperches,

kyphosids,
mola,
other
senoritas1,4,5,6,7,8,9,10,12,15,16
infected
by
ectoparasites
from
the
families
Caligidae
and
Chondracanthidae4
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
5
Has
strong
effect
on
the
density
and
spatial
distribution
of
Strongylocentrotus
franciscanus
at
San
Nicolas
I.
2
enclosure
in
mucus
envelope
at
night
may
help
avoid
predation3
infected
by
ectoparasites
from
the
families
Caligidae;
also
gnathiid
larvae
and
monogenetic
trematodes4
infested
by
parasites
from
the
families
Caligidae
and
Chondracanthidae14
may
destroy
small
Macrocystis
beds
when
feeding
on
the
bryozoan
Membranipora
13
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants11
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49
50
51
52
53
DF
DG
Schooling
Territoriality
No1,3,9,10
Yes6
Yes1,2,4,5,7,9
No1,3,8,9
No1
1­
Limbaugh
1955
1­
CDFG
1982
2­
Hobson
1971
3­
Hobson
&
Chess
1986
4­
Burge
&
Schultz
1973
5­
Turner
et
al.
1969
6­
Stephens
et
al.
1994
7­
Hobson
&
Chess
1976
8­
Crozier
1966
9­
Feder
et
al.
1974
10­
Love
1996
Yes,
but
solitary
and
paired
individuals
also
found
10
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78
79
80
A
B
C
D
E
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68
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76
77
78
79
80
F
G
H
I
J
K
L
MNO
P
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76
77
78
79
80
Q
R
S
T
U
V
W
X
Y
Z
AA
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63
64
65
66
67
68
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70
71
72
73
74
75
76
77
78
79
80
AB
AC
AD
AE
AF
AG
AH
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64
65
66
67
68
69
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71
72
73
74
75
76
77
78
79
80
AI
AJ
AK
AL
AM
AN
AO
AP
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63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
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64
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66
67
68
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70
71
72
73
74
75
76
77
78
79
80
BD
BE
BF
BG
BH
BI
BJ
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64
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66
67
68
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70
71
72
73
74
75
76
77
78
79
80
BK
BL
BM
BN
BO
BP
BQ
BR
BS
BT
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64
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
BU
BV
BW
BX
BY
BZ
11­
Stephens
et
al.
1994
12­
Hartney
1996
13­
Hobson
et
al.
1981
14­
Allen
1985
15­
Ebeling,
Larson,
Alevizon,
&
Bray
1980
16­
Bray
&
Ebeling
1975
17­
Johnson
et
al.
1994
18­
Ebeling
&
Bray
1976
19­
Bodkin
1986
20­
Hobson
&
Chess
1976
21­
DeMartini
1981
22­
Crozier
1966
23­
Love
et
al.
1986
24­
Ebeling,
Larson,
&
Alevizon
1980
25­
Bernstein
&
Jung
1979
26­
Quast
1968a
27­
Feder
et
al.
1974
28­
Love
1996
Page
678
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Labridae
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
CA
CB
CC
CD
CE
CF
11­
Cowen
1986
11­
Bray
&
Ebeling
1975
12­
Wicksten
1978
12­
Ebeling
&
Bray
1976
13­
Hobson
1971
13­
Hobson
&
Chess
1976
14­
Hobson
&
Chess
1986
14­
Bernstein
&
Jung
1979
15­
Fitch
&
Lavenberg
1975
15­
Love
1996
16­
Turner
et
al.
1969
17­
Thomson
&
Lehner
1976
18­
Hartney
1996
19­
Sikkel
1995a
20­
Helvey
1985
21­
Bray
&
Ebeling
1975
22­
Johnson
et
al.
1994
23­
Ebeling
&
Bray
1976
24­
Hobson
&
Chess
1976
25­
Tegner
&
Dayton
1977
26­
Tegner
1980
27­
Norris
1963
28­
Holbrook
et
al.
1997
29­
Crozier
1966
30­
Winget
1968
31­
Bernstein
&
Jung
1979
32­
Quast
1968a
33­
Quast
1968d
34­
Feder
et
al.
1974
35­
Allen
1916
36­
Love
1996
37­
Mitchell
1953
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64
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
CG
CH
CI
CJ
CK
CL
CM
CN
CO
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
CP
CQ
CR
CS
CT
CU
CV
CW
CX
CY
CZ
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65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
DA
DB
DC
DD
DE
11­
Hose
et
al.
1983
12­
Limbaugh
19??

13­
Bernstein
&
Jung
1979
14­
Love
&
Moser
1983
15­
Feder
et
al.
1974
16­
Limbaugh
1956
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68
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71
72
73
74
75
76
77
78
79
80
DF
DG
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
A
B
C
D
E
F
Total
Length
(
cm)

Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)
to
weight
(
g)
equation
Malacanthidae1,2
Caulolatilus
princeps
1,2
ocean
whitefish
1shallow
water
to
136
m
NF
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Eschmeyer
and
Herald
1983
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
3­
Hart
1973
4­
Love
1996
TAXONOMY
RANGE
Vancouver
Island,
British
Columbia
to
Peru,
including
Gulf
of
California
and
Islas
Galapagos1
blanquillo,
whitefish,
blanka,
pez
blanco
1,2,3,4
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
G
H
I
J
K
L
M
N
O
P
Q
R
Max.
reported
Geographic
area
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
Female
Both
Male
Female
Both
Male
Female
Both
NF
1102
cm
TL
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Love
et
al
2000
Linf
in
cm
(
SE)
AGE
and
GROWTH
k
(
SE)
to
(
SE)
Growth
P
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33
34
35
36
37
38
39
40
41
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43
44
45
46
47
48
49
50
51
52
53
54
S
T
U
V
W
X
Y
Z
AA
r2
Age
composition
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
Length)
information
NF
NF
NF
NF
NF
NF
NF
1,213
yrs.
NF
313
yrs
(
50­
53
cm)

1­
Love
1996
2­
Fitch
and
Lavenberg
1971
3­
Kato
1991
Parameters
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20
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22
23
24
25
26
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
AB
AC
AD
AE
AF
AG
AH
AI
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
dioecious
promiscuous
oviparous1
No
NF
NF
148­
56
cm
(
4­
5
yrs)
41­
48
cm
(
3­
4
yrs)

1­
Moser
1996a
1­
Love
1996
REPRODUCTION
Total
length
in
cm
Breeding
Behavior
Sexual
dimorphism
1st
maturity
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
AJ
AK
AL
AM
AN
AO
AP
AQ
AR
AS
Fertilized
eggs
present
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
NG
NG
NG
NG
NG
NF
CalCOFI
area1
California
current
region4
California2,3,5
Bay
of
La
Paz,
Mexico
1­
Moser
1996a
2­
Fitch
and
Lavenberg
1971
3­
Dooley
1978
4­
Kramer
and
Smith
1973
5­
CDFG
1982
50%
maturity
m
(
age
in
yrs)
at
Fertilization
and
Spawning
Period
(^^^

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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
Mean
Fecundity
t
Spawning
females
present
Both
reproductive
states
present
eggs/
g
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
(
weight
range)

multiple
spawners1
NF
*****
1­
Elorduy­
Garay
and
Ramirez­
Luna
1994
^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BE
BF
BG
BH
BI
BJ
Fecundity
Size
specific
Number
of
eggs
fecundity
eq
Size
at
settlement
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
Larval
duration
&
location
Annual
cohort
success
NF
NF
NF
few
months
?
1
NF
NF
1­
Waples
and
Rosenblatt
1987
DISPERSAL
&
RECRUITMENT
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BK
BL
BM
BN
BO
BP
BQ
BR
Size
(
diameter
in
mm)
Habitat
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
NF
eggs
planktonic1,2,3
NF
1<
2.6­
7.0
mm
Sl
planktonic1,2
NF
NF
NF
inshore/
offshore4
1­
Moser
1996a
1­
Moser
1996a
1­
Moser
1996a
2­
C.
D.
F.
G.
1982
2­
Dooley
1978
3­
Feder
et
al
1974
3­
Moser
et
al
1986
4­
Waples
and
Rosenblatt
1987
5­
Kato
1991
Food
habits
Larvae
Eggs
coastal
waters
to
>
200
miles
offshore3
distribution
of
larvae
indicates
populations
off
s.
California
are
recruited
from
Baja,
California3,5
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22
23
24
25
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BS
BT
BU
BV
BW
BX
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
albacore1
NF
NF
NF
NF
NF
1­
Feder
et
al
1974
Food
habits
Juveniles
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46
47
48
49
50
51
52
53
54
BY
BZ
CA
CB
Predators
Habitat
Foraging
guild
Foraging
method
usually
offshore
rocky
reefs
and
around
islands1
carnivores
1­
Kato
1991
1­
Moser
1996a
1­
Fitch
and
Lavenberg
1971
1­
C.
D.
F.
G.
1982
2­
Love
1996
2­
Dooley
1978
2­
Elorduy­
Garay
and
Caraveo­
Patino
1994
3­
Fitch
and
Lavenberg
1971
3­
Elorduy­
Garay
and
Caraveo­
Patino
1994
4­
Limbaugh
1955
4­
C.
D.
F.
G.
1982
5­
Feder
et
al
1974
5­
Feder
et
al
1974
6­
Elorduy­
Garay
and
Caraveo­
Patino
1994
6­
Kato
1991
demersal
fish,
inhabits
the
other
continental
shelf
and
upper
slope,
normally
occurs
on
rocky
bottoms6
waits
for
the
approach
of
zooplankton
masses
or
feeds
on
organisms
from
trophic
levels
close
to
primary
producers
2
observed
dropping
to
the
bottom
to
feed
1
(
crabs,
shrimp,
other
crustaceans,
octopus,
squid
and
fishes)
4,5
Food
habits
Adults
found
in
midkelp
and
rocky­
bottom
kelp­
bed
habitat4
larger
fishes,
including
albacore
tuna1
passive
predatory
teleost,
feeds
maily
on
organisms
associated
with
the
seafloor2
(
shrimps,
pelagic
red
crabs,
hermit
crabs
euphausiids,
small
octupi,
squid,
anchovies,
and
blue
laternfish)
1,2,6
found
in
aggregations
over
rocks,
kelp,
or
other
high
relief2,3,4,5
opportunistic
omnivore
(
ostracods,
hyperiid
amphipods,
euphausiids,
galatheids,
pteropod
molluscs,
cephalopods,
and
fishes)
3
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31
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34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CC
CD
CE
CF
CG
Haplotype
diversity
When
/
Frequency
Predators
Substocks
(
Geographic
Area)

diurnal
feeder1
NF
NF
1­
Elorduy­
Garay
and
Caraveo­
Patino
1994
1­
Fitch
and
Lavenberg
1971
1­
Waples
and
Rosenblatt
1987
2­
Kato
1991
1
No
substocks
found
between
La
Jolla,

CA,
Channel
Islands,
CA,
Punta
Eugenia,
CA,
and
Isla
de
Guadalupe,

CA
using
allozymes
giant
sea
bass,
sharks,
and
other
large
predators1,2
Nucleotide
diversity
(#)

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22
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31
32
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34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CH
CI
CJ
CK
CL
CM
CN
Genetic
variance
Mean
number
of
%
Polymorphic
First
Recruitment
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Length
(
age)

NF
NF
NF
NF
NF
NF
NF
GENETICS
Mean
heterozygosity
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31
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34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CO
CP
CQ
CR
CS
CT
CU
CV
CW
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
Full
Recruitment
Method
of
Fishery
/
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
NF
N
F
NF
Nf
NF
NF
NF
NF
NF
MORTALITY
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CX
CY
CZ
DA
DB
DC
DD
Rate
of
Exploitation
Age
range
used
Rate
of
Exploitation
Interactions
/
associations
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
NF
NF
NF
NF
parasites:
4
families,
4
spp.
(
see
reference)
1
NF
NF
1­
Love
and
Moser
1983
BEHAVIOR
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34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
DE
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Homing
ability
(
Home
range)
Schooling
Territoriality
NF
NF
1­
Limbaugh
1955
2­
Feder
et
al
1974
loosely
gregarious
and
wander
almost
continuously
4­
15
feet
above
the
substrate,
rarely
dropping
to
the
bottom
to
1
2
Movement
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72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
A
B
C
D
E
F
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70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
G
H
I
J
K
L
M
N
O
P
Q
R
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64
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66
67
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
S
T
U
V
W
X
Y
Z
AA
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67
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
AB
AC
AD
AE
AF
AG
AH
AI
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64
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
AJ
AK
AL
AM
AN
AO
AP
AQ
AR
AS
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66
67
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
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64
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66
67
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
BE
BF
BG
BH
BI
BJ
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
BK
BL
BM
BN
BO
BP
BQ
BR
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66
67
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
BS
BT
BU
BV
BW
BX
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67
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
BY
BZ
CA
CB
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64
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
CC
CD
CE
CF
CG
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64
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
CH
CI
CJ
CK
CL
CM
CN
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
CO
CP
CQ
CR
CS
CT
CU
CV
CW
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
CX
CY
CZ
DA
DB
DC
DD
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
DE
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
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33
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38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Mugilidae1,2
Mugil
cephalus
1,2
striped
mullet
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Eschmeyer
and
Herald
1983
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
3­
Eggold
and
Motta
1991
4­
Moyle
1976
5­
Thomson
1963
mullet,
grey
mullet,
black
mullet,

jumping
mullet,
sea
mullet,

tainhas,
chi
yue,
muge
cephale,
cefalo,
ora,
minghach,

aguas,
laban,
haarder,
loban,
ca
doi
1,2,3,4,5
circumglobal;
San
Francisco
Bay,
Northern
California
to
Chile
and
Islas
Galapagos
1
1
shallow
waters
of
marine
and
estuaries
to
122m
TAXONOMY
RANGE
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31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
F
G
H
I
JKL
M
NO
Total
Length
(
cm)
Max.
reported
Geographic
area
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
1W=
0.00808L3.025
NF
1
91
cm
TL
western
Australia1
NG
NG
68.869
NG
NG
­
0.21962
1W=
0.02817L2.9755
eastern
Australia2
NG
NG
60.46
NG
NG
0.344
2W=
0.0000688L3.2754
2W=
0.0000722L3.2746
3Both:
W=
0.000020L2.943
3Female:
W=
0.000065L2.737
3Male:
W=
0.000082L2.694
3Juveniles:
W=
0.000020L2.946
4W=
0.0000688L3.27541(
zooplankton
feeder)

4W=
0.0000287L3.553191(
mixed
food
feeder)

4W=
0.0000722L3.27455(
vegetable
feeder)

1­
Thomson
1973
1­
Love
et
al
2000
1­
Thomson
1963
2­
De
Silva
1980
2­
Grant
and
Spain
1975
3­
Pafford
1983
4­
Suzuki
1965
AGE
and
Linf
in
cm
(
SE)
k
(
SE)

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41
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43
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45
46
47
48
49
50
51
52
53
P
Q
R
S
T
U
V
W
X
Y
Z
r2
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)

NG
NG
0.0332
NG
NG
NG
NG
NG
NG
1,2,3,4,7,8,9,10,11
27
yrs
(
60.46
cm)

NG
NG
0.14
NG
NG
NG
scales/
read
annual
marksNo
No
111­
21
yrs
313­
14
yrs
1­
Radtke
1984
1­
Thomson
1963
2­
Moyle
1976
2­
Grant
and
Spain
1975
3­
Hendricks
1961
3­
Thomson
1966
4­
Thomson
1963
5­
De
Silva
1980
6­
Walker
1961
7­
Thomson
1966
8­
Anderson
1958
9­
Pafford
1983
10­
Jacot
1920
11­
Broadhead
1958
Growth
Parameters
d
GROWTH
to
(
SE)
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AA
AB
AC
AD
AE
AF
AG
AH
Age
composition
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
1
dioecious1,2
polyandrous1,3,4
oviparous1
No
females
larger1,2,3,4,5
sexually
dimorphic
color1
123­
35
cm
SL
(
2­
3
yrs.)

promiscous2
223­
29
cm
(
3
yrs)

223.6
cm
(
2
yrs)

233
cm
(
1
yr.)

233
cm
(
2
yr.)

230­
34
cm
(
3
yrs.)

231­
35
cm
(
3
yrs.)

240
cm
(
5
yrs.)

231­
37
cm
(
6­
8
yrs.)

323.5
cm
(
2
yrs.)

426.3
cm
(
3
yrs.)

5(
3
yrs.)

632
cm
(
3
yrs.)

1­
Thomson
1963
1­
Thomson
1963
1­
Breder
and
Rosen
1966
1­
Moser
1996a
1­
Fitch
and
Lavenberg
1971
1­
Breder
and
Rosen
1966
1­
Moyle
1976
2­
Thomson
1966
2­
Thomson
1963
2­
Breder
and
Rosen
1966
2­
Breder
1940
2­
Thomson
1963
3­
Thomson
1966
3­
Breder
1940
3­
Anderson
1958
4­
Breder
1940
4­
Fitch
and
Lavenberg
1975
4­
Pafford
1983
5­
Pafford
1983
5­
Broadhead
1958
6­
Thomson
1957
some
indivicuals
may
be
hermaphrodites1,2
males
have
a
pearly
sheen
overlying
lengthwise
striping,

which
females
lack2
Sexual
dimorphism
1st
ma
REPRODUCT
Breeding
Behavior
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24
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AI
AJ
AK
AL
AM
AN
AO
AP
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
23­
35
cm
SL
(
2­
3
yrs.)
NG
NG
NG
NG
NG
1
eastern
Pacific1,4
24­
31
(
3
yrs.)
NG
NG
scales
NG
NG
North
Carolina3
25.5
(
3
yrs.)
NG
NG
NG
NG
NG
Florida1,2,3,6
35
cm
(
1
yr.)
NG
NG
scales
NG
NG
Texas3
35
cm
(
2
yr.)
NG
NG
scales
NG
NG
Sea
of
Marmora1
30­
34
cm
(
3
yrs.)
NG
NG
scales
NG
NG
Sicily1
31­
35
cm
(
3
yrs.)
NG
NG
scales
NG
NG
Black
Sea1
41.5
cm
(
5
yrs,)
NG
NG
otoliths/
NG
NG
NG
Southern
India1
31­
37
cm(
6­
8
yrs.)
NG
NG
NG
NG
NG
east
Australia1,5
23.5
cm
(
2
yrs.)
NG
NG
NG
NG
NG
west
Australia1,7
25.8
cm
(
3
yrs.)
NG
NG
scales
and
otoliths
NG
structure
comparison
Hong
Kong1
(
3
yrs)
NG
NG
NG
NG
NG
35
cm
(
3
yrs.)
NG
NG
scales
NG
NG
1­
Thomson
1966
1­
Fitch
and
Lavenberg
1971
2­
Breder
1940
3­
Anderson
1958
4­
Fitch
and
Lavenberg
1975
5­
Grant
and
Spain
1975
6­
Broadhead
1958
7­
Thomson
1957
Fer
aturity
50%
maturity
Total
length
(
age)
at
ION
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22
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24
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31
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33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
states
present
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
1single
spawner
*****
*****
*****
*****
1­
Thomson
1963
2­
Grant
and
Spain
1975
rtilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
may
spawn
in
alternate
years
after
maturity1,2
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BD
BE
BF
BG
BH
BI
BJ
Mean
Fecundity
Size
specific
eggs/
g
Number
of
eggs
fecundity
eq
Size
at
settlement
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
Larval
duration
&
location
Annual
cohort
success
1,2,51,275,000­
2,781,000
eggs
11,320,000
eggs
(
50
cm)
1F=
0.009L3.16
1,22
days
1approx.
3­
4
weeks
NF
NF
12,000,000­
2,500,000
eggs
23,600,000­
7,200,000
eggs
3300,000­
1,600,000
eggs
41,572,000­
4,774,000
eggs
1­
Thomson
1963
1­
Thomson
1963
1­
Grant
and
Spain
1975
1­
Thomson
1963
1­
Anderson
1958
2­
Thomson
1966
2­
Thomson
1966
3­
Pafford
1983
4­
Grant
and
Spain
1975
5­
Thomson
1957
Fecundity
DISPERSAL
&
RECRUITMENT
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22
23
24
25
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BK
BL
BM
BN
BO
BP
BQ
Eggs
Size
(
diameter
in
mm)
Habitat
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
10.74­
0.82
mm
planktonic1,6
NF
12.2­
12.0
mm
SL
planktonic1,3
planktivore1
NF
20.65­
0.78
mm
spawned
offshore
over
deep
water2,3,5,6,9
22.0­
20.0
mm
20.91­
1.08
mm
spawning
takes
place
in
schools4
20.72
mm
(
avg.)

20.60­
0.72
mm
20.65­
0.85
mm
1­
Moser
1996a
1­
Moser
1996a
1­
Moser
1996a
1­
Moser
1996a
1­
Thomson
1963
2­
Thomson
1966
2­
Fitch
and
Lavenberg
1971
2­
Thomson
1966
2­
Thomson
1963
3­
Arnold
and
Thomson
1958
3­
Duffy
1987
4­
Moyle
1976
4­
Johnson
and
McClendon
1970
5­
Thomson
1963
5­
Anderson
1958
6­
Duffy
1987
6­
Powles
1981
7­
Anderson
1958
8­
Hu
1981
9­
Fitch
and
Lavenberg
1975
10­
Grant
and
Spain
1975
Larvae
M.
cephalus
unable
to
ovulate
in
freshwater8
remain
at
sea
until
they
are
about
18­

28
mm,
at
which
time
they
move
to
the
coast5
larvae
drift
from
spawning
grounds
with
prevailing
currents,
until
large
enough
to
swim
against
tidal
streams,

enter
estuaries
between
20­
30
mm2
found
in
offshore
waters,
move
shoreward
with
increased
size3,6
spawning
occurs
at
sea,
ranging
from
close
to
shore
to
20
miles
at
sea
7
optimal
salinity
for
eggs
at
22­
24
C
is
22­
23 
8
Food
habits
may
spawn
in
the
inshore
shallows,

possible
in
estuaries10
some
found
in
freshwater,
indicating
a
freshwater
spawning4
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24
25
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BR
BS
BT
BU
BV
When
/
Frequency
Predators
Fish
Size
Habitat
Foraging
guild
NF
121
cm
to
maturity
planktivore1
223
mm
to
maturity
(
microcrustacean
and
zooplankton)
2
dwell
primarily
in
shallow
estuaries2,3,4,5,6
(
microinvertebrates
and
plant
material)
5
1­
Fitch
and
Lavenberg
1971
1­
Moyle
1976
1­
Fitch
and
Lavenberg
1971
1­
Thomson
1963
2­
Fitch
and
Lavenberg1975
2­
Jacot
1920
2­
Moyle
1976
2­
De
Silva
1980
3­
Thomson
1963
3­
De
Silva
and
Wijeyaratne
1977
4­
Thomson
1966
4­
Thomson
1966
5­
Powles
1981
5­
Horn
and
Allen
1985
6­
Suzuki
1965
6­
Hiatt
1944
7­
Odum
1973
8­
Pafford
1983
9­
Suzuki
1965
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Juveniles
carnivorous
(
mosquito
larvae,
copepods,
and
other
zooplankton)
7
(
planktonic
micro­
crustceans,
diatoms,
and
other
micro­
algae,
sandy
muds
and
organic
(
diatoms,
blue­
green
algae,
and
other
plant
material)
6
(
detritus
and
associated
microflora,
epiphytes
and
filamentous
algae)
8
(
planktonic
and
benthic
forms
of
amphipods
and
copepods)
4
surface
feeding
predators,
including
tropical
tuna
1,2
(
primarily
diatoms,
green
algae,
desmids,
and
blue­
green
algae;
no
detritus
found
in
fish
<
25
mm,
amount
of
detritus/
sand
increased
with
increasing
body
length)
3
surface
waters
offshore,
move
inshore
as
they
increase
in
size1
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BW
BX
BY
BZ
CA
Foraging
method
When
/
Frequency
Predators
Habitat
Foraging
guild
primarily
bottom
feeders1
diurnal
feeders1
omnivore/
bottom
feeder4
feeds
in
water
column2
catadromous
fish3,6,8
migrate
to
salt
water
to
spawn4,5
concentrate
around
freshwater
inlets7
euryhaline9
iliophagous
and
herbivorous
diet6
(
diatoms,
blue­
green
algae,
and
other
plant
material)

(
benthic
microplant
material
and
microplant
detritus)
8
1­
De
Silva
1980
1­
De
Silva
and
Wijeyaratne
1977
1­
Fitch
and
Lavenberg
1971
1­
Moser
1996a
1­
Eggold
and
Motta
1991
2­
De
Silva
and
Wijeyaratne
1977
2­
Fitch
and
Lavenberg
1971
2­
Fitch
and
Lavenberg
1971
2­
Fitch
and
Lavenberg
1975
3­
De
Silva
and
Perera
1976
3­
Hendricks
1961
4­
Hendricks
1961
4­
Thomson
1963
5­
Thomson
1963
5­
Walker
1961
6­
De
Silva
1980
6­
Thomson
1966
7­
Walker
1961
7­
Duffy
1987
8­
Johnson
and
McClendon
1970
8­
Odum
1973
9­
Pafford
1983
9­
Fitch
and
Lavenberg
1975
Food
habits
fish­
eating
birds
(
pelicans,

terns,
gulls,
and
frigate
1
2
schools
in
coastal
waters
including
bays
and
estuaries,
may
enter
fresh
1
2
5
(
diatoms,
blue­
green
algae,
green
filamentous
algae,
cladocerans,
and
other
microscopic
plants
and
animals)
2,3,7,9
(
diatoms,
blue­
green
algae,
green
filamentous
algae,
plant
fibers,
detritus,
mud,
and
occasionally
a
copepod
or
other
small
animal)
5
(
diatoms,
desmids,
green
algae,
crustaceans,

nematodes,
invertebrate
eggs,
polychaetes,

foraminiferans,
and
silaceous
spines)
1
ingests
the
surface
scum
which
contains
numerous
diatoms2
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22
23
24
25
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CB
CC
CD
CE
CF
CG
Haplotype
diversity
Foraging
method
When
/
Frequency
Predators
Substocks
(
Geographic
Area)

no
diel
pattern1
NF
NF
NF
silk
shark2
1­
Moyle
1976
1­
Marais
1980
1­
Thomson
1963
2­
Thomson
1963
2­
Odum
1973
2­
Springer
1957
3­
Odum
1968
3­
Thomson
1966
4­
Walker
1961
5­
Thomson
1966
6­
Hiatt
1944
7­
Odum
1973
8­
Pafford
1983
Nucleotide
diversity
(#)

Adults
piscivorous
fishes,
diving
birds,

human
beings,
and
sharks3
M.
cephalus
will
feed
at
water
surface
when
thick
concentrations
of
micro­
algae
are
present7
contiuous
feeders,
with
increased
ingestion
on
rising
and
high
tides;

feeding
ceases
during
adverse
weather
and
offshore
spawning
migrations2
mud
and
sand
are
ingested
during
feeding,
the
roughest
portions
are
rejected
and
a
considerable
quantity
is
retained,
presumably
aiding
in
5
7
Food
habits
piscivorous
fishes
and
certain
diving
birds1
unique
pharyngeal
filtering
device
prohibits
swallowing
larger
food
items4,5,6,8
prefers
fine
particles
when
sediments
are
involved
in
feeding,
may
be
due
to
richness
in
organic
matter
and
microorganisms3
feeding
takes
place
on
muddy
bottoms
in
shallow
water,
mullet
swim
at
an
angle,
scooping
up
soft
surface
material
with
stiff
lower
jaws,
coarse
material
is
ejected
through
mouth
and
gills,

organic
material
is
retained1,2,5,6,7
sucks
up
the
surface
layer
of
mud
or
grazes
on
submerged
surfaces
such
as
rocks
or
Zosterna
leaves3,7,8
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22
23
24
25
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CH
CI
CJ
CK
CL
CM
CN
Genetic
variance
Mean
number
of
%
Polymorphic
First
Recruitment
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Total
Length
(
age)

NF
NF
NF
NF
NF
NF
120­
30
cm
1­
Walker
1961
GENETICS
Mean
heterozygosity
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CO
CP
CQ
CR
CS
CT
CU
CV
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
Full
Recruitment
Fishery
/
Total
Length
(
age)
Sampling
gear
eing
structure
/
Meth
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)

NG
baited
hook
NG
NG
NG
1,20.33
0.2
0.53
1
0.58
0.003
0.583
1NG
NG
0.995
1NG
NG
0.79
1
0.5
0.29
0.79
2
0.29
0.5
0.79
1­
Thomson
1963
2­
Thomson
1966
Page
726
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CW
CX
CY
CZ
DA
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
short­
term
tagging
NG
NG
0.377
long­
term
tagging
NG
NG
0.000537
short­
term
tagging
NG
NG
NG
market
sizes
NG
NG
NG
short­
term
tagging
NG
NG
0.367
NG
NG
NG
0.633
1­
Thomson
1963
MORTALITY
herbivorous,
iliophagous
and
omnivorous
marine
animals
compete
with
M.
cephalus
for
food,
such
as
milkfish,
other
Mugilidae,

estuarine
fish,
crustacea,
and
mollusks
1
Girella
triscupidata,
Mylio
australis,

Rhabdosargus
sara,
Myxus
elongatus,

Planiprora
fusca
,
and
Sillago
maculata
are
known
to
compete
for
space
with
M.
cephalus
1
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1234
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44
45
46
47
48
49
50
51
52
53
DB
DC
DD
DE
Interactions
/
associations
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

Resident,
with
observed
spawning
migration1
NF
some
resident,
some
transient2
mullet
fry
migrate
from
sea
toward
freshwater4
parasite:
Ancyrocephlus
vanbenedenii
2
juveniles
associated
with
Lepomis
macrochirus
3
parasites:
44
families,
152
spp.
(
see
reference)
4
parasite:
isopod
(
Lironeca
californica
)
5
1­
Thomson
1963
1­
Thomson
1955
1­
Thomson
1963
2­
De
Silva
1980
2­
Pafford
1983
2­
Thomson
1966
3­
Allen,
L.
G.
1982
3­
Horn
and
Allen
1985
4­
Love
and
Moser
1983
4­
Thomson
1955
5­
Bennett
1993
5­
Pafford
1983
6­
Suzuki
1965
BEHAVIOR
movement
southward
in
the
fall
with
a
limited
northward
movement
in
spring
and
summer,

with
a
tendency
to
move
into
freshwater
areas
during
spring
and
summer5
young
30­
60
mm
long
migrate
from
the
open
sea
into
brackish
water
of
estuaries
and
ascend
rivers,
where
they
grow
up
to
200­
250
mm
before
autumn
of
1st
yr.,
when
they
return
to
sea6
Movement
spawns
at
sea
in
the
fall
followed
by
a
return
of
the
prejuveniles
and
adults
to
the
estuary
in
winter3
tag/
recapture
from
3.7
km
(
212
days)
to
611
km
(
393
days),

45%
were
caught
within
5
km
of
place
of
tagging2
spawning
migration
with
downstream
movement
toward
the
sea4
spawning
migration
evident
from
tagging
study
(
Queensland,

Australia)
1
parasites:
fungus(
Saprolengnia
sp.
),
Myosporidia
(
Myxobolus
esiguus
),

Actinomyces
(
Actinomyces
sp
.),
Trematoda
(
Microcotyle
mugilis,

Microcotyle
psuedomugilis,
Microcotylye
macracantha,
Mesostephanus
appendiculatoides,
Phagicola
longa,
Stephanostomum
sp.,
Hysterolecitha
elongata,
Dicrogaster
sp.,
Haplorchis
taichui,
Stellantchionus
falcatus,

Centrocestus
formosanus,
Centrocaecum
spiculigerum,
Strictodora
sawakinensis,
Parascotyle
longa,
Haploporus
longicolum,
Haploporus
benedeni,
Heterophyes
herterophyes,
Heterophyes
nocens
),
Cestoda
(
Otobothriu
mugilis
),
Acanthocephala
(
Neoechinorhynchus
chilkaensis,

Atactorhynchus
mugilis
),
Copepoda
(
Lernanthropus
shiskidai,

Clavellopsis
longimanus,
Ergasulus
lizae
)
1
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24
25
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Schooling
Territoriality
Yes1,2,3,4,5,6,7,8
NF
1­
Moser
1996a
2­
Fitch
and
Lavenberg
1971
3­
Moyle
1976
4­
Springer
1957
5­
Walker
1961
6­
Thomson
1966
7­
Thomson
1955
8­
Fitch
and
Lavenberg
1975
tagging
indicates
schools
have
a
permanence
of
approx.
6
months,

but
some
emmigration
and
immigration
to
the
school
occurs7
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71
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73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
A
B
C
D
E
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81
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83
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85
86
87
88
89
90
91
92
93
94
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97
98
99
100
101
102
103
104
105
F
G
H
I
JKL
M
NO
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85
86
87
88
89
90
91
92
93
94
95
96
97
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100
101
102
103
104
105
P
Q
R
S
T
U
V
W
X
Y
Z
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74
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
AA
AB
AC
AD
AE
AF
AG
AH
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
AI
AJ
AK
AL
AM
AN
AO
AP
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64
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66
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68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Page
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63
64
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66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
BD
BE
BF
BG
BH
BI
BJ
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63
64
65
66
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68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
BK
BL
BM
BN
BO
BP
BQ
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63
64
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66
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68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
BR
BS
BT
BU
BV
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64
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71
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73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
BW
BX
BY
BZ
CA
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68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
CB
CC
CD
CE
CF
CG
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66
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68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
CH
CI
CJ
CK
CL
CM
CN
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73
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80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
CO
CP
CQ
CR
CS
CT
CU
CV
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64
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66
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68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
CW
CX
CY
CZ
DA
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
DB
DC
DD
DE
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69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Page
745
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19
20
21
22
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Muraenidae1
Gymnothorax
mordax
1
1shallow
to
40
1­
Robins
et
al.
1991
1­
Eschmeyer
1998
1­
Gates
&
Frey
1974
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Robins
et
al.
1991
3­
Roedel
1948
TAXONOMY
RANGE
California
moray
eel;
moray;
conger
eel;
marina;
muraena1,2,3
CA
(
Pt.
Conception)
to
s.
Baja
CA
(
Bahia
Magdalena)
1
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14
15
16
17
18
19
20
21
22
F
G
H
I
JKLMNOPQR
Length
weight
equation
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)
to
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
Female
Both
1Both:
W=
0.00000011SL3.43
NF
1
152
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Quast
1968b
1­
Love
et
al
2000
AGE
&
GROWTH
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
Page
747
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12
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14
15
16
17
18
19
20
21
22
S
T
U
V
W
X
Y
Z
AA
AB
AC
AD
Age
composition
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
NF
NF
NF
NF
NF
NF
NF
1,3>
26
NF
NF
NF
Oviparous1
2
30
1­
Fitch
&
Lavenberg
1971
1­
Charter
&
Moser
1996
2­
Love
1996
3­
Fitch
&
Lavenberg
1975
r2
Breeding
Beha
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748
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
AE
AF
AG
AH
AI
AJ
AK
AL
AM
AN
AO
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
avior
Sexual
dimorphism
1st
maturity
50%
maturity
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
Page
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
states
present
Spawning
Frequency
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
(
per
year)

NF
NF
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
REPRODUCTION
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10
11
12
13
14
15
16
17
18
19
20
21
22
BD
BE
BF
BG
BH
BI
Size
specific
fecundity
equation
Size
at
settlement
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
Larval
duration
(
SL
in
mm)

NF
NF
NF
NF
NF
NF
DISPERSAL
&
RECRUITME
Mean
(
range)
fecundity
in
eggs/
g
female
wt.
Fecundity
Fecundity
range
in
#
eggs
(@
cm,
grams,
or
age)
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14
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18
19
20
21
22
BJ
BK
BL
BM
BN
BO
BP
BQ
BR
Size
Fish
Size
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
(
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
NF
13
to
4
planktonic1
NF
1to
>
108
planktonic1
NF
NF
NF
1­
Charter
&
Moser
1996
1­
Charter
&
Moser
1996
1­
Charter
&
Moser
1996
1­
Charter
&
Moser
1996
ENT
Food
habits
Larvae
Eggs
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16
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18
19
20
21
22
BS
BT
BU
BV
BW
BX
BY
BZ
CA
CB
Adults
Fish
Size
Food
habits
Food
habits
Predators
(
SL
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Habitat
Foraging
guild
Foraging
method
NF
NF
NF
NF
NF
NF
NF
carnivore
NF
1­
Fitch
&
Lavenberg
1971
1­
Quast
1968d
2­
Love
1996
2­
Fitch
&
Lavenberg
1971
3­
Feder
et
al.
1974
3­
Love
1996
4­
Limbaugh
1964
5­
Fitch
and
Lavenberg
1975
6­
Feder
et
al.
1974
Life
Stage
(
crustaceans,
especially
crabs;
small
fish,
such
as
blacksmith,
kelp
bass,
and
black
surfperch;

mollusks,
such
as
octopi)
1,2,3,4,5,6
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
rocky
reefs;
kelp
forest
bottoms
in
crevices
1,2,3
Juveniles
Page
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20
21
22
CC
CD
CE
CF
CG
CH
CI
CJ
CK
CL
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
When
/
Frequency
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
diurnal
1
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Fitch
&
Lavenberg
1975
Nucleotide
diversity
(#)
Mean
heterozygosity
GENETICS
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19
20
21
22
CM
CN
CO
CP
CQ
CR
CS
CT
CU
CV
CW
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
First
Recruitment
Full
Recruitment
Method
of
Effective
population
size
Total
length
(
age)
Total
length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTA
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19
20
21
22
CX
CY
CZ
DA
DB
DC
DD
DE
DF
Rate
of
Exploitation
Age
range
used
Rate
of
Exploitation
Interactions
/
associations
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
NF
NF
NF
NF
NF
NF
NF
NF
1­
Hobson
1971
2­
Limbaugh
1961
3­
Love
&
Moser
1983
BEHAVIOR
Movement
infected
by
parasites
from
the
families
Hatschekiidae,
Bucephalidae,
Hemiuridae,

and
Piscicolidae3
cleaned
by
the
shrimp
Hippolysmata
californica
2
Iinfected
by
ectoparasites
from
the
family
Dichelesthiidae1
ALITY
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14
15
16
17
18
19
20
21
22
DG
Territoriality
NF
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22
23
24
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Osmeridae1,2
Thaleichthys
pacificus
1
eulachon
1
surface
to
300
m
Osmeridae1,2
Spirinchus
thaleichthys
1,2
longfin
smelt
Bering
sea
to
Monterey
Bay,
central
California
1
2in
sea
usually
inshore
Osmeridae
1
Spirinchus
starksi
1
night
smelt
1
shallow
water
Osmeridae
1
Hypomesus
pretiosus
1,2
surf
smelt
1
inshore
and
surf
zones
smelt,
surf
fish,
nightfish,
perlin,

silver
smelt,
dayfish,
pacific
surf
smelt
1,2,3,4
Sacramento
smelt,
delta
smelt
1,2,3
candlefish,
ulchen,
oolakan,

oolachon,
hoolakan,
hoolikan,

olachan,
ollachan,
oolichan,

hooligan,
ulichan,
uthlecan,

yshuh,
smallfish,
sak,
ssag,

yuurachon
1,2,3,4,5
Pribilof
Island,
Bering
Sea
(?)
south
to
Pt.
Conception,

central
California
1
TAXONOMY
RANGE
whitebait,
sand
smelt,
nightfish
1,2,3
Prince
William
Sound,
Gulf
of
Alaska
to
Long
Beach,

southern
California
1
Sheikof
Bay,
southeastern
Alaska
to
Pt.
Arguello,

central
California
1
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31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
F
G
H
I
JKL
Total
Length
(
cm)
Max.
reported
Geographic
area
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
Female
Both
2females:
W=
0.84L­
95.5
(
r=
0.85;
r2=
0.72)
NF
1
31
cm
TL
NF
NF
NF
NF
1W=
0.00000237L3.3732
(
freshwater
pop.)
1SL=
0.966
x
TL(
0.823);
SE:
1.814
1
22.8
cm
TL
NF
NF
NF
NF
1TL=
0.516
x
SL(
1.190);
SE:
2.181
1TL=­
3.494
x
FL(
1.150);
SE:
1.199
1FL=
3.476
x
TL(
0.864);
SE:
1.040
NF
1SL=­
6.478
x
TL(
0.876);
SE:
0.841
1
23
cm
TL
NF
NF
NF
NF
1TL=
8.031
x
SL(
1.135);
SE:
0.957
1TL=
12.395
x
FL(
0.990);
SE:
1.196
1FL=­
11.487
x
TL(
1.002);
SE:
1.204
NF
1SL=­
1.940
x
TL(
0.863);
SE:
2.099
1
30.5
cm
TL
Puget
Sound
1
NG
NG
NG
1TL=
3.031
x
SL(
1.153);
SE:
2.426
1TL=
0.161
x
FL(
1.084);
SE:
1.880
1FL=
0.293
x
TL(
0.920);
SE:
1.732
Linf
in
cm
(
SE)

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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
M
N
O
P
Q
R
S
T
U
V
W
r2
Male
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NG
NG
NG
NG
NG
NG
NG
NG
NG
Scales
/
whole
and
length
frequencies
No
k
(
SE)
to
(
SE)

AGE
and
GROWTH
Growth
Parameters
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
X
Y
Z
AA
AB
AC
Age
composition
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
NF
4
14
yrs.
2
NF
65­
6
yrs
NF
1,3,4
b2
yrs.
NF
dioecious2
promiscuous3
53
yrs.

NF
NF
43
yrs.
NF
dioecios
dioecious
No
2,4,5
1,35
yrs.
1
dioecious2
polyandrous1
73
yrs.
polygamous2
8
3
yrs
(
185
mm
SL)
majority
dioecious,
some
hermaphrodites1
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21
22
23
24
25
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28
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30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AD
AE
AF
AG
AH
AI
AJ
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
oviparous2,4,5
No
NF
2,3NG
(
3
yrs)
NG
(
3
yrs)
NG
4NG
16.9
cm
(
3
yrs)
NG
oviparous5
No
males
larger2
58.8­
15.2
cm
TL
(
2
yrs)
8.8­
15.2
cm
TL
(
2
yrs)
NG
oviparous
No
NF
NF
NF
NF
NF
oviparous1,2,3
No
females
larger1,3,4,5,6,7
1NG
1­
2
yrs.
NG
51­
2
yrs.
1­
2
yrs.
NG
61­
2
yrs.
1­
2
yrs.
NG
7
90
mm
SL
(
1
yr)
NG
NG
sexually
dimorphic
color2,6,8
adult
males
have
larger,
rounder
anal
fin
and
are
usually
darker
than
females7
males
have
tubercles
on
the
sides,
head,

and
fins6
Sexual
dimorphism
1st
maturity
50%
m
Breeding
Behavior
at
spawning
time
males
are
distinguished
by
having
tubercles
on
the
scales,
the
muscles
of
the
body
wall
considerably
more
developed,
and
paired
fins
are
longer6
sexual
dimorphism
in
coloration
and
enlargement
of
the
fins
in
breeding
fish4
adult
males
have
thickened
ridge
along
side,
bluish
brown
above
with
fine
black
speckling
on
back,
shading
to
silver
7
REPRODUCTION
Total
length
(
age)
at
males
in
spawning
condition
are
dull
olive
green
with
yellow­
silver
abdomen,
and
develop
wart­
like
excrescences,
called
pearl
organs1,3,5
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22
23
24
25
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AK
AL
AM
AN
AO
AP
AQ
AR
AS
AT
Fertilized
eggs
present
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
NG
NG
NG
NG
NF
British
Columbia2,3,7,8,15,18
*****

NG
otoliths/
surface
burn
No
interreader
comparison
Columbia
River7,11,21
Klamath
River(
northern
CA)
northward20
NG
NG
NG
NG
NF
British
Columbia18
Lake
Washington,
WA13,14
*****
*****

Sacramento­
San
Joaquin
Delta,
CA16
central
California4,5,7
California9
west
coast17
NF
NF
NF
NF
NF
California5
NG
NG
NG
NG
NF
British
Columbia12,18
NG
NG
NG
NG
Puget
Sound,
WA6,19
NG
scales/
annual
mark
reading
No
No
Puget
Sound,
WA22
NG
scales/
length
frequency
No
No
central
California4
California5,10
CalCOFI
area1
maturity
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertiliz
Page
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17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
Mean
Fecundity
Spawning
females
present
Both
reproductive
states
present
eggs/
g
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
(
weight
range)

most
die
after
spawning2,13,17
1,5,6,725,000
eggs
(
avg.)

27,000­
40,000
eggs
83,242­
47,798
eggs
822,943
eggs
(
avg.)

97,000­
31,000
eggs
1017,000­
40,000
eggs
117,000­
31,000
eggs
118,100
eggs
(
avg.)

19,621­
23,634
eggs
75,000­
24,000
eggs
918,000­
24,000
eggs
multiple
spawners3,5
NF
*****
*****
*****
*****
multiple
spawners1,2,4,9,10,15,16,18
1,31,320­
29,950
eggs/
batch
single
spawners6
2,91,300­
37,000
eggs
annual
spawner14
34,020­
13,714
eggs
more
frequent
late
afternoon,
near
high
water
on
ebbing
tides
18
4,51,500­
30,000
eggs
more
than
once
per
day,
over
several
days,
more
than
once
during
season
18
6<
18,000­>
39,000
eggs
915,000­
20,000
eggs
102,500
to
37,000
eggs
12
1,400­
30,000
eggs/
batch
most
die
after
spawning,
but
some
live
to
spawn
a
second
time
8,11,12,14
ation;*****=
peak
spawn;
#####
=
doubtful
spawning
records
most
die
after
spawning,
but
some
live
to
spawn
a
second
time
5,7,8
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22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BE
BF
BG
BH
Size
specific
Number
of
eggs
fecundity
eq
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
Larval
duration
117,300
(
14.5
cm
SL)­
39,600
(
18.8
cm
SL)
1F=
726.9W­
9645
(
r=
0.62;
r2=
0.39)
(
W=
grams)
130­
40
days(
4.4­
7.2
C)
NF
320,000­
60,000
eggs
(
140­
190
cm
TL)
219­
40
days
614
days
714­
21
days
921­
28
days
(
4­
5
C)

1335­
39
days
1821
days
NF
414­
21
days
7,1440
days
(
7
C)

5535
eggs
(
46
mm
SL)­
2,425
eggs
(
61
mm
SL)
1225
days
(
9.6­
10.6
C)

NF
Nf
3,5,814
days
NF
2
eggs
=
396.2
(
SL)
­
40160
[
length
in
mm]
110­
12
days
(
summer)
NF
28­
28
days
8,1714­
28
days
108.5
days
(
17.6
C)
to
13
days
(
12.1
C)

119­
10
days
148.5­
30
days
1510­
11
days
1614­
21
days
6,575
(
12
cm
SL),
7,139
(
12.4
cm
SL),

22,169
(
15.2
cm
SL),
17,848
(
15.3
cm
SL),

19,163
(
15.9
cm
SL)
4
6
1,400
eggs
(
105
mm
SL);
17,000
(
145
mm
SL);
29,000
(
175
mm
SL)
1419
days
(
8.5­
11.5
C)
and
30­
40
days
(
4.4­
7.2
C)

Fecundity
517,336
eggs
(
89
mm
SL)­
22,205
eggs
(
126
mm
SL)
(
freshwater
pop.)
metamorphosis
to
juvenile
30­
60
days,
depending
on
temperature1
DISPERSAL
&
RECRUIT
24,020
eggs(
15
cm
TL),
7,260
eggs
(
15.5
cm
TL),
8,880
eggs
(
15.9
cm
TL),
7,980
eggs
(
16.5
cm
TL),
and
13,714
eggs
(
16.9
Page
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BI
BJ
BK
BL
BM
BN
Eggs
Size
at
settlement
Size
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
Fish
Size
NF
NF
20.08­
1.0
mm
birds2
5<
1.0
mm
8,91.0
mm
(
avg.)
eggs
spawned
at
night9,18
spawns
in
shoals10,18
NF
NF
30.9­
1.1
mm
stream
dwelling
cottids1
35.3­
9.8
mm
at
hatching
71.0
mm
(
avg.)

81.2
mm
(
avg.)
spawned
in
fresh
or
slightly
brackish
water13,19
NF
NF
NF
NF
NF
demersal,
adhesive
eggs13
NF
NF
11.1
mm
(
avg.)
13.0­
40
mm
SL
30.93­
1.24
mm
24.0­
70
mm
SL
21.0
mm
daytime
spawners3,7,9,20
60.88­
1.30
mm
81.0­
1.2
mm
101.02
mm
egg
hatching
aided
by
stirring
of
gravel
by
waves21
eggs
are
laid
intertidally
on
beaches
of
coarse
sand
with
fine
gravel18
spawned
in
the
surf
zone
of
beaches,
esp.
during
periods
of
high
tides7,15,16
35.0­
6.5
mm
(
at
hatching),

postlarvae
are
17­
35
mm
TL
shore­
dwelling
sculpins,

gulls,
and
other
birds;
the
staghorn
sculpin,
starry
flounder,
and
shiner
perch
may
be
predators3
demersal
eggs
attached
to
course
sand
grains
in
the
upper
tidal
zone1,3,4,5,7,9,10,14,15,16,20,21
demersal
eggs
attached
to
submerged
vegetation
or
rocks4,12
eggs
are
deposited
in
areas
of
pea­
sized
gravel
and/
or
semi­
sandy
areas
with
sticks
and
debris;

occur
in
freshwater18
34­
7
mm
at
hatching,

postlarval
length
is
probably
35
mm
spawning
substrate
thought
to
be
sandy­
gravel
areas
with
sand
or
aquatic
plants18
eggs
spawned
in
rivers
and
streams;
demersal,

attached
to
sand
grains3,8,11,12,17,19,23
TMENT
eggs
spawned
nocturnally
in
surf
at
beaches
composed
of
coarse
grain
sand1,6,8,13,22
benthic
in
riverine
or
upper
estuarine
environments18
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45
46
47
48
49
50
51
52
BO
BP
BQ
BR
BS
BT
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Fish
Size
found
in
rivers,
drift
out
into
ocean
5,6
planktivores5
NF
NF
birds1
135­
45
mm
TL
(
at
hatching)

230­
140
mm
pelagic3
carnivorous
planktivores5
NF
NF
135­
45
mm
TL
(
at
hatching)

surface
waters
of
bays
and
estuaries2
(
zooplankton
and
phytoplankton)
5
222­
88
mm
TL
NF
NF
NF
NF
NF
135­
45
mm
TL
(
at
hatching)

planktonic1
planktivores
NF
NF
NF
135­
45
mm
TL
(
at
hatching)

(
dipterous
larvae
and
mayfly
nymphs)
2
235­
85
mm
TL
planktivorous
carnivores5
predatory
fishes,
birds
(
such
as
gulls
and
terns),
and
marine
mammals
(
such
as
harbor
seals)
2
(
copepods
(
consumed
by
eulachon
of
24­
29
mm
FL),
copepod
nauplii
(
10­
29
mm
FL),
and
copepod
eggs
(
26­
29mm
FL))
4
(
phytoplankton,
copepods,
mysids,
ostracods,

barnacle
larvae,
cladocerans,
worm
larvae,
and
larvae
of
their
own
species)
5
(
copepods
(
consumed
by
eulachon
of
22­
157
mm
FL),
trematodes
(
36
mm
FL),
decapod
(
52
mm
FL),
larvacea
(
52
mm
FL),
cirripedia
(
118
mm
FL)
3
(
copepods
(
comsumed
by
surf
smelt
of
63­
129
mm
FL),
decapods,
larvacea
(
54­
129
mm
FL)
3
(
copepods
(
consumed
by
surf
smelt
of
155­

166
mm
FL),
amphipods
(
155­
162
mm
FL),

decapods
(
155
mm
FL),
and
euphausiids
(
155
4
found
in
bays
and
estuaries,
but
primarily
offshore4
(
phytoplankton,
copepod
eggs
copeods,

mysids,
ostracods,
barnacle
larvae,
Cladocera,

and
worm
larvae)
1
pelagic
in
riverine­
marine
waters,
most
often
found
in
estuaries6
Larvae
pelagic
along
Pacific
coast
,
bays
and
estuaries2
HABIT
Food
habits
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31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BU
BV
BW
BX
BY
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
coastal
waters5
planktivores6
NF
NF
found
in
deeper
waters
than
adults6
(
euphausiids
and
copepods)
1
(
primarily
euphausiids)
3
carnivorous
planktivores6
NF
NF
(
copepods)
2
pelagic
in
estuaries4
NF
NF
NF
NF
NF
plantivores
NF
NF
estuaries
used
as
nursery
grounds3
planktivorous
carnivores6
(
phytoplankton,
copepod
eggs,
copepods,

trematodes,
mysiids,
ostracods,
cirripedia
nauplii,
diatoms,
fish,
and
polychaete
larvae)
8
(
cladocera,
copepods
,
cirrepedia
eggs,

trematodes,
amphipods,
and
copepod
eggs)
9
(
copepods
(
consumed
by
surf
smelt
of
155­

166
mm
FL),
amphipods
(
155­
162
mm
FL),

decapods
(
155
mm
FL),
and
euphausiids
(
155
mm
FL))
5
predatory
fishes,
birds
(
such
as
gulls
and
terns),
and
marine
mammals
(
such
as
harbor
2
(
calanoid
copepods,
cladocerans,
amphipods,

and
other
small
crustaceans)
6
(
copepods
(
consumed
by
surf
smelt
of
63­
129
mm
FL),
decapods,
larvacea
(
54­
129
mm
4
found
in
estuaries,
inhabit
middle
and
bottom
strata
of
water
column2
along
Pacific
coast
and
in
euryhaline
waters
of
estuaries1
(
copepods
(
consumed
by
eulachon
of
22­
157
mm
FL),
trematodes
(
36
mm
FL),
decapod
(
52
mm
FL),
larvacea
(
52
mm
FL),
cirripedia
(
118
mm
FL)
4
(
copepods
(
consumed
by
eulachon
of
24­
29
mm
FL),
copepod
nauplii
(
10­
29
mm
FL),
and
copepod
eggs
(
26­
29mm
FL))
5
(
euphausiids,
copepods,
and
other
planktonic
crustaceans)
6
predatory
fish,
marine
mammals,

and
birds1
predatory
fish,
marine
mammals,

and
birds1
Juveniles
TAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Food
habits
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22
23
24
25
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30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BZ
CA
CB
CC
CD
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
planktivore2,10
zooplanktivore4
NF
harbor
seal2
(
euphausiids
and
copepods)
1
(
krill
and
copepods)
2
(
copepods)
9
(
eggs
and
copepods)
11
anadromous2,6,8
carnivores
NF
northern
squawfish9
harbor
seals10
(
mysid
shrimp)
4
primarily
estuarine
species5
found
in
euryhaline
conditons6,7
(
primarily
zooplankton,
esp.
mysids)
8
carnivorous
planktivores10
NF
planktivores5
NF
NF
birds,
seals,
and
a
wide
variety
of
fish4
(
small
shrimplike
crustaceans)
3
planktivores2,5,7
NF
spring
salmon1
(
copepods,
amphipods,
krill,
and
larval
fish)
2
rhinoceros
auklet8
planktivorous
carnivores10
pelagic,
but
often
found
near
the
bottom
in
estuarine
and
marine
waters8
(
crustacean
zooplankton,
small
bottom
crustaceans,
and
insect
larvae)
1
(
crustaceans,
copepods,
amphipods,
larvae,

euphausiids,
shrimp
larvae,
marine
worms,
and
insects)
1
two
resident
freshwater
populations:

Harrison
Lake,
British
Columbia
and
Lake
Washington,
Washington
4
marine,
nearshore;
sometimes
enter
freshwater1
nearly
continuous
feeding
with
a
slight
diel
variation1
(
calanoid
copepods,
cladocerans,
amphipods,

mysids,
and
other
small
crustaceans)
10
fishes
(
such
as
Pacific
salmon,
lingcod,

opportunistic,
possibly
feeding
in
monospecific
prey
patches1
salmon,
halibut,
and
other
large
predatory
species5
(
calanoid
copepods,
hyperiid
amphipods,
decapod
larvae,
larvaceans,
sand
lance
and
flatfish
larvae)
7
halibut,
stugeon,
cod,
porpoise,
finback
whales,
seals,
sea
lions,
spring
salmon,

and
fur
seals13
(
euphausiids,
copepods,
and
other
planktonic
crustaceans)
10
spiny
dogfish,
sturgeon,
Pacific
halibut
and
arrowtooth
founder,
salmon,
harbor
seals,
orcas,
finback
whales,
northern
fur
seals,
Pacific
white­
sided
dolphins,
and
various
sea
birds3,12
dogfish,
salmon,
hake,
Pacific
cod,

lingcod,
and
marine
mammals;
harbor
seals
and
fur
seals7
(
opposum
shrimp,
Neomysis
mercedis
,

copepods,
and
other
crustaceans)
6
predatory
fishes,
birds
(
such
as
gulls
and
terns),
and
marine
mammals
(
such
as
harbor
seals)
12
adults
do
not
feed
during
their
spawning
migration2
salmon,
sturgeon,
and
other
predatory
marine
animals6
spends
most
of
life
in
salt
water,

moving
up
rivers
to
spawn
in
large
numbers
in
spring7
salmon,
birds
(
ie.
Terns),
and
marine
mammals
(
incl.
Harbor
seals,
northern
fur
seals,
and
Dall's
porpoise)
3
lives
in
dense
school
on
the
outer
continental
shelf;
anadromous
fish3
Adults
predatory
species
of
marine
mammals,

fish
(
such
as
sturgeon),
and
birds11
anadromous,
found
nearshore
and
in
costal
inlets
and
rivers8
Food
habits
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25
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CE
CF
CG
CH
CI
CJ
CK
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Puget
sound
stocks
are
genetically
different
than
coastal
stocks1
Nucleotide
diversity
(#)
Mean
heterozygosity
GENETICS
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CL
CM
CN
CO
CP
CQ
CR
CS
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Approach
Effective
population
size
Fishery
/
Total
Length
(
age)
Fishery
/
Total
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CT
CU
CV
CW
CX
CY
CZ
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
DA
DB
DC
DD
DE
Interactions
/
associations
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

NF
parasites:
1
family,
1
spp.
(
see
reference)
3
NF
No
geographic
homing1
Pacific
herring
and
yellowfin
goby1
NF
NF
parasites:
1
family,
1
spp.
(
see
reference)
3
NF
NF
NF
NF
NF
NF
NF
parasites:
7
families,
10
spp.
(
see
reference)
3
Movement
viability
related
to
protection
of
historic
spawning
grounds
from
pollution
and
habitat
alterations2
migrates
from
rivers
to
coastal
feeding
areas,
and
later,
from
the
coastal
feeding
areas
to
riverine
spawning
areas2
specific
spawning
sites
are
used,
there
is
no
information
regarding
whether
fish
return
to
their
natal
spawning
grounds3
adults
migrate
from
salt
and
brackish
water
to
freshwater
during
winter
to
spawn1
juveniles
and
adults
move
to
lower
estuarine/
marine
areas
in
spring
and
summer,
and
to
upper
estuarine
areas
in
fall;
in
winter,
adults
move
to
freshwater
i
3
found
in
association
with
Cottus
sp.
and
Oncorhynchus
nerka
1
competes
with
sockeye
salmon
for
food2,3
parasites:
digenea
(
Lecithaster
gibbosus
),

cestoda
(
Phyllobothrium
sp.
larval),
and
nematoda(
Contracaecum
sp.
larval)
4
BEHAVIOR
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30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
Schooling
Territoriality
Yes1,4
NF
schools
at
depths
of
150­
750
feet4
NF
NF
Yes2,3
NF
NF
Yes
8
surf
perch
nearing
the
spawning
grounds
may
segregate
by
size8
Yes;
school
often
segregated
by
sex1,5,7
during
spawning
run
schools
are
segregated
by
sex6
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
A
B
C
D
E
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Eschmeyer
and
Herald
1983
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
2­
Eschmeyer
and
Herald
1983
3­
Hart
1973
4­
Love
1996
5­
Hart
and
McHugh
1944
Page
775
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
F
G
H
I
JKL
1­
Dryfoos
1965
1­
Karpov
and
Kwiecien
1988
1­
Love
et
al
2000
1­
Schaefer
1936
Page
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67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
M
N
O
P
Q
R
S
T
U
V
W
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
X
Y
Z
AA
AB
AC
1­
Moulton
1974
1­
Love
1996
1­
Yap­
Chiongco
1941
1­
Pederson
et
al
1995
1­
Emmett
et
al
1991
2­
Yap­
Chiongco
1941
2­
Moulton
1974
2­
Delacy
and
Batts
1963
2­
Emmett
et
al
1991
2­
Pentilla
1978
3­
Dryfoos
1965
3­
Moyle
1992
3­
Moulton
1974
4­
Hart
and
McHugh
1944
4­
Fitch
and
Lavenberg
1975
5­
Pentilla
1978
5­
Moyle
1992
6­
Hay
et
al
1997
7­
Yap­
Chiongco
1941
8­
Schaefer
1936
Page
778
of
1167
5­
2220e
­
Osmeridae
53
54
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64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
AD
AE
AF
AG
AH
AI
AJ
1­
Moser
1996a
1­
Hart
1973
1­
Garrison
and
Miller
1982
1­
Moyle
1992
2­
Love
1996
2­
Moulton
1974
2­
Breder
and
Rosen
1966
2­
Barraclough
1964
3­
Garrison
and
Miller
1982
3­
Moyle
1992
3­
Yap­
Chongco
1941
3­
Hay
et
al
1997
4­
Pederson
et
al
1995
4­
Baxter
1999b
4­
Dryfoos
1965
4­
Pederson
et
al
1995
5­
Emmett
et
al
1991
5­
Levy
1990
5­
Emmett
et
al
1991
5­
Emmett
et
al
1991
6­
Hart
and
McHugh
1944
6­
Hart
and
McHugh
1944
6­
Pentilla
1978
7­
Pentilla
1978
7­
Eschmeyer
and
Herald
1983
7­
Schaefer
1936
8­
Schaefer
1936
Page
779
of
1167
5­
2220e
­
Osmeridae
53
54
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
AK
AL
AM
AN
AO
AP
AQ
AR
AS
AT
1­
Moser
1996a
2­
Hart
1973
3­
Love
1996
4­
Wang
1981
5­
Moyle
1992
6­
Garrison
and
Miller
1982
7­
Hearne
1983
8­
Barraclough
1964
9­
Moyle
1976
10­
Baxter
1999b
11­
Hay
et
1997
12­
Yap­
Chiongco
1941
13­
Dryfoos
1965
14­
Moulton
1970
15­
Pederson
et
al
1995
16­
Radtke
1966
17­
Emmett
et
al
1991
18­
Hart
and
McHugh
1944
19­
Penttila
1978
20­
Eschmeyer
and
Herald
1983
21­
Parente
and
Snyder
1970
22­
Schaefer
1936
Page
780
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
1­
Hart
1973
1­
Hart
1973
2­
Love
1996
2­
Love
1996
3­
Fitch
and
Lavenberg
1971
3­
Wang
1981
4­
Fitch
and
Lavenberg
1975
4­
Fitch
and
Lavenberg
1975
5­
Moyle
1992
5­
Moyle
1992
6­
Breder
and
Rosen
1966
6­
Breder
and
Rosen
1966
7­
Barraclough
1964
7­
Moyle
1976
8­
Moyle
1976
8­
Pederson
et
al
1995
9­
Baxter
1999b
9­
Emmett
et
al
1991
10­
Yap­
Chiongco
1941
10­
Hart
and
McHugh
1944
11­
Dryfoos
1965
11­
Parente
and
Snyder
1970
12­
Moulton
1970
13­
Hay
1998
14­
Emmett
et
al
1991
15­
Hart
and
McHugh
1944
16­
CDFG
1982
17­
Eschmeyer
and
Herald
1983
18­
Schaefer
1936
Page
781
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
BE
BF
BG
BH
17approx.
14
days
19
10
days
in
summer,
longer
in
winter
1­
Hart
1973
1­
Pederson
et
al
1995
1­
Hart
1973
1­
Emmett
et
al
1991
2­
Thompson
et
al
1936
2­
Schaefer
1936
2­
Love
1996
3­
Hearne
1983
3­
Fitch
and
Lavenberg
1971
4­
Yap­
Chiongco
1941
4­
Fitch
and
Lavenberg
1975
5­
Dryfoos
1965
5­
Moyle
1992
6­
Schaefer
1936
6­
Breder
and
Rosen
1966
7­
Moyle
1976
8­
Baxter
1999b
9­
Hay
et
al
1997
10­
Middaugh
et
al
1988
11­
Yap­
Chiongco
1941
12­
Moulton
1970
13­
Pederson
et
al
1995
14­
Emmett
et
al
1991
15­
Hart
and
McHugh
1944
16­
C.
D.
F.
G.
1982
17­
Pentilla
1978
18­
Parente
and
Snyder
1970
19­
Schaefer
1936
Page
782
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
BI
BJ
BK
BL
BM
BN
1­
Moser
1996a
1­
Moser
1996a
1­
Moulton
1970
1­
Moser
1996a
2­
Hart
1973
2­
Hart
1973
2­
Pederson
et
al
1995
2­
Yap­
Chiongco
1941
3­
Wang
1981
3­
Love
1996
3­
Emmett
et
al
1991
4­
Breder
and
Rosen
1966
4­
Wang
1981
5­
Hay
et
al
1997
5­
Thompson
et
al
1936
6­
Yap­
Chinogco
1941
6­
Fitch
and
Lavenberg
1971
7­
Dryfoos
1965
7­
Fitch
and
Lavenberg
1975
8­
Emmett
et
al
1991
8­
Moyle
1992
9­
Parente
and
Snyder
1970
9­
Hearne
1983
10­
Schaefer
1936
10­
Breder
and
Rosen
1966
11­
Barraclough
1964
12­
Moyle
1976
13­
Baxter
1999b
14­
Middaugh
et
al
1987
15­
Yap­
Chiongco
1941
16­
Penttila
1997
17­
Pederson
et
al
1995
18­
Emmett
et
al
1991
19­
Hart
and
McHugh
1944
20­
C.
D.
F.
G.
1982
21­
Pentilla
1978
22­
Eschmeyer
and
Herald
1983
23­
Parente
and
Snyder
1970
Page
783
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
BO
BP
BQ
BR
BS
BT
1­
Moser
1996a
1­
Hart
1973
1­
Pederson
et
al
1995
1­
Hearne
1983
2­
Wang
1981
2­
Yap­
Chiongco
1941
2­
Emmett
et
al
1991
2­
Emmett
et
al
1991
3­
Stevens
and
Miller
1983
3­
Robinson
et
al
1968a
4­
Pearcy
and
Myers
1974
4­
Barraclough
et
al
1968
5­
Hay
et
al
1997
5­
Emmett
et
al
1991
6­
Pederson
et
al
1995
7­
Emmett
et
al
1991
mm
FL))
4
Page
784
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
BU
BV
BW
BX
BY
(
copepods
(
64
mm
FL))
7
1­
Wang
1981
1­
Hart
1973
1­?????????????

2­
Baxter
1999b
2­
Moyle
1992
2­
Emmett
et
al
1991
3­
Pearcy
and
Myers
1974
3­
Barraclough
1964
4­
Emmett
et
al
1991
4­
Robinson
et
al
1968a
5­
Barraclough
1964
5­
Barraclough
et
al
1968
6­
Eschmeyer
and
Herald
1983
6­
Emmett
et
al
1991
7­
Barraclough
1967a
8­
Barraclough
1967c
9­
Robinson
et
al
1968a
(
copepods,
fish
mysiids,
and
insects(
74­
80
mm
FL))
8
(
trematodes,
euphausiid
eggs,
cladocera,

copepoods,
fish
larvae,
and
fish)
9
Page
785
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
BZ
CA
CB
CC
CD
Caspian
terns
and
Fosters'
terns14
1­
Moser
1996a
1­
Hart
1973
1­
Brodeur
et
al
1987
1­
Dryfoos
1965
1­
Hart
1973
2­
Hart
1973
2­
Love
1996
2­
Emmett
et
al
1991
2­
Olesiuk
et
al
1990
3­
Love
1996
3­
Fitch
and
Lavenberg
1971
3­
Love
1996
4­
Moulton
1974
4­
Moyle1992
4­
Fitch
and
Lavenberg
1971
5­
Moyle
1992
5­
Fitch
and
Lavenberg
1975
5­
Fitch
and
Lavenberg
1975
6­
Hearne
1983
6­
Moyle
1976
6­
Moyle
1992
7­
Moyle
1976
7­
Brodeur
et
al
1987
7­
Barraclough
1964
8­
Eschmeyer
and
Herald
1983
8­
Dryfoos
1965
8­
Davoren
and
Burger
1999
9­
Barraclough
and
Fulton
1968
9­
Moulton
1970
10­
Emmett
et
al
1991
10­
Treacy
1985
11­
Barraclough
1967a
11­
Hay
1998
12­
Emmett
et
al
1991
13­
Hart
and
McHugh
1944
14­
Harvey
1982
(
copepods,
fish,
such
as
Theragra
chalcogrammus
and
Stichaeidae)
11
and
striped
bass),
birds,
and
marine
mammals12
(
planktonic
crustaceans,
including
amphipods,

euphausiids,
copepods,
crustacean
larvae,
and
some
larval
fish)
10
Page
786
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
CE
CF
CG
CH
CI
CJ
CK
1­
Emmett
et
al
1991
Page
787
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
CL
CM
CN
CO
CP
CQ
CR
CS
Page
788
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
CT
CU
CV
CW
CX
CY
CZ
Page
789
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
DA
DB
DC
DD
DE
1­
Baxter
1999b
1­
Dryfoos
1965
1­
Stevens
and
Miller
1983
1­
Delacy
and
Batts
1963
2­
Dryfoos
1965
2­
C.
D.
F.
G.
1982
2­
Hay
1998
3­
Moulton
1970
3­
Love
and
Moser
1983
3­
Emmett
et
al
1991
4­
Arai
1969
Page
790
of
1167
5­
2220e
­
Osmeridae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
1­
Love
1996
2­
Fitch
and
Lavenberg
1971
3­
Fitch
and
Lavenberg
1975
4­
Moyle
1992
5­
Breder
and
Rosen
1966
6­
Emmett
et
al
1991
7­
CDFG
1982
8­
Schaefer
1936
Page
791
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
A
B
C
D
E
Page
792
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
F
G
H
I
JKL
Page
793
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
M
N
O
P
Q
R
S
T
U
V
W
Page
794
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
X
Y
Z
AA
AB
AC
Page
795
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
AD
AE
AF
AG
AH
AI
AJ
Page
796
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
AK
AL
AM
AN
AO
AP
AQ
AR
AS
AT
Page
797
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
Page
798
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
BE
BF
BG
BH
Page
799
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
BI
BJ
BK
BL
BM
BN
Page
800
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
BO
BP
BQ
BR
BS
BT
Page
801
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
BU
BV
BW
BX
BY
Page
802
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
BZ
CA
CB
CC
CD
Page
803
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
CE
CF
CG
CH
CI
CJ
CK
Page
804
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
CL
CM
CN
CO
CP
CQ
CR
CS
Page
805
of
1167
5­
2220e
­
Osmeridae
107
108
109
110
111
112
113
114
115
116
CT
CU
CV
CW
CX
CY
CZ
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806
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Osmeridae
107
108
109
110
111
112
113
114
115
116
DA
DB
DC
DD
DE
Page
807
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1167
5­
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Osmeridae
107
108
109
110
111
112
113
114
115
116
DF
DG
DH
DI
DJ
DK
DL
DM
DN
DO
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Osmeridae
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5
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Polyprionidae1,3
Stereolepis
gigas
1
15.5
to
46
Percichthyidae2
1­
Roberts
1986
1­
Eschmeyer
1998
1­
Robins
et
al.
1991
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Robins
et
al.
1991
2­
Gates
&
Frey
1974
3­
Eschmeyer
1998
3­
Roedel
1948
TAXONOMY
RANGE
giant
sea
bass;
black
sea
bass;
jewfish;
giant
bass;
California
jewfish;
California
black
sea
bass1,2,3
n.
CA
(
Humboldt
Bay)
to
S.
Mexico
(
Oaxaca);
Japan1
Page
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Polyprionidae
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5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
F
G
H
I
J
K
L
M
N
O
P
Length
weight
equation
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
NF
NF
1
22.6
NF
NF
NF
NF
NF
NF
NF
NF
1­
Love
et
al
2000
AGE
&
GROWTH
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
Page
810
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2220e
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Polyprionidae
1234
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
Q
R
S
T
U
V
W
X
Y
Z
AA
AB
AC
to
(
SE)
Age
composition
Female
Both
Male
Female
Both
eing
structure
/
Met
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
NF
NF
NF
NF
NF
NF
NF
NF
NF
1,3,4
72
to
75
NF
NF
NF
2
90
to
100
1­
Fitch
&
Lavenberg
1971
2­
CDFG
1972
3­
Fitch
1960
4­
Love
1996
r2
Page
811
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Polyprionidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
AD
AE
AF
AG
AH
AI
AJ
AK
AL
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Oviparous1
NF
NF
NF
1,3,5,6
NG
(
11
to
13)
NG
(
11
to
13)
NF
NF
NF
2NG
(
NG)
NG
(
7
to
8)
NF
NF
NF
4
NG
(
11
to
12)­
sex
unspecified
NG
NG
NG
1­
Shane
et
al.
1996
1­
Feder
et
al
1974
2­
Crooke
1992a
3­
Fitch
&
Lavenberg
1971
4­
CDFG
1982
5­
Fitch
1960
6­
Love
1996
1st
maturity
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
50%
maturity
Breeding
Behavior
Sexual
dimorphism
Page
812
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1167
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2220e
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Polyprionidae
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
state
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.

NF
NF
NF
California1,2,3,4,5
NF
NF
NG
NG
1­
Crooke
1992a
2­
Fitch
&
Lavenberg
1971
3­
CDFG
1982
4­
Fitch
1960
5­
Love
1996
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
REPRODUCTION
Page
813
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Polyprionidae
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5
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
BB
BC
BD
BE
BF
BG
BH
BI
es
present
Spawning
Frequency
Size
specific
fecundity
equation
Size
at
settlement
Dec.
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
Larval
duration
(
SL
in
mm)

NF
NF
1,2,3,4
60
million
(~
50
yrs.)
NF
NF
NF
NF
1­
Crooke
1992a
2­
Fitch
&
Lavenberg
1971
3­
Fitch
1960
4­
Love
1996
DISPERSAL
&
RECRUITME
Fecundity
range
in
#
eggs
(@

cm,
grams,
or
age)
Fecundity
Mean
(
range)
fecundity
in
eggs/
g
female
wt.
Page
814
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
BJ
BK
BL
BM
BN
BO
BP
BQ
BR
BS
Size
Fish
Size
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
(
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
NF
11.5
to
1.6
planktonic1,2
NF
13.0
to
18.8
pelagic1
NF
NF
NF
NF
1­
Shane
et
al.
1996
1­
Shane
et
al.
1996
1­
Shane
et
al.
1996
1­
Shane
et
al.
1996
2­
Limbaugh
1955
NT
Food
habits
Eggs
Larvae
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815
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Polyprionidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
BT
BU
BV
BW
BX
BY
BZ
Fish
Size
Food
habits
(
SL
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Habitat
118.8
to
maturity
carnivore
NF
NF
NF
(
anchovy,
sardine,
and
squid)
1,2,3
1­
Shane
et
al.
1996
1­
Crooke
1992a
1­
Crooke
1992a
1­
Feder
et
al
1974
2­
Love
1996
2­
Fitch
&
Lavenberg
1971
2­
Crooke
1992a
3­
Fitch
1960
3­
Carlisle
et
al.
1964
4­
Turner
et
al.
1969
5­
Fitch
&
Lavenberg
1971
6­
Limbaugh
1955
7­
CDFG
1982
8­
Love
1996
kelp
beds;
sandy
bottoms;
hard
bottoms1,4
rocky
bottoms
near
kelp
beds;

artificial
reefs;
smaller
individuals
over
sandy
bottoms1,2,3,4,5,6,7,8
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Juveniles
Life
Stage
Page
816
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Polyprionidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
CA
CB
CC
CD
CE
CF
CG
CH
CI
Adults
Genetic
variance
Food
habits
Haplotype
diversity
Mean
number
of
%
Polymorphic
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
(
mega)
carnivore3
NF
NF
large
sharks,
marine
mammals1,2,3
NF
NF
NF
NF
NF
1­
Feder
et
al
1974
1­
Crooke
1992a
2­
Crooke
1992a
2­
Fitch
&
Lavenberg
1971
3­
Quast
1968a
3­
Fitch
1960
4­
Fitch
&
Lavenberg
1971
5­
Limbaugh
1995
6­
Winget
1968
7­
Fitch
1960
8­
Love
1996
(
spiny
lobster;
squid;
fishes,
such
as
kelp
bass,
sheephead,
mackerel,

bonito,
and
small
sharks)
1,2,3,4,5,6,7,8
GENETICS
Nucleotide
diversity
(#)

Page
817
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2220e
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Polyprionidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
CJ
CK
CL
CM
CN
CO
CP
CQ
CR
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Total
Length
(
age)
Fishery
/
Total
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
NF
NF
NF
NF
NF
NF
NF
NF
NF
Mean
heterozygosity
Page
818
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Polyprionidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
CS
CT
CU
CV
CW
CX
CY
CZ
DA
DB
DC
Mortality
Estimates
Rate
of
Exploitation
Method
of
ge
range
use
Rate
of
Exploitation
Interactions
/
associations
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimationalculate
mort
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
NF
NF
NF
NF
NF
NF
NF
NF
NF
cleaned
by
Oxyjulis
californica
1,2,3
NF
1­
Limbaugh
1955
2­
Hobson
1971
3­
Limbaugh
1961
4­
Love
&
Moser
1983
infested
by
parasites
from
the
families
Dasyrhynchidae,
Caligidae,

Cymothoidae,
and
Diplectanidae4
MORTALITY
Page
819
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
DD
DE
DF
DG
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
Transient1
NF
Yes1,2
NF
1­
Turner
et
al.
1969
1­
Feder
et
al
1974
2­
Limbaugh
1955
3­
CDFG
1982
Yes,
during
breeding
season3
BEHAVIOR
Movement
Page
820
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Polyprionidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Pomacentridae1
Chromis
punctipinnis
1,2
1inshore
to
61
Hypsypops
rubicundus
1,2
1intertidal
to
29
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Love
et
al.
2000
1­
Love
et
al.
2000
2­
Eschmeyer
1998
2­
Gates
&
Frey
1974
3­
Roedel
1948
garibaldi;
ocean
goldfish;
garibaldi
perch;
golden
perch
1,2,3
central
California
(
Monterey
Bay)
to
central
Baja
California
(
Punta
San
Pablo)
1
central
California
(
Monterey
Bay)
to
southern
Baja
California
(
Bahia
Magdalena)
1
blacksmith;
perch;
blue
perch;
kelp
perch;
rock
bass;
black
perch1,2,3
TAXONOMY
RANGE
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
F
G
H
I
J
K
L
M
N
O
P
Length
weight
equation
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
1*
Both:
W=
0.0191TL3.178
(
r2=
0.99)
NF
1
30
NF
NF
NF
NF
NF
NF
NF
NF
2Both:
W=
0.0000185SL3.09
1Both:*
W=
0.1099TL3.077
(
r2=
0.96)
NF
1
36
NF
NF
NF
NF
NF
NF
NF
NF
2Both:
W=
0.000036SL3.10
1­
DeMartini
et
al.
1994
1­
Love
et
al.
2000
2­
Quast
1968b
AGE
&
GROWTH
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
Page
822
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
Q
R
S
T
U
V
W
X
Y
Z
AA
AB
to
(
SE)
Age
composition
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Dioecious
NF
NF
NF
NF
NF
NF
NF
NF
1,2
1,4
13
NF
Dioecious
2
17;
29
in
captivity
3
12
5
up
to
17
1­
DeMartini
et
al.
1994
1­
Limbaugh
1955
2­
Clarke
1970
2­
Fitch
&
Lavenberg
1975
3­
Clarke
1971
4­
Neal
1993
5­
Clarke
1970
r2
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823
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Pomacentridae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AC
AD
AE
AF
AG
Mating
system
Mode
of
reproduction
Parental
Care
Male
vs.
Female
length
Secondary
Characteristics
Promiscuous1,2
Oviparous1,2,4
NF
Promiscuous3,4,5,6,7,8,9,11
Oviparous1,2,3
1Males>
Females
males
more
reddish
than
females2
Polygynous12
1­
Fitch
and
Lavenberg
1971
1­
Watson
1996e
1­
Limbaugh
1955
1­
Limbaugh
1964
1­
Turner
&
Ebert
1962
2­
Turner
&
Ebert
1962
2­
Turner
&
Ebert
1962
2­
Watson
1996e
2­
Limbaugh
1964
3­
Fitch
&
Lavenberg
1975
3­
Limbaugh
1964
3­
Fitch
&
Lavenberg
1971
3­
Sikkel
1993
4­
Sikkel
1988
4­
Love
1996
4­
Turner
&
Ebert
1962
5­
Alcalay
&
Sikkel
1994
5­
Limbaugh
1964
6­
Sikkel
1995a
6­
Hobson
1971
7­
Sikkel
1995b
7­
Pondella
&
Stephens
1994
8­
Knapp
et
al.
1995
8­
Fitch
&
Lavenberg
1975
9­
Pankhurst
1995
9­
CDFG
1982
10­
Sikkel
1998
10­
Clarke
1971
nest­
guarding
males
turn
a
pale,
mottled
grey,

with
dark
bands
dorsal
and
anterior
to
the
eyes1
Breeding
Behavior
Sexual
dimorphism
females
have
visibly
swollen
abdomens
during
breeding
season3
males
are
nest­
builders;
guard
eggs
for
a
few
weeks1,2,3,4,5,9,11,23,24
17,18nest
size
about
300
to
3000
cm2
males
are
nest­
builders
and
guarders;
farm
algae
within
nest1,2,4,5,6,7,8,10,12,13,14,15,16,17,18,1
9,20,21,22,23,24
Female
mate
choice
determined
by
visitng
the
nest
of
multiple
males;
choice
affected
by
nests
with
eggs
in
the
early
stages
(
yellow­
colored,
as
opposed
to
gray)
of
development6,7,10,11,12
Page
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12
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15
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AH
AI
AJ
AK
AL
AM
AN
AO
AP
AQ
AR
AS
AT
Fertilized
eggs
present
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
1,2,3,4,7,8
14
(
2)­
unspecified
sex
NF
NG
(
NG)
NG
(
NG)
NG
NG
NG
NF
California2,3
s.
California1
1,4
20.3
(
3)­
unspecified
sex
20.3
(
3)­
unspecified
sex
NG
NG
NG
NF
s.
California1,3,4,5,6,7,8,9,10
5
NG
(
5)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NG
NG
NG
6,8
21.7
(
6)
21.8
(
6)
NG
(
NG)
NG
(
NG)
scales
NG
NG
1­
Limbaugh
1955
1­
Limbaugh
1955
2­
Fitch
&
Lavenberg
1971
2­
Fitch
&
Lavenberg
1971
3­
Turner
&
Ebert
1962
3­
Limbaugh
1964
4­
Limbaugh
1964
4­
Fitch
&
Laveneberg
1975
5­
Fitch
&
Lavenberg
1975
5­
Clarke
1971
6­
Clarke
1970
6­
Sikkel
1988
7­
Bray
1978
7­
Neal
1993
8­
Love
1996
8­
Foster
1972
9­
Clarke
1970
10­
Feder
et
al.
1974
50%
maturity
REPRODUCTION
1st
maturity
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
Fertilization
and
Spawning
Period
(^^^^^=
peak
fe
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Spawning
females
present
Both
reproductive
states
present
Spawning
Frequency
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
(
per
year)

NF
NF
NF
Multiple1,2,3,4,5,6
1
35
±
13.2
(
NG)
1
15,000
(
NG)
to
80,000
(
NG)

1­
Sikkel
1995a
1­
DeMartini
et
al.
1994
1­
Fitch
&
Lavenberg
1975
2­
Knapp
et
al.
1995
2­
Clarke
1970
3­
DeMartini
et
al.
1994
3­
Love
1996
4­
Sikkel
1998
5­
Sikkel
1994
6­
Sikkel
1993
7­
Clarke
1970
8­
Sikkel
1989
Fecundity
Mean
(
range)
fecundity
in
eggs/
g
female
wt.
Fecundity
range
in
#
eggs
(@

cm,
grams,
or
age)

ertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
females
may
spawn
only
once
a
year;
males
up
to
7
times
7
males
may
finish
up
to
four
brood
cycles
per
season4,8
2
129,000
(
34,000
to
190,000)

per
male
nest
clutch
1
12,546
±
5,294
(
NG)
eggs
per
batch
males
every
6
days
during
breeding
season;
24
per
year3
2,3
44,000
(
15,000
to
88,000)

eggs
per
batch
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11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BF
BG
BH
BI
BJ
BK
BL
Size
specific
fecundity
equation
Size
at
settlement
Size
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
Larval
duration
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
NF
NF
NF
2
15
to
20
NF
11.1
to
1.3
X
0.6
to
0.7
2
1.22
(
1.14
to
1.32)

3
1.5
NF
1,3,4,6,7,8,9,11
2
to
3
weeks
NF
1
20.5
NF
3
1
to
2
2,5,10
12
to
23
days
4,5
1
X
2
1­
Fitch
&
Lavenberg
1975
1­
Clarke
1970
1­
Watson
1996e
1­
Watson
1996e
2­
Sikkel
1988
2­
Bray
1978
2­
Turner
&
Ebert
1962
2­
Turner
&
Ebert
1962
3­
Alcalay
&
Sikkel
1994
3­
Limbaugh
1964
3­
Limbaugh
1964
4­
Sikkel
1995b
4­
Sikkel
1988
4­
CDF
&
G
1982
5­
Knapp
et
al.
1995
5­
Clarke
1970
5­
Sikkel
1988
6­
Sikkel
1998
6­
Alcalay
&
Sikkel
1994
7­
Sikkel
1994
7­
Sikkel
1995a
8­
Sikkel
1993
8­
Knapp
et
al.
1995
9­
Clarke
1970
10­
Sikkel
1989
demersal,
attached
to
substrate
in
a
nest
via
adhesive
filaments1,2,3,4
Eggs
DISPERSAL
&
RECRUITMENT
demersal,
attached
to
substrate
in
a
nest
via
adhesive
filaments1,3,5,6,7,8
hatching
occurs
from
dusk
to
dawn3
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BM
BN
BO
BP
BQ
BR
BS
BT
Fish
Size
Fish
Size
Predators
(
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
(
SL
mm)

12.7
to
3.1
to
13.4
planktonic1
NF
NF
NF
blacksmith1
NF
2
3.06
to
some,
but
not
all
planktonic2
within
kelp
forest
cover3
beyond
4
to
5
km
of
shore5
some,
but
not
all
planktonic2
other
garibaldi1,3,4
12.9
to
3.5
to
9.0
planktonic
(
neuston)
1,6,7,8
NF
NF
NF
NF
NF
Lythrypnus
dalli
4
senorita
and
black
perch5
1­
Limbaugh
1955
1­
Watson
1996e
1­
Watson
1996e
1­
Turner
&
Ebert
1962
2­
Turner
&
Ebert
1962
2­
Turner
&
Ebert
1962
2­
Fitch
&
Laveneberg
1971
3­
Limbaugh
1964
3­
Turner
&
Ebert
1962
4­
Sikkel
1994
4­
Limbaugh
1964
5­
Clarke
1970
5­
Barnett
et
al.
1984
6­
Sikkel
1994
7­
Sikkel
1993
8­
Clarke
1970
Food
habits
open
ocean,
sometimes
entering
kelp
beds4
Larvae
blacksmiths;
senoritas;

sheephead;
garibaldis2
males
tend
to
cannabalized
eggs
deposited
on
older
broods
or
empty
nests4
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BU
BV
BW
BX
BY
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
carnivore/
planktivore1,4,7
sucker1
diurnal1,2,3,4
surfgrass
beds10
carnivores
NF
NF
kelp
bass4
deep
rock
pools9
surfgrass
beds10
1­
Limbaugh
1955
1­
Limbaugh
1955
1­
Hobson
&
Chess
1976
1­
Tegner
&
Dayton
1981
1­
Fitch
&
Lavenberg
1971
2­
Turner
&
Ebert
1962
2­
Bray
1981
2­
Bray
1981
2­
Bray
1981
3­
Limbaugh
1964
3­
Tegner
&
Dayton
1981
3­
Hobson
&
Chess
1976
3­
Love
&
Ebeling
1978
4­
Pondella
&
Stephens
1994
4­
Hartney
1996
4­
Bray
1978
4­
Neal
1993
5­
Coyer
1979
5­
Hobson
&
Chess
1976
5­
Bray
1978
6­
CDF
&
G
1982
6­
Clarke
1970
7­
Turner
et
al.
1969
7­
Bray
1978
8­
Hobson
&
Chess
1976
9­
Smith
1883
10­
DeMartini
1981
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Juveniles
Life
Stage
comorants;
adult
blacksmiths1
(
;

cyclopoid
copepods
and
cladocerans;
larvaceans;
bryozoan
larvae;
fish
eggs)
2,3,5
kelp
bass;
marine
mammals;
birds2,3
(
planktonic
crustacea,
such
as
copepods,
amphipods,
and
isopods)
6
kelp
bass;
olive
rockfish;
lingcod
5
neritic;
tide
pools;
rocky
reefs;
kelp
forests;
in
caves
and
crevices
3,4,12
open
ocean,
under
kelp
patties;

shallow
sandy
areas
with
rocks;
kelp
canopy
and
rocky
shores;
caves
and
crevices
1,2,3,4,5,6,7,8,11,12,13
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BZ
CA
CB
CC
Adults
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
carnivore;
planktivore4,8,9,13,17,19,22,23,24,25,32/
microcarnivore30
sucker2
diurnal1,2,3,5,7,8,9,10,12
pecker3
in
caves
and
crevices
at
night
3,7,8,12,14,15,19,32,34
omnivore7,26/
microcarnivore22
browser1
diurnal4,6,8,10,11,13
mornings6
scavange
sea
urchins
form
sheephead
attacks
2
1­
Limbaugh
1955
1­
Limbaugh
1955
1­
Davis
&
Anderson
1989
1­
Bray
et
al.
1986
2­
Fitch
&
Lavenberg
1971
2­
Nelson
&
Vance
1979
2­
Hobson
&
Chess
1976
2­
Hobson
&
Chess
1986
3­
Bray
et
al.
1986
3­
Fitch
&
Lavenberg
1971
3­
Green
&
McFarland
1994
3­
Bray
et
al.
1981
4­
Limbaugh
1964
4­
Bray
et
al.
1986
4­
Clarke
1971
5­
Stephens
et
al.
1984
5­
Turner
&
Ebert
1962
5­
Hartney
1996
6­
CDFG
1982
6­
Limbaugh
1964
6­
Sikkel
1995a
7­
Bray
et
al.
1981
7­
Davis
&
Anderson
1989
7­
Bray
1981
8­
Tricas
1979
8­
Pondella
&
Stephens
1994
8­
Hobson
et
al.
1981
9­
Carlisle
et
al.
1964
9­
Hobson
&
Chess
1986
9­
Ebeling,
Larson,
Alevizon,
&
Bray
1980
10­
Turner
et
al.
1969
10­
Fitch
&
Lavenberg
1975
10­
Ebeling
&
Bray
1976
(
small
fish;
squid;
crustaceans,
such
as
cladocerans
and
copepods;
larval
fish,
including
blacksmith;
fish
eggs,

including
blacksmith
eggs;
larvaceans)

1,3,5,6,11,14,17,18,20,23,28,31,32,33,34
kelp
forest;
rocky
areas,
using
holes
and
crevices
as
cover;
tide
pools;
surfgrass
beds;
artificial
reefs1,4,5,9,13,16,17,18,19,22,23,24,25,26,27,28,29,30,31,33,34
kelp
forest;
shallow
rocky
areas
and
breakwaters;

surfgrass
beds
(
rarely);
midwater
over
rocky
and
artificial
reefs
1,2,4,5,6,9,10,11,12,15,16,17,18,19,20,21,22,30,31,32,33,34
(
bryozoans;
sea
anemones;
sponges;
polychaetes;
small
crustaceans
and
gooseneck
barnacles;
gastropod
eggs;

bivalves;
fish
eggs,
such
as
garibaldi
and
blacksmith
eggs)
1,5,6,10,12,15,16,21,27,28,29,31,33,34,35
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CD
CE
CF
CG
CH
CI
CJ
CK
CL
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
sea
lions1,10
NF
NF
NF
NF
NF
1,2
NG
0.009
allozymes
harbor
seals4
blue
sharks5
None2
NF
NF
NF
NF
NF
NF
NF
NF
giant
sea
bass;
sharks;
sea
lions
3
harbor
seals7
harbor
seals
ans
sea
lions8
1­
Fitch
&
Lavenberg
1971
1­
Waples
&
Rosenblatt
1987
2­
Limbaugh
1964
2­
Waples
1987
3­
Fitch
&
Lavenebrg
1975
4­
Haaker
et
al.
1984
5­
Tricas
1979
6­
Hubbs
et
al.
1970
7­
Sikkel
1995aa
8­
Neal
1993
9­
Clarke
1970
10­
Hawes
1983
Nucleotide
diversity
(#)

sea
lions;
groupers;
sharks;
moray
eels9
kelp
bass;
moray
eels;
lingcod;

Brandt's
and
double­
breasted
cormorants;
bald
eagles;
least
terns11
Mean
heterozygosity
GENETICS
moray
eels;
kelp
bass;
lingcod;

Brandt's
comorants2,3,6
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12
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CM
CN
CO
CP
CQ
CR
CS
CT
CU
CV
CW
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
First
Recruitment
Full
Recruitment
Method
of
Effective
population
size
Total
length
(
age)
Total
length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTA
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CX
CY
CZ
DA
DB
DC
Rate
of
Exploitation
Age
range
used
Rate
of
Exploitation
Interactions
/
associations
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
NF
NF
NF
NF
forms
mixed
aggragations
with
Sebastes
mystinus
10
clean
ectoparasites
from
Mola
mola
8
NF
NF
NF
young
associated
with
red
sea
urchins3
NF
1­
Sikkel
1995a
1­
Limbaugh
1955
1­
Bray
1981
2­
Sikkel
1998
2­
Bray
et
al.
1986
2­
Hobson
et
al.
1981
3­
Clarke
1970
3­
Limbaugh
1964
3­
Hobson
&
Chess
1976
4­
Hobson
1971
4­
Green
&
McFarland
1994
5­
Bray
et
al.
1981
5­
Bray
1978
6­
Sigalit
&
Rainboth
1981
7­
Clarke
1971
8­
Turner
et
al.
1969
9­
Love
et
al.
1981
10­
Ebeling
&
Bray
1976
share
territories
with
moray
eels
and
rockfish,
such
as
gopher,
kelp,
and
treefish3
infested
with
ectoparasites
from
the
families
Bomolochidae
and
Caligidae;
also
gnathiid
larvae
and
monogenetic
trematodes
4
agonsitic
towards
territory
invaders
such
as
labrids,

embiotocids,
kyphosids,
scorpaenids,
and
pomacentrids,

including
both
juvenile
and
adult
garibaldis7,12,14,17
cultivate
living
turf­
forming
algae,
such
as
Tiffaniella
and
Murrayellopsis
16
return
trips
between
spawning
bouts
allows
females
to
expell
food
competitors
form
their
territory2
competes
with
Girella
nigricans
,
Medialuna
californiensis
,
Embiotoca
jacksoni
,

Halichoeres
semicinctus
,
and
Semicossyphus
pulcher
for
food1,2
supply
young
Macrocystis
with
ammonium
from
nighttime
excretions2
infested
with
ectoparasites
from
the
families
Caligidae
and
Lerneopodidae;
also
gnathiid
larvae
4
infested
by
parasites
from
the
familes
Caligidae,

Lernaeopodidae,
Gnathiid
larvae,
Microcotylidae,
and
Ceratomyxidae18
infested
by
parasites
from
the
families
Bomolochidae,

Caligidae,
Pennellidae,
Hemiuridae,
and
Gnathiid
larvae18
cleaned
by
fishes
such
as
Rhacochilus
vacca
and
Oxyjulis
califonica
1,3,4,8,15,20
external
parasites
removed
by
Oxyjulis
californica,

Brachyistius
frenatus,
and
the
shrimp
Hippolysmata
californica
3,6,15,20
infected
by
a
possible
species­
specific
bacteria
Vibrio
damsela
9
may
supply
organic
carbon,
via
nightly
defecation,
to
benthic
reefs
organisms5,19
ALITY
adults
have
daily
movements
from
sleeping
crevices
to
the
incurrent
side
of
reef
to
forage1,4,5
daily
movement
from
diurnal
feeding
spots
to
nocturnal
sleeping
holes2,3
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants11,13
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
DD
DE
DF
DG
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
Residential2,3,4
Yes­
to
same
shelter
each
night
1,3
Yes2
Yes2,3,4,6,7,8,10,11,12,13,14,15,16,17,19,21,22
Residential1,2,3,5
Yes­
back
to
their
territories4
solitary5,12,14
forms
occasional
aggregations9
Yes2,4,5,6,9,16,17,20,23,26,27
25territory
size
10
to
15
m2
1­
Limbaugh
1955
1­
Bray
et
al.
1981
1­
Limbaugh
1955
1­
Limbaugh
1955
2­
Larson
&
DeMartini
1984
2­
Hartney
1996
2­
Watson
1996e
2­
Watson
1996e
3­
Stephens
et
al.
1994
3­
Ebeling
&
Bray
1976
3­
Fitch
&
Lavenberg
1971
3­
Turner
&
Ebert
1962
4­
Helvey
1985
4­
Clarke
1970
4­
Limbaugh
1964
4­
Limbaugh
1964
5­
Clarke
1970
5­
Hobson
1971
5­
Hobson
1971
6­
Pondella
&
Stephens
1994
6­
Davis
&
Anderson
1989
7­
CDFG
1982
7­
Sigalit
&
Rainboth
1992
8­
Tricas
1979
8­
Clarke
1971
9­
Clarke
1971
9­
Stephens
et
al.
1994
10­
Carlisle
et
al.
1964
10­
Sikkel
1988
Movement
some
non­
territorial
"
floaters"
or
"
wanderers"
18,24
areas
defended
include
grazing
area,

shelter,
and
nest
site;
adult
males
most
strictly
territorial8,11,12,13,15,19,21,22,24
both
sexes
maintain
territories
yearround
males
more
aggrssive
towards
adults
during
breeding
season,
to
juveniles
the
rest
of
the
year
7,8,10,11,12,13,14,15,18,19,21,22,24,25
Yes,
during
breeding
season
(

eggguarding
1,3
Yes,
during
breeding
season
(

eggguarding
1,4
Yes­
schools
have
more
individuals
at
the
incurrent
end
of
the
school
than
the
rear
end;
individuals
tend
to
move
to
the
incurrent
front
end
of
the
school
18,20
BEHAVIOR
YOY­
Yes;
older
yearlings
more
solitary;
adults
found
aggregating
1
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834
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57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
A
B
C
D
E
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835
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
F
G
H
I
J
K
L
M
N
O
P
Page
836
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
Q
R
S
T
U
V
W
X
Y
Z
AA
AB
Page
837
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
AC
AD
AE
AF
AG
11­
Sikkel
1994
11­
Turner
et
al.
1969
12­
Sikkel
1989
12­
Sikkel
1988
13­
Alcalay
&
Sikkel
1994
14­
Sikkel
1995a
15­
Sikkel
1995b
16­
Knapp
et
al.
1995
17­
Sikkel
1998
18­
Sikkel
1998
19­
Sikkel
1993
20­
Foster
1972
21­
Clarke
1970
22­
Sikkel
1989
23­
Feder
et
al.
1974
24­
Love
1996
Page
838
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
AH
AI
AJ
AK
AL
AM
AN
AO
AP
AQ
AR
AS
AT
Page
839
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Page
840
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
BF
BG
BH
BI
BJ
BK
BL
11­
Love
1996
Page
841
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
BM
BN
BO
BP
BQ
BR
BS
BT
Page
842
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
BU
BV
BW
BX
BY
11­
Kritzler
et
al.
1950
12­
Feder
et
al.
1974
13­
Love
1996
Page
843
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
BZ
CA
CB
CC
11­
Stephens
et
al.
1994
11­
Bray
et
al.
1981
11­
Clarke
1970
12­
Hartney
1996
12­
Clarke
1971
12­
Bray
1978
13­
Sikkel
1995b
13­
Hartney
1996
13­
Love
1996
14­
Bray
1981
14­
Love
et
al.
1981
15­
Shrode
et
al.
1982
15­
Sikkel
1988
16­
Allen
1985
16­
Sikkel
1995a
17­
Ebeling,
Larson,
Alevizon,
&
Bray
1980
17­
Bray
1981
18­
Johnson
et
al.
1994
18­
Shrode
et
al.
1982
19­
Ebeling
&
Bray
1976
19­
Ebeling,
Larson,
Alevizon,
&
Bray
1980
20­
Hobson
&
Chess
1976
20­
Helvey
1985
21­
Green
&
McFarland
1994
21­
Johnson
et
al.
1994
22­
DeMartini
1981
22­
Ebeling
&
Bray
1976
23­
Sikkel
1998
23­
Hobson
&
Chess
1976
24­
Sikkel
1994
24­
Green
&
McFarland
1994
25­
Kritzler
et
al.
1950
25­
Hose
&
Stoffel
1980
26­
Sikkel
1993
26­
Sikkel
1998
27­
Neal
1993
27­
Kritzler
et
al.
1950
28­
Norris
1963
28­
Holbrook
et
al.
1997
29­
Clarke
1970
29­
Clarke
1970
30­
Ebeling,
Larson,
&
Alevizon
1980
30­
Quast
1968a
31­
Quast
1968a
31­
Quast
1968d
32­
Bray
1978
32­
Bray
1978
33­
Feder
et
al.
1974
33­
Feder
et
al.
1974
34­
Love
1996
34­
Love
1996
35­
Mitchell
1953
Page
844
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1167
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­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
CD
CE
CF
CG
CH
CI
CJ
CK
CL
11­
Love
1996
Page
845
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
CM
CN
CO
CP
CQ
CR
CS
CT
CU
CV
CW
Page
846
of
1167
5­
2220e
­
Pomacentridae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
CX
CY
CZ
DA
DB
DC
11­
Hose
&
Stoffel
1980
12­
Sikkel
1998
13­
Hose
et
al.
1983
14­
Neal
1993
15­
Limbaugh
19??

16­
Foster
1972
17­
Clarke
1970
18­
Love
&
Moser
1983
19­
Bray
1978
20­
Feder
et
al.
1974
Page
847
of
1167
5­
2220e
­
Pomacentridae
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55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
DD
DE
DF
DG
11­
Turner
et
al.
1969
11­
Alcalay
&
Sikkel
1994
12­
Love
et
al.
1981
12­
Sikkel
1995a
13­
Bray
1981
13­
Sikkel
1995b
14­
Hobson
et
al.
1981
14­
Knapp
et
al.
1995
15­
Ebeling,
Larson,
Alevizon,
&
Bray
1980
15­
Pankhurst
1995
16­
Ebeling
&
Bray
1976
16­
Hobson
et
al.
1981
17­
Hobson
&
Chess
1976
17­
DeMartini
1981
18­
Green
&
McFarland
1994
18­
Sikkel
1998
19­
Hose
&
Stoffel
1980
19­
Sikkel
1994
20­
Bray
1978
20­
Kritzler
et
al.
1950
21­
Feder
et
al.
1974
21­
Sikkel
1993
22­
Love
1996
22­
Neal
1993
23­
Foster
1972
24­
Clarke
1970
25­
Sikkel
1989
26­
Feder
et
al.
1974
27­
Love
1996
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848
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­
Pomacentridae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Sciaenidae1
Atractoscion
nobilis
1,2
Cynoscion
nobilis
3
Genyonemus
lineatus
1,2
Menticirrhus
undulatus
1,2
1surf
to
45
Roncador
stearnsi
1,2
1surf
to
18
1inshore
to
236
California
(
Point
Conception)
to
the
Gulf
of
California1
California
(
Point
Conception)
to
Mexico
(
Mazatlan),
including
the
Gulf
of
California1
Calfiornia
corbina;
California
whiting;

corvina;
surf
fish;
bagre;
sucker;
corbina
1,2,4
spotfin
croaker;
spot;
golden
croaker;
surf
fish;
roncador
1,2,4
white
seabass;
sea
trout
(
juveniles);

weakfish;
king
croaker;
white
croaker
1,2,4,5
white
croaker;
tomcod;
kingfish3;
shiner;

chenfish;
herring;
Pasadena
trout;
carbinette;

roncador;
tommy;
cognard;
little
bass;
little
roncador;
tommy
croaker
1,2,3,4,5
1inshore
to
122
southeastern
Alaska
(
Juneau)
to
southern
Baja
California
(
Bahia
Magdalena)
and
northern
Gulf
of
California1
British
Columbia
(
Barkley
Sound)
to
southern
Baja
California
(
Bahia
Magdalena)
1
TAXONOMY
RANGE
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12
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
F
G
H
I
JK
L
Length
weight
equation
Max.
reported
Geographic
area
Linf
in
cm
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
1*
Both:
W=
0.00001549TL2.92
(
round
wt.)
NF
1
152
California
(
TL)
1
NG
NG
14.65
(
95%
C.
I.=
1.02)

1*
Both:
W=
0.00001227TL2.94
(
dressed
head­
on
wt.)

3Both:
W=
0.000000542TL3.315
6
Both:
Dry
W=
0.0111SL4.495
larvae
only
6
Both:
Dry
W=
0.0131SL4.466
larvae
only
6
Both:
Dry
W=
0.0125SL4.476
larvae
only
7
Both:
Dry
W=
0.245SL3.668
larvae
only
7
Both:
Dry
W=
0.0012SL5.152
larvae
only
8
Both:
Dry
W
(
mg)=
0.000000004SL4.426
larvae
<
21
days
only
8
Both:
Dry
W
(
mg)=
0.00000286SL3.07
larvae
21
to
60
days
only
8
Both:
Dry
W
(
mg)=
0.00000989SL2.87
larvae
100
to
190
days
only
8
Both:
Dry
W=
0.0005SL2.91
fish
128
to
344
mm
only
8
Both:
Dry
W=
0.000875SL2.79
fish
439
to
2090
mm
only
4Males:
W=
0.0111TL3.011
(
r2=
0.951)
1TL=
3.01SL1.203
(
SE=
2.942)
1
41
s.
California
(
TL)
2
59.17
(
0.29)
60.72
(
0.23)
NG
4Females:
W=
0.0109TL3.024
(
r2=
0.967)
1TL=
0.338FL1.03
(
SE=
4.417)

5Both:
W=
0.0000272SL2.943
1SL=
­
2.253TL0.83
(
SE=
2.444)

5Both:
W=
0.000033TL2.804
1SL=
­
5.019FL0.879
(
SE=
1.832)

1FL=
0.698TL0.966
(
SE=
4.417)

1FL=
5.868SL1.137
(
SE=
2.084)

3TL=
0.892
+
1.19SL
3TL=
0.023
+
1.04FL
3SL=
0.442
+
0.79TL
3SL=
0.379
+
0.82FL
3FL=
0.088
+
0.96TL
3FL=
0.849
+
1.14SL
4TL=
4.2
+
1.1933SL
4Both:
W=
0.000001133SL3.068
2SL=
0.852TL­
4.72
1
83.6
NF
NF
NF
NF
4Both:
W=
0.0000321SL2.944
1TL=
22.25SL1.131
(
SE=
4.13)
1
70
NF
NF
NF
NF
1TL=
4.852FL1.022
(
SE=
4.56)

1SL=
­
18.66TL0.881
(
SE=
3.64)

Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)
equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
Page
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
M
N
O
P
Q
R
S
T
U
V
W
X
Growth
Parameters
k
(
SE)
to
(
SE)

Male
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
NG
NG
0.128
(
95%=
0.021)
NG
NG
­
0.281
NG
NG
NG
scales
No
interscale
comparisons
0.033
(
0.03)
0.037
(
0.02)
NG
­
8.66
(
1.3)
­
7.54
(
1.1)
NG
NG
NG
NG
otoliths;
section
No
intrareader
comparison
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
AGE
&
GROWTH
r2
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
Y
Z
AA
AB
AC
AD
AE
Age
composition
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
10
>
20
NF
dioecious1
NF
oviparous1
NF
4,5­
larval
growth
1,4,5,8,9
12
to
15
1
dioecious1
NF
oviparous1,2
NF
6
12
7­
Larval
10
13
1­
monthly
and
annual
growth
2,8>
8
NF
NF
NF
oviparous
NF
1­
monthly
and
annual
growth
3,7,8,10
at
least
15
NF
NF
NF
oviparous
NF
Breeding
Behavior
6­
larval
growth
fitted
to
a
Laird­
Gompertz
growth
curve
3­
otoliths
used
to
give
size
ranges
for
age
groups
2­
juvenile
growth
fit
to
Gompertz,
validated
by
larval
rearing
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11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AF
AG
AH
AI
AJ
AK
AL
Male
vs.
Female
length
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
NF
Females
may
develop
dark
vertical
bars
during
breeding
season
3
1,17
NG(
NG)
NG(
NG)
61
(
NG)
71
(
NG)
NA
2,10>
50
(
NG)
60
(
NG)
60
(
NG)
70
(
NG)
NA
3,551
(
NG)
61
(
NG)
NG
(
NG)
NG
(
4)
NG
4NG
(
NG)
NG
(
3)
NG
(
NG)
NG
(
4)
NG
7
51
(
3)
61
(
4)
81.3
(
6)
81.3
(
6)
NG
1551
(
2)
61
(
3)
61
(
NG)
71
(
NG)
NG
1651
(
NG)
NG
(
NG)
NG
(
NG)
79
(
5
to
6)
NG
1,2Males<
Females
None2
1,2,1613
to
15
(
NG)­
unspecified
sex
NG
(
NG)
NG
(
NG)
NA
3,11
12
(<
1)
13
(<
1)
14
(
1)
15
(
1)
otoliths;
section
9,17
NG
(
NG)
NG
(
NG)
14
(
1)
15.2
(
1)
NG
1214.7
to
16.4
(
3
to
4)­
unspecified
sex
NG
(
NG)
NG
(
NG)
otoliths
NF
NF
223
(
NG)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NA
4,13,16,17
NG
(
NG)
NG
(
NG)
25
(
2)
33
(
3)
NG
NF
2NG
(
NG)
23
to
24
(
NG)
NG
(
NG)
NG
(
NG)
NA
8,14,17
22.9
(
2)
NG
(
NG)
NG
(
NG)
31.8
(
3)
NG
large
males
ate
golden
in
color1
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
1st
maturity
50%
maturity
Sexual
dimorphism
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
Fertilized
eggs
present
Spawning
females
present
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.

NA
NA
NF
Canada10
NA
NA
California1,2,6,10,12,17,18,25,30,31,33
*****
*****
*****
*****

NG
NG
s.
California5,7,9,21
*****
*****

NG
NG
NG
NG
NG
NG
NG
NG
NA
NA
NF
British
Columbia1
No
intrareader
comparison
California2,31,32,33
*****
*****
*****

NG
NG
n.
California7,16
Petersen
length­
frequency
NG
c.
California7,15,20
*****
*****

s.
California3,4,7,15,19,20,24,26,27
*****
*****
*****

NA
NA
NF
California2,8,13,28,31,33
*****
*****

NG
NG
NA
NA
NF
California2,14,29,31
NG
NG
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
recor
REPRODUCTION
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
AZ
BA
BB
BC
BD
BE
BF
Both
reproductive
states
present
Spawning
Frequency
Size
specific
fecundity
equation
Oct.
Nov.
Dec.
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)

Multiple9,10
NF
NF
NF
every
3
weeks
for
4
to
5
months10
Multiple1,2,4,5,11
NF
NF
2,4,6,11
18
to
24
times/
season
once
every
five
days4,6,11
NF
NF
NF
NF
NF
NF
NF
NF
Fecundity
range
in
#
eggs
(@
cm,
grams,

or
age)

rds
Mean
(
range)
fecundity
in
eggs/
g
female
wt.
1,3,4,6
800
(
15.5
TL)
to
37,200
(
26
TL)

Fecundity
larger
females:
7
months;
smaller
females:
3
to
4
months6
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BG
BH
BI
BJ
BK
BL
BM
BN
Size
at
settlement
Size
Fish
Size
Egg
duration
Larval
duration
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
(
mm
SL)

1
3
days
1
35
days
1,2
~
5
NF
1,2,51.24
to
1.32
planktonic1,2,3,6.10,12
NF
1,42.8
to
69.7
2
6
to
7
22.8
to
33
3
7
to
10
5
2.6
to
101.29
to
1.33
7
3.14
to
11
1.28
±
0.013
(
1.2
to
1.4)
8
2.49
(
2.44
to
2.76)

1,3
52
hours
2
3
months
NF
NF
10.79
to
0.92
planktonic2,3,4,5,9,13
NF
11.5
to
1.8
to
17.2
2,4,7
0.85
(
0.5
to
0.9)
21.8
to
2.8
to
13
8
0.81
(
SD=
0.02)
3
1.5
to
2.0
to
9
0.9
6
1.57
to
<
19.2
12
0.5
to
0.9
92.2
to
2.8
to
13
(
TL)

NF
NF
NF
NF
10.80
to
0.82
planktonic1,2,7,12
NF
11.4
to
11.5
NF
NF
NF
NF
10.66
to
0.76
planktonic1,2,12
NF
1<
1.9
to
14.0
Eggs
DISPERSAL
&
RECRUITMENT
2
Combined
egg
&
larval
duration
is
40
days
8at
depths
of
25
to
120
among
the
largest
of
scianid
eggs5
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10
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BO
BP
BQ
BR
BS
BT
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
(
SL
mm)

planktonic1,3,23
carnivore2
seeker1,2
NF
1
33.0
to
maturity
3
69.7
to
maturity
other
larval
white
seabass3
planktonic1,5,6,8
carnivore
NF
1
13
to
maturity
2
17.2
to
maturity
nocturnal2
4<
19.2
to
maturity
sandy
bottoms3
diurnal4
within
4
to
5
km
of
shore15,17
channels
and
bays22
planktonic1
carnivore
NF
NF
NF
NF
(
bivalve
veligers)
6
planktonic1
NF
NF
NF
NF
NF
Food
habits
Larvae
cyclopoid
copepods
and
euphausids2
(
planktonic
crustaceans,
such
as
shrimp
nauplii,
amphipods,
zoea;
bivlave
veligers;

trochophores;
tintinnids;
dinoflagellates;

larval
white
seabass)
2,6,7,8,9
possibly
northern
anchovy,

sardine.
mackerel,
and
juvenile
white
seabass1
larval
feeding
success
increases
with
size,

attributed
to
learning7,8
mostly
diurnal,
with
reduced
nocturnal
feeding1
shallow
inshore
waters
and
bays9,14
inshore,
lower
water
column
and
epibentic2,3,7,10,11,13,15,17,18,20,21
(
tintinnids;
bivalve
and
gastropod
veligers;

small
copepods,
especially
Euterpina
acutifrons
;
rotifers;
dinoflagellates;

polychaeete
larvae;
invertebrate
eggs)
1,2,4,5,6
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BU
BV
BW
BX
BY
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
carnivore1
engulpher1
NF
lizardfish;
halibut1
(
mysids,
copepods,
and
gammerids)
4
carnivore/
zooplanktivore2
NF
Caspian
terns3
diurnal2
shallow
sandy
bottoms
nearshore1,12
carnivore
NF
all
day6
NF
surf
zone12,14
carnivore
NF
NF
NF
(
clams;
polychaetes;
small
crustaceans)
10
kelp
forest;
bays
and
shallow
nearshore
waters,
near
surf­
zone;

harbors;
under
drift
algae
and
debris;

open
waters;
bays2,3,5,6,9,10,11,12,13,14
Juveniles
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
(
crustaceans,
such
as
caldocerans,

amphipods,
mysids,
and
euphasiids;

polychaetes;
chaetognath;
fish
larvaeand
small
fish)
1,7,9,11
shallow
sandy
habitat
with
vegetation8
epibenthic;
sandy
bottoms;
kelp
beds2,4,7,14
nocturnal,
but
some
midwater
feeding
diurnally1,5
(
small
fish,
such
as
anchovy,
sardine,

queenfish,
white
croaker,
white
sea
bass
larvae
and
mackerel;
squid)
1,4
(
veneroid
pelecypods
and
clam
siphons;

small
crustaceans,
such
as
amphipods
and
mysids)
3,11,12
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
BZ
CA
CB
CC
CD
Adults
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
carnivore3
NF
nocturnal1
marine
mammals;
sharks1,2,14
carnivore/
omnivore13,21
searcher1
nocturnal1,3,9
epibentic
searcher2
Western
gull6
Caspian
terns11
harbor
seals12,13
benthic
carnivore32
NF
all
day10
carnivore
NF
NF
NF
(
clams;
polychaetes;
crabs)
1,2,25,28,30,31
Pacific
bottlenose
dolphin;

comorants;
angel
shark2
deeper
waters
during
some
months
of
the
year2
kelp
forest;
sandy
bottoms;
artificial
reefs1,2,3,5,7,11,12,18,19,22,24,25
shallow
sandy
or
sandy
mud
bottoms;
surf
zone;
bays1,2,4,5,7,8,13,14,17,19,20,22,24,25,26
sandy
bottoms
and
sloughs;
near
breakwaters
and
bay
entrances;
in
holes
or
depressions;
artificial
reefs
some
midwater
feeding3
sea
lions;
Pacific
bottlenose
dolphin;
California
halibut;
black
sea
bass;
bluefin
tuna1,4,5,7,10
neritic;
sandy
nearshore
bottom
and
midwaters;
kelp
beds;
artificial
reefs2,3,5,6,10,11,12,13,14,15,19,22,23,25
spiny
dogfish;
harbor
seals
and
sea
lions;
California
halibut
and
barred
sandbass;
Brandt's
and
double­
crested
cormorant,

western
gulls,
and
least
terns15
(
fish,
such
as
anchovies,
herring,
and
smelts;
squid;

crayfish;
pelagic
red
crab;
isopods)

1,2,3,4,5,12,23,26,28,29,30,31
(
crustaceans,
such
as
gammarids,
caprellids,
and
decapods;
tubiculous
polychaetes;
calms,

scaphopods,
squid
and
octopus;
small
fish,
such
as
anchovy;
chaetognaths;
algae)
2,3,8,11,13,14,18,20,21,28,31
(
veneroid
pelecypods,
such
as
bean
clams;

crustaceans,
such
as
Emerita,
ghost
shrimp,
and
decapods;
polychaetes;
grunion
eggs)

2,4,9,22,24,28,30,31,32
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CE
CF
CG
CH
CI
CJ
CK
CL
CM
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
NF
NF
NF
NF
2NG
0.033
to
0.064
allozymes
2NG
0.024
to
0.06
allozymes
NF
NF
NF
NF
1
NG
0.03
(
NG)
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
none
in
California2
NF
NF
NF
NF
NF
NF
NF
NF
possible
substock
in
northern
Gulf
of
California,
separate
from
the
northeast
Pacific
population1,2
no
stock
structure
found
within
the
Southern
California
Bight5
need
74
to
satisfy
hatchery
conservation
goals1
Mean
heterozygosity
maybe
isolated
substocks
from
California
to
Mexico3
none
between
central
and
southern
California4
GENETICS
Nucleotide
diversity
(#)
Page
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15
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18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CN
CO
CP
CQ
CR
CS
CT
CU
CV
CW
CX
RECRUITMENT
INTO
FISHERY
Mortality
Estimates
First
Recruitment
Full
Recruitment
Method
of
ge
range
use
Total
length
(
age)
Total
length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
alculate
mort
NF
NF
NF
NF
NF
NF
1
0.303
0.589
0.892
Silliman
(
1943)/
Ricker
(
1958)
5
to
15
1
0.303
0.518
0.821
Silliman
(
1943)/
Ricker
(
1958)
4
to
15
1
0.303
0.259
0.562
Silliman
(
1943)/
Ricker
(
1958)
3
to
15
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
CY
CZ
DA
DB
DC
DD
DE
Rate
of
Exploitation
Rate
of
Exploitation
Interactions
/
associations
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

NG
0.66
NF
Juveniles
residential2
No1
0.19
0.631
0.33
0.461
NF
NF
NF
occurs
with
S.
politus
1,2
move
offshore
in
winter5
Transient1,4
NF
parasitized
by
copepod
family
Naobranchia
6
NF
NF
NF
1
up
to
93
km
Residential1,3,5
NF
2
up
to
94.5
km
NF
NF
NF
NF
Transient1
NF
infested
by
roundworms
from
genera
Anisakis
8
infested
by
parasites
from
the
families
Caligidae,
Lernaeopodidae,
Dasyrhynchidae,

Acanthocolpidae,
Anisakidae,
Ceratomyxidae,

and
Tetracapsulidae15
share
same
resources
as
S.
politus
and
Phanerodon
furcatus
14
flesh
found
to
contain
high
levels
of
DDT,

PCBs
in
Los
Angeles
and
southern
California
areas7,10,11,12
pathological
evidence
of
stress
near
power
plant
outfalls14
infested
by
parasites
from
the
families
Rhadinorhynchidae,
Bomolochidae,
Caligidae,

Hatschekiidae,
Bucephalidae,
Diplangidae,

Monorchiidae,
and
Opecoelidae15
possibly
not
fit
for
human
consumption
in
areas
of
waste
discharge3
move
to
deeper
water
during
winter7,8
Movement
juveniles
infested
by
the
gliding
bacteria
Flexibacter
maritimus
13
infested
by
parasites
from
the
families
Lacistorhynchidae,
Bomolochidae,
Caligidae,

Lernaeopodidae,
Acanthocolpidae,
and
Anisakidae15
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
8
increasing
risk
of
hepatic
lesions
in
older
fish,

fish
exposed
to
chemical
contaminants9
infested
by
parsites
from
the
families
Rhadinorhynchidae,
Tetraphyllidea,

Bomolochidae,
Lernaeopodidae,
Diplangidae,
BEHAVIOR
move
to
deeper
waters
in
winter6
reproductive
success
decreased
by
contaminants
introduced
by
humans4,5
Page
862
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1167
5­
2220e
­
Sciaenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
DF
DG
Schooling
Territoriality
Yes1,3,4,6,15,17
NF
Yes2,3,4,5,7,14,18,19
NF
Yes1,6,17
NF
Yes1,6,16,17
NF
Both
schooling
and
solitary3,8,19
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863
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1167
5­
2220e
­
Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
A
B
C
D
E
Seriphus
politus
1,2
Umbrina
roncador
1,2
1surf
to
10
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Love
et
al.
2000
1­
Love
et
al.
2000
2­
Eschmeyer
1998
2­
Gates
&
Frey
1974
3­
Love
et
al.
2000
3­
Skogsberg
1939
4­
Roedel
1948
5­
Starks
1919
yellowfin
croaker;
surf
fish;
yellowfinned
roncador;
Catalina
croaker;
yellowtailed
croaker;
golden
croaker
1,2,4
British
Columbia
(
Burrard
Inlet,
Vancouver
I.)

to
the
southern
Gulf
of
California1
1inshore
to
181
California
(
Point
Conception)
to
southern
Baja
California
(
Bahia
Magdalena)
1
queenfish;
kingfish;
tomcod;
sea
trout;
shiner;

herring;
white
croaker;
herring
croaker
1,2,4,5
Page
864
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1167
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Sciaenidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
F
G
H
I
JK
L
1SL=
­
14.96FL0.902
(
SE=
1.832)

1FL=
­
4.239TL0.977
(
SE=
2.707)

1FL=
5.868SL1.137
(
SE=
2.084)

2SL=
0.833TL­
4.382
2Both:
W=
0.00000879SL3.111
NF
1
53.5
NF
NF
NF
NF
NF
NF
1
55.6
NF
NF
NF
NF
1­
Thomas
1968
1­
Karpov
&
Kwiecien
1988
1­
Love
et
al.
2000
1­
Thomas
1968
2­
DeMartini
&
Fountain
1981
2­
Joseph
1962
2­
Love
et
al.
1984
3­
Clark
1930b
3­
Love
et
al.
1984
4­
Love
et
al.
1984
4­
Isaacson
1964
5­
Isaacson
1964
6­
Kim
1987
7­
Dutton
1989
Page
865
of
1167
5­
2220e
­
Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
M
N
O
P
Q
R
S
T
U
V
W
X
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
866
of
1167
5­
2220e
­
Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
Y
Z
AA
AB
AC
AD
AE
NF
NF
NF
NF
NF
oviparous2
NF
NF
10
>
10
NF
NF
NF
oviparous
NF
1­
Joseph
1962
1­
Emmett
et
al.
1991
1­
Fleming
1999b
1­
Emmett
et
al.
1991
1­
Emmett
et
al.
1991
2­
Donohoe
1997
2­
Fitch
&
Lavenberg
1971
2­
Love
1996
3­
Issacson
1967
3­
Oliphant
1992b
4­
Kim
1987
4­
Wild
1992
5­
Dutton
1989
5­
Fleming
1999b
6­
Orhun
1989
6­
Love
et
al.
1984
7­
Tenera
Environmental
Services
2000
7­
Carlisle
1960c
8­
Maxwell
1975
9­
Tenera
Environmental
Services
2000
10­
Love
1996
Page
867
of
1167
5­
2220e
­
Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
AF
AG
AH
AI
AJ
AK
AL
NF
NF
217
(
NG)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NA
6NG
(
NG)
10
to
10.5
(
1)
NG
(
NG)
11
(
2)
otoliths;
section
17
NG
(
NG)
NG
(
NG)
12.7
(
1)­
sex
unspecified
NF
NF
2,16,17
23
(
NG)­
unspecified
sex
NG
(
NG)
NG
(
NG)
NA
1­
Wild
1992
1­
Oliphant
1992b
1­
Hart
1973
2­
Love
et
al.
1984
2­
Love
et
al.
1984
2­
Skogsberg
1939
3­
Love
1996
3­
Emmett
et
al.
1991
4­
Fitch
&
Lavenberg
1971
5­
Thomas
1968
6­
DeMartini
&
Fountain
1981
7­
Vojkovich
1992
8­
Oliphant
1992b
9­
Wild
1992
10­
Clark
1930b
11­
Love
et
al.
1984
12­
Issacson
1967
13­
Carlisle
1960b
14­
Carlisle
1960c
15­
Pinkas
1960b
16­
Maxwell
1975
17­
Love
1996
Page
868
of
1167
5­
2220e
­
Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
NA
NA
NF
California2,10,22,33
*****
*****
*****

NG
NG
s.
California3,4,5,11,19,20,23,27,31
*****
*****
monthly
peaks
during
moon's
first
quarter11
NA
NA
NF
California5,31
1­
Hart
1973
2­
Moser
1996b
3­
Lavenberg
et
al.
1986
4­
Goldberg
1987
5­
Skogsberg
1939
6­
Emmett
et
al.
1991
7­
Monaco
et
al.
1990
8­
Fitch
&
Lavenebrg
1971
9­
Thomas
1968
10­
Breder
&
Rosen
1966
11­
DeMartini
&
Fountain
1981
12­
Vojkovich
1992
13­
Oliphant
1992a
14­
Oliphant
1992b
15­
Wild
1992
16­
Fleming
1999b
17­
Vojkovich
&
Reed
1983
18­
Bartley
et
al.
1995
19­
Schlotterbeck
&
Connally
1982
20­
Love
et
al.
1984
21­
Donohoe
1997
22­
Demartini
et
al.
1985
23­
DeMartini
1991a
24­
Watson
1982
25­
Kramer
&
Smith
1973
26­
Issacson
1967
27­
Lavenberg
et
al.
1986
28­
Carlisle
1960b
29­
Carlisle
1960c
30­
Pinkas
1960b
31­
Maxwell
1975
32­
Wang
1981
33­
Love
1996
Page
869
of
1167
5­
2220e
­
Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
AZ
BA
BB
BC
BD
BE
BF
Multiple1,7
1
287
(
218
to
494)
1F=
0.0007153SL3.3809
(
r2=
0.664)

1F=
92.897(
W
in
grams)
1.302
(
r2=
0.816)

2F=
192.5(
W
in
grams)
1.125
(
r=
0.729)

2F=
152.9(
W
in
grams)
1.19
(
r=
0.712)

2F=
107.8(
W
in
grams)
1.212
(
r=
0.444)

2F=
119.1(
W
in
grams)
1.249
(
r=
0.666)

7
300,000
(
NG)
2F=
62.8(
W
in
grams)
1.41
(
r=
0.542)

NF
NF
NF
NF
1­
Goldberg
1987
1­
DeMartini
&
Fountain
1981
1­
Emmett
et
al.
1991
1­
DeMartini
&
Fountain
1981
2­
Emmett
et
al.
1991
2­
DeMartini
&
Fountian
1981
2­
DeMartini
1991a
3­
DeMartini
&
Fountain
1981
3­
Wild
1992
4­
Wild
1992
4­
Love
et
al.
1984
5­
Fleming
1999b
5­
DeMartini
1991a
6­
Love
et
al.
1984
6­
Tenera
Environmental
Services
2000
7­
DeMartini
et
al.
1985
7­
Love
1996
8­
DeMartini
1991a
9­
Kent
et
al.
1993
10­
Orhun
1989
11­
Tenera
Environmental
Services
2000
spawn
every
7.4
days;
spawning
duration:

small
individuals­
3
months,
large
individual­

up
to
6
months3
25,000
(
10.5
SL)
to
90,000
(
25
SL)
per
batch
20
batches
per
season
over
6
months
8
511,784
(
NG)
to
15,019
(
NG)
per
batch
260,000
(
10.5
SL)
to
2,200,000
(
25
SL)
per
season
Page
870
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Sciaenidae
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65
66
67
68
69
70
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73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BG
BH
BI
BJ
BK
BL
BM
BN
NF
2
3
months
NF
NF
10.73
to
0.78
planktonic1,2,4,11,12
NF
11.6
to
17.2
NF
NF
NF
NF
10.74
to
0.80
planktonic2
NF
11.5
to
20.7
1­
Emmett
et
al
.1991
1­
Cordes
1992
1­
Allen
&
Franklin
1992
1­
Moser
1996b
1­
Limbaugh
1955
1­
Moser
1996b
2­
Bartley
et
al.
1995
2­
Beckwitt
1983
2­
Donohoe
1997
2­
Emmett
et
al.
1991
2­
Moser
1996b
2­
Emmet
et
al.
1991
3­
Watson
1982
3­
Allen
&
Franklin
1988
3­
DeMartini
&
Fountain
1981
3­
Emmet
et
al.
1991
3­
Matarese
et
al.
1989
4­
Matarese
et
al.
1989
4­
Allen
1982
4­
Moser
et
al.
1983
5­
Moser
et
al.
1983
5­
Matarese
et
al.
1989
5­
Margulies
1989
6­
DeMartini
1991a
6­
Vojkovich
1992
6­
Watson
1982
7­
Watson
1982
7­
Oliphant
1992a
7­
Kim
1987
8­
Gadomski
&
Caddell
1996
8­
Wild
1992
8­
Orhun
1989
9­
Morris
1956a
9
­
Fleming
1999b
9­
Wang
1981
10­
Kim
1987
10­
Vojkovich
&
Reed
1983
11­
Orhun
1989
11­
DeMartini
1991a
12­
Wang
1981
12­
Feder
et
al.
1974
13­
Wang
1981
60.478
to
0.523;
eggs
larger
in
larger
females
30.68;
eggs
larger
in
larger
females
Page
871
of
1167
5­
2220e
­
Sciaenidae
51
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60
61
62
63
64
65
66
67
68
69
70
71
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73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BO
BP
BQ
BR
BS
BT
planktonic1,5,16
carnivore
NF
NF
NF
diurnal3
within
2
km
from
shore15
vertically
dispersed
at
night19
planktonic1
NF
NF
NF
NF
NF
1­
Moser
1996b
1­
Watson
&
Davis
Jr.
1989
1­
Dutton
1989
1­
Watson
&
Davis
Jr.
1989
1­
Margulies
1989
1­
Emmet
et
al.
1991
2­
Watson
&
Davis
Jr.
1989
2­
Emmett
et
al.
1991
2­
Orhun
1989
2­
Emmett
et
al.
1991
2­
Brewer
et
al.
1984
2­
Moser
1996b
3­
Emmet
et
al.
1991
3­
DeMartinin
&
Fountian
1981
3­
Jahn
&
Lavenberg
1986
3­
Orhun
1989
3­
Moser
et
al
1983
4­
DeMartini
&
Fountain
1981
4­
Schlotterbeck
&
Connally
1982
4­
Brewer
&
Kleppel
1986
4­
Watson
1982
5­
Allen
1982
5­
Jahn
et
al.
1988
6­
Matarese
et
al.
1989
6­
Brewer
&
Kleppel
1986
7­
Wild
1992
7­
Dutton
1992
8­
Fleming
1999b
8­
Dutton
1989
9­
Eldridge
1977
9­
Orhun
1989
10­
Schlotterbeck
&
Connally
1982
11­
Love
et
al.
1984
12­
Jahn
et
al.
1988
13­
Moser
&
Pommeranz
1999
14­
Donohoe
1997
15­
Barnett
et
al.
1984
16­
DeMartini
1991a
17­
Watson
1982
18­
Brewer
et
al.
1984
19­
Jahn
&
Lavenberg
1986
20­
Brewer
&
Kleppel
1986
21­
Beckwitt
1983
22­
Leithiser
1981
23­
Orhun
1989
24­
Tenera
Environmental
Services
2000
inshore,
lower
water
column
and
epibentic2,10,13,15,19
mostly
diurnal,
with
reduced
nocturnal
feeding1
(
bivalve
veligers;
small
copepods,
Euterpina
acutifrons;
mysids)
1,3,4,6
<
30
m
and
within
4
to
5
km
of
shore4
Page
872
of
1167
5­
2220e
­
Sciaenidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BU
BV
BW
BX
BY
carnivore
NF
nocturnal3,4
NF
NF
NF
NF
NF
1­
Limbaugh
1955
1­
Emmett
et
al.
1991
1­
Margulies
1989
1­
Emmett
et
al.
1991
1­
Donohoe
1997
2­
Emmett
et
al.
1991
2­
Allen
1982
2­
Allen
1982
2­
DeMartini
et
al.
1985
3­
Thomas
1968
3­
Oliphant
1992a
3­
DeMartini
et
al.
1985
3­
Parkin
1998
4­
Love,
Stephens,
et
al.
1996
4­
Donohoe
1997
4­
Hobson
&
Chess
1976
5­
CDFG
1982
5­
DeMartini
et
al.
1985
5­
Love
1996
6­
Vojkovich
1992
6­
DeMartini
1991a
6­
O'Brien
&
Valle
2000
(
in
press)

7­
Wild
1992
7­
Brewer
et
al.
1984
8­
Allen
&
Franklin
1992
8­
Hobson
&
Chess
1976
9­
Vojkovich
&
Reed
1983
9
­
Ware
1979
10­
Donohoe
1997
10­
Maxwell
1975
11­
Allen
&
Franklin
1988
11­
Love
1996
12­
Feder
et
al.
1974
12­
O'Brien
&
Valle
2000
(
in
press)

13­
Maxwell
1977
14­
Love
1996
(
holoplanktonic
copepods,
mysids,
and
other
small
crustaceans;
small
fish
and
fish
larvae,
such
as
anchovy)
5,6,7,8
shallow
sandy
bottoms,
moving
deeper
during
winter4,12
California
scorpionfish;

Pacific
mackerel;
Pacific
bonito;
halibut;
kelp
bass;

barracuda2
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873
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­
Sciaenidae
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65
66
67
68
69
70
71
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73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
BZ
CA
CB
CC
CD
carnivore/
planktivore24
pursuer1
nocturnal3,4,5,6,7,8
halibut3,5
Brandt's
cormorants6
kelp
bass8
carnivore/
benthivore7
NF
nocturnal2,5,10
NF
1­
Limbaugh
1955
1­
Limbaugh
1955
1­
Allen
1982
1­
Emmet
et
al.
1991
1­
Emmet
et
al.
1991
2­
Skogsberg
1939
2­
Skogsberg
1939
2­
Ware
1979
2­
Hobson
&
Chess
1986
2­
Fitch
&
Lavenberg
1971
3­
Emmet
et
al.
1991
3­
Emmett
et
al.
1991
3­
Allen
1982
3­
Limbaugh
1955
4­
Fitch
&
Lavenberg
1971
4­
Fitch
&
Lavenberg
1971
4­
DeMartini
et
al.
1985
4­
Wild
1992
5­
Love,
Stephens
et
al.
1996
5­
Thomas
1968
5­
Hobson
et
al.
1981
5­
Wertz
&
Domeier
1997
6­
Allen
1982
6­
Stephens
et
al.
1984
6­
Helvey
1985
6­
Hubbs
et
al.
1970
7­
CDF&
G
1982
7­
Hobson
&
Chess
1986
7­
Hobson
&
Chess
1976
7­
Morejohn
et
al.
1978
8­
Oliphant
1992a
8­
Allen
1982
8­
Hobson
1994
8­
Young
1963
9­
Oliphant
1992b
9­
CDF&
G
1982
9­
Love
1996
9­
DeMartini
et
al.
1985
10­
Wild
1992
10­
Oliphant
1992a
10­
O'Brien
&
Valle
2000
(
in
press)
10­
Ware
1979
11­
Carlisle
et
al.
1964
11­
Wild
1992
11­
Parkin
1998
12­
Turner
et
al.
1969
12­
Vojkovich
&
Reed
1983
12­
Torok
1994
13­
Love
et
al.
1984
13­
Love
et
al.
1984
13­
Trumble
1995
14­
Allen
1985
14­
Morejohn
et
al.
1978
14­
Pinkas
1960b
15­
Beckwitt
1983
15­
DeMartini
et
al.
1985
15­
Love
1996
16­
Hobson
&
Chess
1976
16­
DeMartini
1991a
17­
Klingbeil
1972
17­
Hobson
et
al.
1981
18­
Orhun
1989
18­
Helvey
1985
19­
Love
et
al.
1986
19­
Hobson
&
Chess
1976
20­
Carlisle
1960b
20­
McCain
et
al.
1992
21­
Carlisle
1960c
21­
Ware
1979
22­
Maxwell
1975
22­
Klingbeil
1972
23­
Quast
1968a
23­
Orhun
1989
24­
Feder
et
al.
1974
24­
Carlisle
1960b
25­
Love
1996
25­
Carlisle
1960c
26­
O'Brein
&
Valle
2000
(
in
press)
26­
Pinkas
1960b
27­
Hobson
1994
28­
Maxewell
1975
29­
Quast
1968d
30­
Feder
et
al.
1974
31­
Love
1996
32­
O'Brein
&
Valle
2000
(
in
press)

1,2,5,7,9,11,13,14,19,21,24,25
sandy
bottoms;
bays
and
sloughs;
surf;

artificial
reefs1,2,5,7,9,11,13,14,19,22,24,25,26
fine
sand
bottoms;
midwater;
artificial
reefs;

kelp
forests
at
night
1,2,5,11,14,15,16,19,22,24,25
(
small
crustaceans,
such
as
mysids,
gammerids,

and
isopods;
polychaetes;
small
fish,
like
anchovy)

1,2,8,15,16,17,19,28,30,31
Pacific
mackerel;
Pacific
bonito;

halibut;
kelp
bass9
(
small
fish
and
fish
eggs;
small
crustaceans,
such
as
gammerids;
mollusks;
tubiculous
polychaetes)

2,6,17,28,31,32
sea
lions;
least
terns;

doublecrested
cormorants15
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874
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Sciaenidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
CE
CF
CG
CH
CI
CJ
CK
CL
CM
none
within
southern
California4
NF
NF
NF
NF
1NG
0.043
(
NG)
NF
NF
none
in
California2
NF
NF
NF
NF
NF
NF
NF
NF
1­
Thomas
1968
1­
Beckwitt
1983
1­
Bartley
et
al.
1995
2­
CDFG
1982
2­
Bartley
&
Kent
1990
3­
Vojkovich
1992
4­
Beckwitt
1983
5­
Bartley
&
Kent
1990
Page
875
of
1167
5­
2220e
­
Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
CN
CO
CP
CQ
CR
CS
CT
CU
CV
CW
CX
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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876
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1167
5­
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Sciaenidae
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
CY
CZ
DA
DB
DC
DD
DE
NF
NF
NF
associated
with
G.
lineatus
1
Transient3,4
NF
move
offshore
in
winter5
NF
NF
NF
NF
Transient1
NF
1­
Skogsberg
1939
1­
CDFG
1982
1­
CDFG
1982
1­
Bartley
et
al.
1995
2­
Emmett
et
al
.1991
2­
Oliphant
1992a
2­
Drawbridge
et
al.
1995
3­
Puffer
et
al.
1982
3­
DeMartini
et
al.
1985
3­
Stephens
et
al.
1994
4­
Cross
&
Hose
1989
4­
DeMartini
&
Fountain
1981
4­
Helvey
1985
5­
Cross
&
Hose
1988
5­
Helvey
1985
5­
Klingbeil
1972
6­
Dojiri
1981
6­
Klingbeil
1972
7­
Castle
&
Woods
1972
7­
Love
1996
8­
Dailey
et
al.
1981
8­
Skogsberg
1939
9­
Myers
et
al.
1994
10­
Gossett
et
al.
1983
11­
Gossett
et
al.
1989
12­
McCain
et
al.
1992
13­
Chen
et
al.
1995
14­
Ware
1979
15­
Love
&
Moser
1983
share
same
resources
as
G.
lineatus
and
Phanerodon
furcatus
14
Monorchiidae,
and
Opecoelidae15
nighttime
offshore
movments
to
spawn3,4
offshore
movements
at
night
to
feed3
infested
by
roundworms
from
genera
Anisakis
8,15
infested
by
parasites
from
the
families
Polymorphidae,
Rhadinorhynchidae,

Acanthocolpidae,
Diplangidae,
Opecoelidae,

Cymothoidae,
and
Diplectanidae15
Page
877
of
1167
5­
2220e
­
Sciaenidae
51
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
DF
DG
Yes1,5,9,10,11,12,13,17
NF
school
during
the
day,
disperse
at
night
to
feed
11
school
during
the
day,
disperse
at
night
to
feed
11
NF
1­
Limbaugh
1955
2­
Hart
1973
3­
Skogsberg
1939
4­
Emmet
et
al.
1991
5­
Allen
1982
6­
CDF&
G
1982
7­
Wild
1992
8­
Vojkovich
and
Reed
1983
9­
Carlisle
et
al.
1964
10­
DeMartini
et
al.
1985
11­
Hobson
et
al.
1981
12­
Helvey
1985
13­
Hobson
&
Chess
1976
14­
McCain
et
al.
1992
15­
Orhun
1989
16­
Carlisle
1960c
17­
Feder
et
al.
1874
18­
Tenera
Environmental
Services
2000
19­
Love
1996
Page
878
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1167
5­
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­
Sciaenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Scorpaenidae1
Scorpaena
guttata
1
1
1
to
183
Sebastes
atrovirens
1
1<
1
to
46
S.
auriculatus
1
1
<
1
to
135
S.
carnatus
1
1
<
1
to
80
California
scorpionfish;
spotted
scorpionfish;
sculpin;
scorpion;
bullhead;

scorpene1,2,4,9,13
northern
Gulf
of
alaska
(
Prince
William
Sound)

to
central
Baja
California
(
Bahia
San
Hipolito)
1
central
CA
(
Santa
Cruz)
to
the
Gulf
of
California1,2
northern
CA
(
Timber
Cove)
to
central
Baja
California
(
Punta
San
Pablo)
1
kelp
rockfish;
sugar
bass;
dumb
bass;

garrupa;
grass
bass;
green
garrupa1,2,4,6,9,11,12
brown
rockfish;
bolina;
garrupa;

chocolate
bass;
P.
D.
bass;
boraccho;

cinnamon
bass;
brown
bass;
ground
owl;

chocolate
spot1,2,3,4,5,6,12
gopher
rockfish;
gopher
cod;
rock
bass;

spotted
rock
bass;
flesh­
colored
rockfish;

butterball;
butter
bass1,2,3,6,7,9,12
northern
CA
(
Eureka)
to
central
Baja
California
(
San
Roque)
1
TAXONOMY
RANGE
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1234
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67
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
F
G
H
I
J
Length­
weight
equation
Max.
reported
Geographic
Area
Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
9Male:
W=
0.0205TL3.0045
(
r2=
0.883)
NF
1
43
s.
California
(
TL)
12
36.3
(
1.6)

9Female:
W=
0.0196TL3.0102
(
r2=
0.865)

21Both:
W=
0.0000398SL2.98
8*
Male:
W=
0.00001327TL3.038
(
r2=
0.927)
1TL/
SL=
1.233
1
42.5
c.
California
(
TL)
11
37.7
8*
Female:
W=
0.000009984TL3.094
(
r2=
0.964)
1SL/
TL=
0.811
c.
California
(
SL)
24
28.2
8*
Both:
W=
0.000006291TL3.172
(
r2=
0.990)

21Both:
W=
0.0000427SL2.92
2Both:
W=
0.044TL2.74
(
r2=
0.94)
4TL=
­
1.423
+
1.24(
SL);
r2=
1.0;
SE=
3.8
1
56
Washington
(
TL)
19
35
14*
Male:
W=
0.00000376TL2.87
(
r2=
0.914)
4TL=
­
0.586
+
1.007(
FL);
r2=
1.0;
SE=
1.6
Washington
(
TL)
37
35
14*
Female:
W=
0.00000261TL2.95
(
r2=
0.940)
4SL=
1.369
+
0.806(
TL);
r2=
1.0;
SE=
3.1
California
(
TL)
6
NG
14*
Both:
W=
0.00000172TL3.02
(
r2=
0.925)
4SL=
0.575
+
0.813(
FL);
r2=
0.998;
SE=
3.4
28Male:
W=
0.0000000376TL2.87
(
r2=
0.914)
4FL=
0.634
+
0.993(
TL);
r2=
1.0;
SE=
1.6
28Female:
W=
0.0000000261TL2.95
(
r2=
0.94)
4FL=
­
0.369
+
1.228(
SL);
r2=
0.998;
SE=
4.1
28Both:
W=
0.0000000172TL3.02
(
r2=
0.925)

8*
Male:
W=
0.00001595TL3.027
(
r\=
0.949)
4TL=
3.676
+
1.201(
SL);
r2=
0.998;
SE=
2.2
1
34.8
c.
California
(
TL)
11
32.9
8*
Female:
W=
0.00001921TL3.01
(
r2=
0.960)
4TL=
­
0.759
+
1.005(
FL);
r2=
1.0;
SE=
0.5
8*
Both:
W=
0.00001299TL3.077
(
r2=
0.986)
4SL=
­
2.866
+
0.832(
TL);
r2=
0.998;
SE=
1.8
4SL=
­
3.613
+
0.836(
FL);
r2=
0.998;
SE=
2.0
4FL=
0.768
+
0.995(
TL);
r2=
1.0;
SE=
0.5
4FL=
4.601
+
1.194(
SL);
r2=
0.998;
SE=
2.4
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)
equation.
*

denotes
TL
=
mm
or
FL,
SL=
cm
Page
880
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1167
5­
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­
Scorpaenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
K
L
M
N
O
P
Q
R
S
T
U
Growth
Parameters
Linf
in
cm
(
SE)
k
(
SE)
to
(
SE)

Female
Both
Male
Female
Both
Male
Female
Both
Male
Female
Both
44.3
(
1.57)
NG
0.12
(
0.02)
0.13
(
0.02)
NG
­
3.86
(
0.68)
­
1.9
(
0.42)
NG
NG
NG
NG
36.9
37.8
0.2522
0.2552
0.2307
­
0.3
­
0.4
­
0.7
NG
NG
NG
28.5
NG
0.3
0.29
NG
­
0.01
­
0.03
NG
NG
NG
NG
47
NG
0.21
0.087
NG
­
1.5
­
4.5
NG
NG
NG
NG
47
53
0.21
0.087
0.06
­
1.5
­
4.5
­
6.2
NG
NG
NG
NG
51.4
NG
NG
0.16
NG
NG
­
0.55
NG
NG
NG
34.1
34.1
0.2753
0.2531
0.2256
0.1
0.1
­
0.5
NG
NG
NG
r2
AGE
&
GROWTH
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1167
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2220e
­
Scorpaenidae
1234
5
67
8
9
10
11
12
13
14
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20
21
22
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24
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
V
W
X
Y
Ageing
structure
/
Method
Validation
Verification
Other
refs.

pterygiophores;
section
Edge
Analysis
Intrareader
comparisons
NF
otoliths;
whole
Edge
Analysis
Interreader
comparisons
NF
otoliths;
whole
Edge
Analysis
Intra/
interreader
comparisons
otoliths;
whole
No
No
16­
larvae
growth
otoliths;
section
NG
NG
otoliths;
section
No
Intrareader
comparisons
otoliths;
whole
Edge
Analysis
Interreader
comparisons;
comparisons
with
operculum
NF
Page
882
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2220e
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Scorpaenidae
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16
17
18
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
Z
AA
AB
AC
AD
AE
AF
Age
composition
Sexual
dim
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male:
Female
SL
ratio
1,35at
least
21
NF
Dioecious
Polygynous1
Oviparous1
No
2
Male<
Female
(
ovuliparity)
3
4,21,33
20
NF
Dioecious
Polygynous2
Viviparous3,7,9
No
NF
17,36>
20
2,18,21
Dioecious
Polygynous2
Viviparous3,7
No
1
0.94
21,31
19
Ovoviviparous10
17>
30
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
1.02
21,33
24
Ovoviviparous12
Breeding
Behavior
Page
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Scorpaenidae
1234
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AG
AH
AI
AJ
AK
AL
AM
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
morphism
1st
maturity
50%
maturity
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
NF
1,53
15
(<
1)
14
(<
1)
17
(
2)
18
(
2)
pterygiophores;
section
Edge
Analysis
5119
(
NG)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NG
NG
None1
2,5225.1
(
4)
N=
251
24.4
(
5)
N=
218
NG
(
NG)
NG
(
NG)
otoliths;
whole
Edge
Analysis
26NG^(
NG)
15.8
(
3)
NG
(
NG)
NG
(
NG)
otoliths;
whole
Edge
Analysis
3016.0^
(
2)­
unspecified
sex
NG
(
NG)
NG
(
NG)
otoliths;
NG
NG
32,52NG
(
NG)
NG
(
NG)
26
(
4
to
5)­
sex
unspecified
NG
NG
None1
3,46,53
26
(
3)
26
(
3)
31
(
5)
31
(
5)
otoliths;
whole
No
423
(
2)
423
(
3)
25
(
3.9)
26.4
(
4.2)
otoliths;
section
No
15,41NG
(
NG)
NG
(
3)
NG
(
NG)
NG
(
4)
otoliths;
whole
No
32NG
(
NG)
NG
(
NG)
31
(
5)­
sex
unspecified
NG
NG
3225
(
3)
25
(
3)
31
(
5)
31
(
5)
NG
NG
None1
3,5217
(
4)
17
(
4)
17
(
4)
17
(
4)
otoliths;
whole
No
2,52
23.7
(
10)
20.7
(
NG)
NG
(
NG)
NG
(
NG)
otoliths;
whole
Edge
Analysis
1013.5^
(
3)
13.5
(
3)
NG
(
NG)
NG
(
NG)
otoliths;
whole
NG
32NG
(
NG)
NG
(
NG)
17
(
4)­
sex
unspecified
NG
NG
5514
to
15^
(
3)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NG
NG
Page
884
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1167
5­
2220e
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Scorpaenidae
1234
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AN
AO
AP
AQ
AR
AS
AT
AU
AV
Fertilized
eggs
present
Spawning
f
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
intrareader
comparisons
NF
s.
California1
NG
California2,3,27,28,77
interreader
comparisons
NF
c.
California4,34,43,49,51
*****
^^^^^
*****

intra/
interreader
comparisons
s.
California55
NG
California57,75,76
*****
*****
*****
*****

NG
No
NF
Washington7,8,9,35,36,42,70
*****
********

intrareader
comparisons
Oregon77
No
n.­
c.
California5
^^^^^^
********

NG
c.
California4,77
NG
s.
California7
*****

California57,58
*****
*****
*****

No
NF
n.­
c.
California5
******

interreader
comparisons;
comparison
with
operculum
c.
California4,34,76,78
^^^^^
^^^^^
*********

NG
s.
California10
NG
California57
*****
*****

NG
REPRODUCTION
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn
Page
885
of
1167
5­
2220e
­
Scorpaenidae
1234
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13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
AW
AX
AY
AZ
BA
BB
BC
BD
BE
females
present
Both
reproductive
states
present
Spawning
Frequency
Number
of
eggs
July
Aug.
Sept.
Oct.
Nov.
Dec.
(
per
year)
at
cm
length
*****
NF
NF
NF
Once1,17,19
1,5374
(
344­
403)
15172,000
(
30.5
TL)

Once6
1139
(
95­
176)
6,9,11,20
51,619
(
31.1
TL)
to
338,540
(
47.7
TL)

Multiple14,15,21,23
21,27
42,500
(
30.5
?
L)
to
266,000
(
46
?
L)

^^^^^^^
^^^^^^

*****
Once1,17,22
1,5
249
(
176­
307)
NF
Mean
(
range)
fecundity
in
eggs/
g
female
wt.

;
#####
=
doubtful
spawning
records
Fecund
Page
886
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1167
5­
2220e
­
Scorpaenidae
1234
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BF
BG
BH
BI
BJ
Size
specific
fecundity
equation
Size
at
settlement
(
TL=
cm;
*
denotes
TL,
SL=
mm)
Egg
duration
Larval
duration
(
SL
in
mm)
Annual
cohort
success
NF
1,43
days
1
~
1
to
2
months
5
20.5
NF
2,7<
5
days
3
2­
3
days
*
9F=
0.000021(
SL)
4.134
(
r2=
0.788)
NA
1
~
1
to
2
months
121
to
48
NF
3
50
to
75
days
15,20
20
to
40
TL
*
2,14F=
0.000363(
TL)
3.34124
NA
1
~
1
to
2
months
1218
to
25
NF
1530
to
35
NF
NA
1
~
1
to
2
months
20
20
to
40
TL
NF
360
to
90
days
3
80
to
90
days;

91.5
days
DISPERSal
&
RECRUITMENT
dity
Page
887
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1167
5­
2220e
­
Scorpaenidae
1234
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BK
BL
BM
BN
BO
BP
Larvae
Size
Fish
Size
Food
habits
(
diameter
in
mm)
Habitat
Predators
(
mm)
Habitat
Foraging
guild
2,9ovoid:
1.16
to
1.119
&
1.22
to
1.29
planktonic1,3,4
NF
6,8,13,14,17,181.9
to
20.5
(
SL)
planktonic1
planktivore
3,4,15
1.2
5
0.42
to
0.76
No
external
eggs
NA
1,17,224
to
23
(
SL)
planktonic1,7
planktivore
74.3
(
SL)
to
14
1.3
to
1.5
No
external
eggs
NA
10,245
(
SL)
to
planktonic1,6
planktivore
17,18,20,235.5
(
SL)
to
20,31
4.7
(
SL)
to
215.8
(
TL)
to
5
0.86
No
external
eggs
NA
7,264.4
(
SL)
to
planktonic1
planktivore
17,224.3
to
25
(
SL)

215.5
(
TL)
to
254.6
(
SL)
to
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
eggs
contained
within
a
gelatinous
floating
mass2,3,4,9,15,18
Eggs
Page
888
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5­
2220e
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Scorpaenidae
1234
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13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BQ
BR
BS
BT
BU
BV
Fish
Size
Foraging
method
When
/
Frequency
Predators
(
mm)
Habitat
Foraging
guild
filter
feeder1
NF
NF
720.5
(
SL)
to
maturity
carnivore
(
crabs;
small
fishes;
octopi)
25
deeper
waters52
NF
diurnal1
siphonophores;
923
(
SL)
to
maturity
zooplanktivore
chaetognaths1
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
NF
diurnal1
siphonophores;
25
(
SL)
to
maturity9
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
calanoids,
shrimp,
and
barnacle
cyprids)
1,4,6,8,10
Life
Stage
Juveniles
shallow
water
in
dense
algae;
rocky
areas;

artificial
reefs21,25,26,54
F
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
pelagic;
kelp
canopy;
artificial
reefs
2,3,9,13,24,26,36,42,44,53,54
(
crustaceans,
such
as
gammarids
and
barnacle
cyprids;
juvenile
fishes)
1,6,7,8,10,18
(
crustaceans,
such
as
barnacle
cyprids)
1,6,20
pelagic;
low
or
high
relief
rock;
sand­
rock
interface;
crevices;
estuaries;
eelgrass;

low­
relief
artificial
reefs
pelagic;
kelp
canopy;
rocky
reefs;
sand
and
low
rock;
artificial
reefs
1,2,3,8,9,12,13,24,26,53
Page
889
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1167
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2220e
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Scorpaenidae
1234
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
BW
BX
BY
BZ
CA
Foraging
method
When
/
Frequency
Predators
Habitat
Foraging
guild
NF
NF
NF
(
meso)
carnivore8
foliage­
searcher3
diurnal4,5
carnivore
NF
NF
carnivore
eelgrass
beds
at
night47
NF
diurnal4,5
(
meso)
carnivore8
shallow
rock/
rocky
shelf;
often
within
crevices;
estuarine;
high
relief
rock;

highrelief
artificial
reefs
5,7,11,17,19,20,27,32,34,43,64,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,7,8,12,13
(
crustaceans;
fish,
especially
juvenile
rockfishes
like
S.
mystinus
;
cephalopods;
gastropods;

ophiuroids)
2,8,9,18,21,22,23,32,41,48,61,63,71,75,76
shallow
rocky
reef
bottoms/
rocky
shelf;
kelp
forest
bottom;
artificial
reefs
2,5,7,11,14,17,28,32,37,50,51,62,64,65,66
rocky
reefs
and
crevices;
occasionally
over
sandy
and
muddy
substrates;
artificial
reefs;

kelp
beds
1,11,17,27,31,32,33,38,40,48,56,58,64,65
(
crustaceans,
such
as
spiny
lobster;
cephalopods;

small
fish,
such
as
anchovy)
1,8,9,19,25,37,63,65,75,76
Food
habits
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
pinnipeds;
porpoise
1,2,3,5,6,8,11,12
(
crustaceans;
small
fish,
such
as
surfperches;

polychaetes)
3,9,19,24,28,44,48,50,63,75,76
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,9,12,20
(
crustceanas,
such
as
caridians
and
amphipods;

polychaetes;
small
fish,
especially
juvenile
S.

mystinus
;
cephalopods;
tunicates)
9,15,18,19,21,22,23,25,31,32,34,41,47,48,49,57,61,63,64,65,71,73,75,76
shallow
rocky
reef;
kelp
canopy;
midwater
nearshore;
artificial
reefs
5,7,14,17,27,28,29,32,37,44,45,50,53,55,62,63,64,65
Page
890
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1167
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2220e
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Scorpaenidae
1234
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CB
CC
CD
CE
Adults
Foraging
method
When
/
Frequency
Predators
Substocks
ambusher3,8
nocturnal8,13,21
NF
NF
foliage­
searcher10
nocturnal4,5,8,11,12,13,17,19,20,21
sharks;
dolphin;
seals1,4
ambusher11
dawn
&
dusk7,9
diurnal/
nocturnal18
California;
Washington7
NF
NF
NF
possibly
river
otters12
ambusher6,9,12
dawn
&
dusk;
nocturnal5,7,13,18
nocturnal12,22
Food
habits
sharks;
dolphin;
seals1,4
sharks;
dolphin;
seals;
salmon1,4,10
northern
CA
(
Ft.
Bragg);
central
CA
(
Monterey
to
San
Luis
Obispo);
southern
CA
(
Ventura/
Pt.
Loma)
1,9
Pt.
Conception
north;
Pt.
Conception
south6
Page
891
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1167
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Scorpaenidae
1234
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CF
CG
CH
CI
CJ
CK
CL
CM
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1
0.23
11.2
(
0.1)
20.8
0.06
(
0.029)
0.033
(
0.022)
allozymes
NF
30.046
to
0.068
NF
NF
11.2
(
0.1)
16.7
0.022
(
0.018)
0.022
(
0.017)
allozymes
NF
GENETICS
Mean
heterozygosity
Nucleotide
diversity
(#)
Page
892
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1167
5­
2220e
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Scorpaenidae
1234
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CN
CO
CP
CQ
CR
CS
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Fishery
/
Length
(
age)
Fishery
/
Length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
812
(
NG)
22
(
NG)
NG
NG
NG
NG
NF
NF
NF
NF
NF
NF
15NG
(
3
to
4)
NG
(
8
to
9)
hook
&
line
otoliths;
whole
NG
NG
NF
NF
NF
NF
NF
NF
Page
893
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1167
5­
2220e
­
Scorpaenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CT
CU
CV
CW
Mortality
Estimates
Method
of
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
NF
NF
NF
NF
NF
NF
NF
NF
4
NG
NG
0.2744
Hencke
survivorship/
ration
21
0.112
0.162
0.274
Hencke
survivorship/
ration/
age
frequency
NF
NF
NF
NF
MORTALITY
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9
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
CX
CY
CZ
DA
Rate
of
Exploitation
Age
range
used
Rate
of
Exploitation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
NF
NF
NF
NF
NA
NF
NF
NF
6
to
15
NG
NG
NF
6
to
15
0.142
0.591
NF
NF
NF
may
compete
with
S.
chrysomelas
1,2
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Scorpaenidae
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11
12
13
14
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
DB
DC
DD
DE
Interactions
/
associations
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

2190
km
Transient2
NF
9>
50
km
22up
to
220
miles
parasitized
by
copepod
family
Naobranchia
19
infested
by
roundworms
from
genera
Anisakis
20
5<
5
nm
Residential+
DE43
NF
no
movement6,3,12,13,20
infested
by
roundworms
from
genera
Anisakis
20
5<
5
nm
Residential3,5,12,19
Yes3,4,8
no
movement6,3,12
sub­
adults
are
residential15
parasitized
by
copepod
family
Neobrachiella19
9>
50
km
infested
by
roundworms
from
genera
Anisakis
20
11<
2
km
18<
1.3
km
5<
5
nm
Residential
5,8,15
NF
10,11<
2
km
2(
5.54
m2)

no
movement6,8
parasitized
by
copepods
Chondracanthus
pinguis
,
Clavella
parva
,
and
Clavellopsis
robusta
11
infected
by
parasites
from
the
families
Chondracanthidae,

Lernaeopodidae,
Sanguinicolidae,
Capsalidae,

Microcotylidae,
and
Anisakidae22
infected
by
parasites
from
the
families
Caligidae,

Naobranchiidae,
Sanguinicolidae,
Gnathiid
larvae,

Capsalidae,
Microcotylidae,
Ceratomyxidae,
and
Myxidiidae22
associated
and
may
compete
for
resources
with
S.

chrysomelas;
usually
found
deeper
than
S.
chrysomelas
5,10,16
competitively
exlcuded
from
bottom
territories
by
S.

carnatus
16,17
competitively
excludes
S.
atrovirens
from
bottom
territories16,17
decrease
in
abundance
as
Macrosytis
beds
are
removed18,21
infected
by
parasites
from
the
families
Caligidae,

Hemiuridae,
Lepocreadiidae,
Opecoelidae,
Sanguinicolidae,

Gnathiid
larvae,
Capsalidae,
Microcotylidae,
and
Anisakidae22
BEHAVIOR
migrates
to
deeper
water
in
winter16
juveniles
migrate
to
open
ocean
as
they
mature20,21
infected
by
parasites
from
the
families
Bomolochidae,

Caligidae,
Chondracanthidae,
Opecoelidae,
Piscicolidae,

Cymothoidae,
Capsalidae,
Trochopodinae,
Anisakidae,

Ceratomyxidae,
and
Myxidiidae22
infected
by
ectoparasites
from
the
families
Bomolochidae,

Caligidae,
Chondracanthidae;
also
monogenetic
trematodes9
infected
by
ectoparasites
from
the
families
Caligidae
and
Lerneopodidae;
also
gnathiid
larvae
and
monogenetic
trematodes9
infected
by
ectoparasites
from
the
families
Caligidae;
also
gnathiid
larvae9
Movement
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21
22
23
24
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
DF
DG
Schooling
Territoriality
No2,10,32
NF
Ocassionally4
No10
No11,35
No23
No12
Yes­
as
juveniles27
No3,11,36
Yes2,3,4,5,8,10,16
aggregate
during
spawning
season34,35
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68
69
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
A
B
C
D
E
S.
caurinus
1
1
<
1
to
183
(
S.
vexillaris)
2
S.
chlorostictus
1
1
30
to
273
S.
chrysomelas
1
1
<
1
to
37
S.
constellatus
1
1
24
to
274
S.
dalli(
i)
1
calico
rockfish;
pretty
rock
cod1,4,9
1
7
to
256
S.
elongatus
1
1
25
to
495
copper
rockfish;
whitebelly;
white;
whisky;

fighting
Bob;
borrego;
bravo;
chucklehead;

barriga
barriga;
never
dies;
palermotana;

northern
rockfish1,3,4,5,6,9,12
greenstriped
rockfish;
poinsetta;

strawberry;
serena;
rainha;
reina;
sugar
bass;
striped
rockfish;
cucumber
1,4,6,8,12,13
starry
rockfish;
scacciatale;
spotted
rockfish;
spotted
corsair;
chinafish
1,4,5,7,9,12,13
greenspotted
rockfish;
bosco;
warthog;

starry
eyes;
scrub;
Santa
Maria;
Callahan;

popeye;
chucklehead;
chinafish;
cernie;

pesce;
vermiglia;
bolina1,3,4,5,6,9,12,13
black­
and­
yellow
rockfish;
zurndicky;

china
cod;
gopher
cod;
cifulatano1,3,4,6,9,12,13
northern
CA
(
Cordell
Bank)
to
southern
Baja
California
(
Thetis
Bank)
1
northern
CA
(
San
Francisco)
to
central
Baja
California
(
Bahia
de
Sebastian
Vizcaino)
1
northern
Gulf
of
alaska
(
Kachemak
Bay)
to
central
Baja
California
(
Islas
San
Benito)
1
WA
(
Copalis
Head)
to
central
Baja
California
(
Isla
Cedros)
1
northern
CA
(
Eureka)
to
central
Baja
California
(
Isla
Natividad)
1
western
Gulf
of
Alaska
(
Chirikof
I.,
Aleutians
Is.)

to
central
Baja
California
(
Isla
Cedros)
1,4
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71
72
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79
80
81
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83
84
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87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
F
G
H
I
J
5Both:
W=
0.00001623FL3.0402
(
Sept.­
Nov.)
4TL=
3.873
+
1.209(
SL);
r2=
0.994;
SE=
5.8
1
58
Washington
(
TL)
19
52
5Both:
W=
0.00002495FL2.9533
(
Dec.­
Mar.)
4TL=
0.629
+
1.01(
FL);
r2=
0.998;
SE=
3.0
Washington
(
TL)
17
70
(
6.075)

8*
Male:
W=
0.00002106TL2.981
(
r2=
0.964)
4SL=
­
0.653
+
0.82(
TL);
r2=
0.994;
SE=
4.6
Washington
(
TL)
18
NG
8*
Female:
W=
0.00001088TL3.100
(
r2=
0.990)
4SL=
­
2.272
+
0.836(
FL);
r2=
0.992;
SE=
5.7
Washington
(
TL)
37
52
8*
Both:
W=
0.000008976TL3.132
(
r2=
0.933)
4FL=
0.005
+
0.998(
TL);
r2=
0.998;
SE=
3.0
c.
California
(
TL)
11
51.7
13*
Both:
W=
0.0000095TL3.1
(
r2=
0.995)
4FL=
5.896
+
1.187(
SL);
r2=
0.992;
SE=
6.8
14*
Male:
W=
0.0000046TL2.84
(
r2=
0.960)

14*
Female:
W=
0.00000299TL2.92
(
r2=
0.951)

14*
Both:
W=
0.00000311TL2.91
(
r2=
0.967)

15*
Both:
W=
0.0000101TL3.09
28Male:
W=
0.000000046TL2.84
(
r2=
0.96)

28Female:
W=
0.00000002991TL2.92
(
r2=
0.951)

28Both:
W=
0.0000000311TL2.91
(
r2=
0.967)

Both1:
W=
0.00905TL3.16321
(
r2=
0.990)
4TL=
5.316
+
1.202(
SL);
r2=
0.998;
SE=
3.7
1
50
c.
California
(
TL)
11
44
8*
Male:
W=
0.000002827TL3.291
(
r2=
0.988)
4TL=
­
0.723
+
1.028(
FL);
r2=
1.0;
SE=
1.9
California
(
SL)
14
NG
8*
Female:
W=
0.000004773TL3.21
(
r2=
0.980)
4SL=
­
3.931
+
0.83(
TL);
r2=
0.998;
SE=
3.0
8*
Both:
W=
0.00001558TL3.001
(
r2=
0.980)
4SL=
­
3.987
+
0.852(
FL);
r2=
0.998;
SE=
3.2
11Both:
W=
0.0000115SL3.18
(
SL=
mm)
4FL=
0.858
+
0.972(
TL);
r2=
1.0;
SE=
1.9
4FL=
5.289
+
1.171(
SL);
r2=
0.998;
SE=
3.7
8*
Male:
W=
0.0000142TL3.07
(
r2=
0.943)
4TL=
1.007
+
1.211(
SL);
r2=
0.996;
SE=
3.2
1
39
c.
California
(
TL)
11
32.6
8*
Female:
W=
0.00001024TL3.13
(
r2=
0.960)
4SL=
­
0.123
+
0.822(
TL);
r2=
0.996;
SE=
2.6
c.
California
(
SL)
26
24.7
8*
Both:
W=
0.00001117TL3.114
(
r2=
0.982)
4SL=
­
0.009
+
0.822(
FL);
r2=
0.994;
SE=
2.6
s.
California
(
SL)
26
19.9
19Both:
W=
0.000017SL3.13
(
r2=
0.98)
4FL=
1.137
+
1.209(
SL);
r2=
0.994;
SE=
3.2
19Both:
W=
0.000174SL2.67
(
r2=
0.96)

1Both:
W=
0.00967TL3.15979
(
r2=
0.957)
4TL=
4.497
+
1.175(
SL);
r2=
0.998;
SE=
3.1
1
46
s.
California
(
TL)
10
38.06
(
1.68)

8*
Male:
W=
0.000001594TL3.371
(
r2=
0.823)
4TL=
­
0.134
+
1.023(
FL);
r2=
1.0;
SE=
1.5
c.
California
(
TL)
11
46
8*
Female:
W=
0.00001486TL3.009
(
r2=
0.960)
4SL=
­
3.204
+
0.849(
TL);
r2=
0.998;
SE=
2.7
8*
Both:
W=
0.000008537TL3.112
(
r2=
0.972)
4SL=
­
2.774
+
0.866(
FL);
r2=
0.998;
SE=
2.6
4FL=
0.301
+
0.977(
TL);
r2=
1.0;
SE=
1.5
4FL=
3.883
+
1.152(
SL);
r2=
0.998;
SE=
3.0
1Both:
W=
0.00945TL3.21542
(
r2=
0.925)
NF
1
20
California
(
SL)
14
NG
1Both:
W=
0.00793TL3.12745
(
r2=
0.986)
4TL=
15.24
+
1.165(
SL);
r2=
0.996;
SE=
3.5
2
43
(
FL)
British
Columbia
(
FL)
2
37
4Male:
W=
0.000003FL3.2597
(
r2=
0.981)
4TL=
­
0.552
+
1.047(
FL);
r2=
1.0;
SE=
1.5
CA
to
WA
(
FL)
9
30.01
(
3.5)

4Female:
W=
0.000004FL3.2145
(
r2=
0.988)
4SL=
­
12.14
+
0.855(
TL);
r2=
0.996;
SE=
3.0
s.
California
(
TL)
10
29.65
(
0.55)

4SL=
­
11.72
+
0.892(
FL);
r2=
0.996;
SE=
3.1
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899
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68
69
70
71
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74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
K
L
M
N
O
P
Q
R
S
T
U
46
NG
0.12
0.16
NG
­
1.9
­
1.3
NG
NG
NG
NG
78
(
1.533)
59
(
1.421)
0.051
(
0.0106)
0.042
(
0.0162)
0.083
(
0.004)
­
2.6
(
1.127)
­
3.9
(
1.746)
­
0.56
(
0.082)
NG
NG
NG
NG
40
NG
NG
0.183
NG
NG
­
0.6061
NG
NG
NG
46
50
0.12
0.16
0.12
­
1.9
­
1.3
­
0.1
NG
NG
NG
57.2
56.5
0.2235
0.1269
0.1354
0.1
­
1.3
­
1
NG
NG
NG
44.2
44.4
0.1359
0.1311
0.1282
0
0
­
0.2
NG
NG
NG
NG
39.2
NG
NG
0.0582
NG
NG
­
0.87
NG
NG
NG
33.9
33.6
0.2409
0.2125
0.2165
­
0.3
­
0.6
­
0.5
NG
NG
NG
25.2
NG
0.24
0.22
NG
­
0.32
­
0.45
NG
NG
NG
NG
21.5
NG
0.28
0.21
NG
­
0.28
­
0.72
NG
NG
NG
NG
45
(
2.87)
NG
0.094
(
0.014)
0.087
(
0.009)
NG
­
1.065
(
0.712)
­
3.107
(
0.822)
NG
NG
NG
NG
45
46.7
0.1208
0.1241
0.1148
­
0.1
0
­
0.1
NG
NG
NG
NG
16
NG
NG
0.12196
NG
NG
­
2.95
NG
NG
NG
35.2
38.6
0.077
0.12
0.075
­
2.7
­
0.1
­
2.5
NG
NG
NG
37.45
(
9.1)
NG
0.108
(
0.01)
0.079
(
0.008)
NG
­
3.273
(
0.7)
­
3.465
(
0.889)
NG
0.826
0.825
NG
37.26
(
0.66)
NG
0.121
(
0.009)
0.098
(
0.006)
NG
­
2.732
(
0.334)
­
2.36
(
0.288)
NG
NG
NG
NG
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Scorpaenidae
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71
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80
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88
89
90
91
92
93
94
95
96
97
98
99
100
101
V
W
X
Y
otoliths;
whole
No
No
6
otoliths;
whole
No
No
15
(
juveniles)

otoliths;
whole
No
Intrareader
comparisons
otoliths;
whole
NG
NG
otoliths;
whole
No
Interreader
comparisons
otoliths;
whole
No
between
readers
NF
otoliths;
whole
No
No
otoliths;
whole
Edge
Analysis
Interreader
comparisons;
comparisons
with
operculum
NF
otoliths;
whole
OTC
Intra/
interreader
comparisons
otoliths;
whole
OTC
Intra/
interreader
comparisons
otoliths;
section
Edge
Analysis;
<
6
yrs
Intrareader
comp.
NF
otoliths;
whole
No
Interreader
comparisons
otoliths;
whole
No
No
NF
otoliths;
whole
No
No
NF
otoliths;
b
&
b
No
Interreader
comparisons
otoliths;
section
Edge
Analysis;
<
6
yrs
Intrareader
comparisons
Page
901
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1167
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Scorpaenidae
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58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
Z
AA
AB
AC
AD
AE
AF
17
55
2,3,7,11,13,18
Dioecious
Polygynous2
Viviparous3,7
No
1
0.98
15,21,32
35
(
matrotrophy)
4
33,34
41
Ovoviviparous10
10
>
21
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
1.02
21
33
4
20
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
1.07
8~
22
Ovoviviparous12
10,21,33
21
10,35
28
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
1.1
21,36
32
21,33
12
NF
Dioecious
Polygynous2
Viviparous3,7
No
NA
35
11
7
46
7
Dioecious
Polygynous2
Viviparous3,7
No
1
0.87
21,33,36
37
(
Males);
28
(
Females)
Ovoviviparous10
Page
902
of
1167
5­
2220e
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Scorpaenidae
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61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
AG
AH
AI
AJ
AK
AL
AM
None1
3,53
30
(
3)
31
(
5)
32
(
4)
34
(
6)
otoliths;
whole
No
237
(
8)
29.5
(
NA)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
15,41NG
(
NG)
NG
(
3)
NG
(
NG)
NG
(
4)
otoliths;
whole
No
18NG
(
5)
NG
(
5)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
25NG
(
NG)
NG
(
NG)
25
(
6)
25
(
6)
NG
NG
30NG
(
NG)
NG
(
NG)
25
(
6)
NG
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
34
(
6)­
sex
unspecified
NG
NG
3331
(
3)
31
(
5)
33
(
4)
34
(
6)
NG
NG
None1
325
(
4)
26
(
5)
27
(
6)
28
(
6)
otoliths;
whole
No
5,5320
(
NG)
15
(
NG)
22
(
NG)
22
(
NG)
NG
NG
233.3
(
11)
29.9
(
10)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
32NG
(
NG)
NG
(
NG)
28
(
6
to
9)­
sex
unspecified
NG
NG
None1
3,5214
(
3)
14
(
3)
16
(
3)
15
(
3)
otoliths;
whole
No
2,5223.9
(
4)
24.3
(
6)
NG
(
NG)
NG
(
NG)
otoliths;
whole
Edge
Analysis
1013.5^
(
3)
13.5
(
3)
NG
(
NG)
NG
(
NG)
otoliths;
whole
NG
2814^
(
3)
16
(
3)
16.5
(
3)
16
(
4)
otoliths;
whole
OTC
2813^
(
3)
14
(
3)
13
(
3)
13.5
(
3)
otoliths;
whole
OTC
32NG
(
NG)
NG
(
NG)
15
(
3)­
sex
unspecified
None1
328
(
6)
23
(
5)
30
(
7)
27
(
6)
otoliths;
whole
No
518
(
NG)
21
(
NG)
19
(
NG)
22
(
NG)
NG
NG
226.5
(
NG)
27
(
7)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
32NG
(
NG)
NG
(
NG)
27
(
6)­
sex
unspecified
NG
NG
53NG
(
NG)
NG
(
NG)
20.3
(
6
to
7)
25.4
(
8
to
9)
NG
NG
None1
57(
NG)
9
(
NG)
9
(
NG)
9
(
NG)
NG
NG
32,53NG
(
NG)
NG
(
NG)
9
(
3
to
4)­
sex
unspecified
NG
NG
None1
3,52
23
(
7)
18
(
5)
23
(
7)
23
(
7)
otoliths;
whole
NG
615*
(
NG)
15
(
NG)
23
(
10)
21
(
7)
otoliths;
b
&
b
No
515
(
NG)
16
(
NG)
18
(
NG)
19
(
NG)
NG
NG
11NG*(
NG)
NG
(
NG)
23
(
NG)
23
(
NG)
NG
NG
Page
903
of
1167
5­
2220e
­
Scorpaenidae
52
53
54
55
56
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
AN
AO
AP
AQ
AR
AS
AT
AU
AV
No
2
Alaska67,71,77
interreader
comparisons
Washington7,8,9,11,12,35,42,53,59,70
^^^^^
*****

No
Oregon66
No
n.­
c.
California5
^^^^^
*****

NG
c.
California4,51
NG
California57,59
NG
NG
No
1
Oregon5,66
NG
n.­
c.
California5,18,77
^^^^^
*****

interreader
comparisons
c.
California4
NG
s.
California13,77
*****

California57
*****
*****

No
NF
n.­
c.
California5,76
^^^^^
*****

interreader
comparisons;
comparison
with
operculum
c.
California4,50,78
NG
s.
California10,14,50,76
intra/
interreader
comparisons
California57
*****
*****

intra/
interreader
comparisons
No
NF
n.­
c.
California5,18
^^^^^
*****

NG
c.
California4,77
interreader
comparisons
s.
California13,77
*****

NG
California57
*****
*****

NG
NG
1
s.
California13,77
*****

NG
California57
*****

NG
NF
Alaksa71
between
readers
British
Columbia15
*****

NG
Oregon16,19,66,70
*****

NG
n.­
c.
California5,77
^^^^^
*****

Page
904
of
1167
5­
2220e
­
Scorpaenidae
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Once6
1183
(
38­
301)
6,9,1115,642
(
24.2
TL)
to
640,052
(
47.4
TL)

22
97,000
(
33
?
L)
to
654,000
(
51
?
L)

Multiple
2
3148
(
NA)
1,2740,961(
27.6
TL)
to
759,040
(
39
TL)

Once17
NF
1699,000
(
20
SL)

^^^^^
16110,000
(
20
SL)

Multiple1,2
1160
(
87­
194)
1,27
32,674
(
24
TL)
to
227,573
(
32.6
TL)

3198
(
NA)

Once2
3155
(
NA)
1,27
3,878
(
11.6
TL)
to
18,006
(
15.5
TL)

26
2,000
(
NG)
to
113,000
(
NG)

4
1,700
(
NG)

Multiple
2,22,23
3325
(
NA)
1,27
10,637
(
16.0
TL)
to
295,153
(
34.4
TL)

Once5
26
26,000
(
NG)
to
344,000
(
NG)

Page
905
of
1167
5­
2220e
­
Scorpaenidae
52
53
54
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56
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58
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
BF
BG
BH
BI
BJ
*
2,7,14F=
0.0000000026TL5.347
NA
1
~
1
to
2
months
9,1040
to
50
NF
*
6,7F=
0.0000000274TL4.957
See
Richards
&
Schnute
(
1990)
for
additional
discussion
3F=
0.000005TL4.971
(
r2=
0.843)
NA
1
~
1
to
2
months
NF
NF
*
10F=
0.0000136(
SL)
5.59
NA
~
1
to
2
months1
940
to
50
(
SL)
NF
*
10F=
0.0000464(
SL)
4.09
3F=
0.000066TL4.2514
(
r2=
0.731)
NA
1
~
1
to
2
months
NF
NF
3F=
0.000211TL4.0981
(
r2=
0.591)
NA
1
~
1
to
2
months
420
to
25
NF
3F=
0.000427TL3.7392
(
r2=
0.898)
NA
1
~
1
to
2
months
11~
30
NF
Page
906
of
1167
5­
2220e
­
Scorpaenidae
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
BK
BL
BM
BN
BO
BP
5
0.62
No
external
eggs
NA
10,22,245
to
23
(
SL)
planktonic1
planktivore
11,17,18,235.3
(
SL)
to
205.1
(
SL)
to
NF
No
external
eggs
NA
20,254.2
(
SL)
to
planktonic1
planktivore
120.78
to
0.91
No
external
eggs
NA
22to
25
(
SL)
planktonic1
planktivore
120.68
to
0.75
5
0.14
to
0.82
No
external
eggs
NA
17,194.0
(
SL)
to
planktonic1
planktivore
254.2
(
SL)
to
NF
No
external
eggs
NA
16,185.0
to
20.0
(
SL)
planktonic1
planktivore
254.9
(
SL)
to
NF
No
external
eggs
NA
95
(
TL)
to
planktonic1
planktivore
204.7
(
TL)
to
254.9
(
SL)
to
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
Page
907
of
1167
5­
2220e
­
Scorpaenidae
52
53
54
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59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
BQ
BR
BS
BT
BU
BV
NF
diurnal1
siphonophores;
23
(
SL)
to
maturity9
zooplanktivore
chaetognaths1
diurnal1
siphonophores;
NA
pelagic?;
NF3,9,22
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
25
(
SL)
to
maturity9
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
pelagic?;
NF3,9,22
zooplanktivore
chaetognaths1
(
crustaceans,
especially
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
6,820.0
(
SL)
to
maturity
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6,15
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids
and
calenoid
copepods;
small
fish)
1,6,11
pelagic;
soft
sediment;
sand­
silt
interface;

artificial
reefs3,9,18,26
(
crustaceans,
such
as
calanoids,
gammarids,

and
barnacle
cyprids;
juvenile
polychaetes;

fish)
1,6,8,9,10,12,16,20
pelagic;
seagrass;
kelp
canopy;
low
growing
algae;
rock
and
cobble;
high
relief
rock;
sand
and
low
rock;
low­
relief
artificial
reefs
2,3,9,14,13,15,16,24,26,27,29,30,33,37,40,44,49,54
pelagic;
kelp
canopy;
rocky
reefs;
artificial
reefs
1,2,3,9,13,26,53
pelagic;
sand
and
mud;
bedrock
and
cobble;
caves
and
crevices;
artificial
reefs
3,4,9,22,23,25,26
Page
908
of
1167
5­
2220e
­
Scorpaenidae
52
53
54
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
BW
BX
BY
BZ
CA
NF
diurnal4,5
fishes,
including
rockfishes,
carnivore
lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,9,12,15
NF
NF
carnivore
(
crustaceans;
cephalopods;
fish)
20,33,48,76
NF
NF
carnivore
NF
NF
(
meso)
carnivore8
(
crustaceans;
cephalopods;
fish)
8,20,48,76
picker1
diurnal11
(
meso)
carnivore8
picker1
diurnal11
carnivore
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
rocky
shelf;
mud­
rock
or
sand­
mud
interface;

fine­
sediments;
high
&
low
relief;
artificial
reefs
5,7,9,18,21,25,32,35,43,62,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
shallow
rocky
reefs;
kelp
beds;
artificial
reefs
2,5,7,11,14,17,27,32,37,50,51,53,63,62,64,65,66
shallow
rock/
high­
relief
rocky
shelf;

vegetation;
rock­
sand;
interface;
eelgrass
beds;
high­
relief
artificial
reefs4,5,7,19,20,27,32,34,35,36,37,43,52,65
(
small
fish,
including
surfpearches,
juvenile
greenlings,
sculpins,
and
gobies;
spiny
dogfish;

crustaceans;
cephalopods;
lingcod
eggs)

3,12,13,17,22,23,24,26,28,29,32,33,38,39,44,48,51,63,76
(
crustaceans;
fish,
especially
juvenile
S.
mystinus
;

cephalopods;
ophiuroids;
chitons)

2,15,18,19,21,22,23,32,41,48,49,61,65,71,73,76
rocky
shelf;
continental
slope/
basin;
high
relief
5,7,18,65
(
pelagic
crustaceans,
such
as
calanoid
copepods)
33,37,48,63,76
estuarine;
rocky/
non­
rocky
shelf;
shallow
sandy/
soft
bottoms;
artificial
reefs
5,17,18,27,31,32,46,49,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,7,8,12
(
crustaceans,
such
as
euphausiids;
fish;

cephalopods)
33,37,48,76
rocky
shelf;
hard
or
mixed
bottom;
high
releif;
cobble­
mub
5,7,18,25,65
Page
909
of
1167
5­
2220e
­
Scorpaenidae
52
53
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55
56
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59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CB
CC
CD
CE
NF
California;
Washington;
Alaska7
dawn2,3
possibly
river
otters12
NF
NF
NF
ambusher6,9,12
dawn
&
dusk;
nocturnal5,7,13,18,20
nocturnal12,22
NF
NF
NF
searcher/
persuer8
nocturnal13
NF
picker/
pursuer8
nocturnal?
13
NF
possibly
river
otters12
sharks;
dolphin;
seals1,4
dawn
&
dusk;
nocturnal1,5,13,14
sharks;
dolphin;
seals;
salmon1,4,10
sharks;
dolphin;
seals1,4
northern
CA
(
Ft.
Bragg);
central
CA
(
Monterey
to
San
Luis
Obispo);
southern
CA
(
Ventura/
Pt.
Loma)
1
sharks;
dolphin;
seals1,4
sharks;
dolphin;
seals1,4
sharks;
salmon;
dolphin;
seals1,4,10
Page
910
of
1167
5­
2220e
­
Scorpaenidae
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CF
CG
CH
CI
CJ
CK
CL
CM
NF
1
1.55
11.2
(
0.1)
12.5
0.047
(
0.03)
0.03
(
0.022)
allozymes
NF
NA
NA
30.035
to
0.067
NA
allozymes
2
1.04
4
&
6.7
0.018
&
0.02
NA
allozymes
NF
NF
11.0
(
0.0)
4.2
0.013
(
0.013)
0.014
(
0.014)
allozymes
NF
NF
NF
11.1
(
0.1)
12.5
0.037
(
0.025)
0.042
(
0.029)
allozymes
NF
NF
NF
11.2
(
0.1)
16.7
0.063
(
0.032)
0.014
(
0.01)
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
11.1
(
0.1)
8.3
0.006
(
0.005)
0.006
(
0.005)
allozymes
NF
2
1.08
8.3
&
6.7
0.034
&
0.042
NA
allozymes
Page
911
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1167
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2220e
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Scorpaenidae
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CN
CO
CP
CQ
CR
CS
3
26.7(
5)
31
(
8)
hook
&
line;
spearfishing
otoliths;
whole
No
No
15NG
(
3
to
4)
NG
(
8
to
9)
hook
&
line
otoliths;
whole
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
711
(
2)
NG
(
7
to
13)
trawls
otoliths;
break­
and­
burn
No
intereader
comparisons
Page
912
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1167
5­
2220e
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Scorpaenidae
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54
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56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CT
CU
CV
CW
1
0.1127
0.0686
0.1813
tag
&
recapture
4
NG
NG
0.2329
Hencke
survivorship/
ration
21
0.131
0.101
0.232
Hencke
survivorship/
ration/
age
frequency
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
9
0.1­
Males;
0.12­
Females
0.33­
Males;
0.21­
Females
0.43Males;
0.31­
Females
Hoenig
estimation/
Z­
formula
9
0.1­
Males;
0.12­
Females
0.33­
Males;
0.19­
Females
0.43Males;
0.31­
Females
catch
curve/
Z­
formula
9
0.15­
Females
0.16­
Females
0.43Males;
0.31­
Females
GSI/
Z­
formula
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913
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1167
5­
2220e
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Scorpaenidae
52
53
54
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56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
CX
CY
CZ
DA
5
to
34
0.0627
0.378
NF
9
to
18
NG
NG
9
to
18
0.09
0.435
NF
NF
NF
NF
NF
NF
NF
may
compete
with
S.
carnatus
1,2
NF
NF
NF
NF
NF
NF
NF
NF
2
to
46
NG
0.77(
M)/
0.68(
F)
NF
2
to
46
NG
0.77(
M)/
0.61(
F)

2
to
46
NG
0.55(
F)
Page
914
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5­
2220e
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Scorpaenidae
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57
58
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60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
DB
DC
DD
DE
may
contain
parasite
Hemiurus
levinseni
6
5,9<
5
nm
Transient/
Residential1,3,5
Yes4,5,8
associated
with
S.
maliger
and
S.
nigrocinctus8
31.5
miles
Residential12
infested
by
roundworms
from
genera
Phocanema
20
no
movement7,8,18
<
1km1
Residential11
NF
5<
5
nm
Residential8,11,15,23
Yes2,6,10
10,11<
2
km
no
movement6,3,8,21
NF
Residential?
11
NF
infested
by
roundworms
from
genera
Anisakis
20
parasitized
by
copepod
family
Neobrachiella19
NF
Residential
5
NF
infected
by
parasites
from
the
genus
Anisakis
22
associated
with
S.
rosenblatti
7
NF
Residential11
NF
co­
occurs
with
S.
chlorostictus
12
migrates
to
deeper
water
in
winter14,16
infected
by
macroparasites
from
the
families
Microcotylidae,

Bothriocephalidae,
and
Anisakide14
Adults
residential;
juveniles
show
significant
movement
before
maturity19
infected
by
ectoparasites
from
the
families
Caligidae
and
Lerneopodidae;
also
gnathiid
larvae
and
monogenetic
trematodes9
infected
by
parasites
from
the
families
Bothriocephalidae,

Philichthyidae,
Microcotylidae,
Anisakidae,
Ceratomyxidae,

and
Myxobolidae22
infected
by
macroparasites
from
the
families
Microcotylidae
and
Anisakide14,20
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
20
associated
and
may
compete
for
resources
with
S.
carnatus
;

usually
found
shallower
than
S.
carnatus
and
thought
to
exclude
it
from
shallower
depths5,10,16
infected
by
parasites
from
the
families
Hemiuridae,
Gnathiid
larvae,
and
Cymothoidae22
infected
by
parasites
from
the
families
Echinorhynchidae,

Polymorphidae,
Argulidae,
Bothriocephalidae,

Phyllobothriidae,
Tentaculariidae,
Bomolochidae,
Caligidae,

Chondracanthidae,
Lernaeopodidae,
Naobranchiidae,

Philichthyidae,
Acanthocolpidae,
Gorgoderidae,
Hemiuridae,

Lepcreadiidae,
Opecoelidae,
Sanguinicolidae,
Zoogonidae,

Piscicolidae,
Capsalidae,
Microcotylidae,
Anisakidae,

Capillariidae,
Cucullanidae,
Cystidicolidae,
and
Myxidiidae22
infected
by
macroparasites
from
the
families
Microcotylidae,

Bothriocephalidae,
Anisakide,
and
Polymorphidae14
infected
by
parasites
from
the
families
Lepocreadiidae,

Opecoelidae,
Sanguinicolidae,
Gnathiid
larvae,
and
Cymothoidae22
Page
915
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1167
5­
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­
Scorpaenidae
52
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55
56
57
58
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62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
DF
DG
No7,23
No12
Yes19
Yes­
as
juveniles27,28
No23,35
NF
No3,4,11,36
Yes2,3,4,5,8,10,13,16
No
23
NF
No16,18
NF
No18,33
NF
Page
916
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Scorpaenidae
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103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
A
B
C
D
E
S.
helvomaculatus
1
1
25
to
549
S.
hopkinsi
1
1
18
to
183
S.
maliger
1
1
<
1
to
274
S.
melanops
1
1
<
1
to
366
quillback
rockfish;
orange­
spotted
rockfish;
yellow­
back
rockfish;
speckled
rockfish;
stickleback1,4,7,9,12
rosethorn
rockfish;
flyfish;
deep­
water
sacciatale;
rosy;
deep­
water
scratchtail1,4,8,12
squarespot
rockfish;
mustard;
perch;

smallmouth
rockfish;
widow;
Belinda
bass1,4,6,7,12
black
rockfish;
bass;
black
bass;
black
snapper;
black
rock
cod;
cherna;
nero;

pesce
pretre1,3,4,5,9,12,13
Gulf
of
Alaska
(
Kenai
Peninsula)
to
southern
CA
(
San
Miguel
I.)
1,7
western
Gulf
of
alaska
(
east
of
Sitkinak
I.)
to
central
Baja
California1
Amchitka
I.,
Aleutian
Is.
to
southern
CA
(
Huntington
Beach)
1
southern
OR
to
Guadalupe
Is.
and
central
Baja
California1,6
Page
917
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Scorpaenidae
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103
104
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106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
F
G
H
I
J
4FL=
0.701
+
0.954(
TL);
r2=
1.0;
SE=
1.4
4FL=
14.19
+
1.116(
SL);
r2=
0.996;
SE=
3.5
4Male:
W=
0.000003FL3.2597
(
r2=
0.981)
NF
1
41
British
Columbia
(
FL)
2
30.6
4Female:
W=
0.000007FL3.2145
(
r2=
0.988)
CA
to
WA
(
FL)
9
27.93
(
7.7)

1Both:
W=
0.01464TL2.96355
(
r2=
0.927)
4TL=
3.059
+
1.2(
SL);
r2=
0.986;
SE=
4.8
1
29
s.
California
(
TL)
10
24.71
(
10.77)

4TL=
0.002
+
1.041(
FL);
r2=
0.998;
SE=
1.9
4SL=
­
0.195
+
0.822(
TL);
r2=
0.986;
SE=
4.0
4SL=
­
0.372
+
0.856(
FL);
r2=
0.988;
SE=
3.8
4FL=
0.428
+
0.959(
TL);
r2=
0.998;
SE=
1.8
4FL=
3.011
+
1.153(
SL);
r2=
0.988;
SE=
4.5
13*
Both:
W=
0.000012TL3.07
(
r2=
0.994)
4TL=
1.463
+
1.22(
SL);
r2=
0.992;
SE=
5.6
1
61
Alaska
(
FL)
13
42
14*
Male:
W=
0.00000497TL2.83
(
r2=
0.925)
4TL=
­
8.696
+
1.034(
FL);
r2=
0.998;
SE=
2.7
Alaska
(
FL)
22
42
to
46
14*
Female:
W=
0.00000305TL2.92
(
r2=
0.910)
4SL=
­
1.12
+
0.813(
TL);
r2=
0.992;
SE=
4.6
Washington
(
TL)
17
45
(
1.396)

14*
Both:
W=
0.00000295TL2.92
(
r2=
0.951)
4SL=
­
8.202
+
0.848(
FL);
r2=
0.994;
SE=
4.1
Washington
(
TL)
18
NG
15*
Both:
W=
0.0000126TL3.064
4FL=
9.075
+
0.965(
TL);
r2=
0.998;
SE=
2.7
Washington
(
TL)
19
45
17*
Both:
W=
0.007711FL3.26
(
r2=
0.99)
4FL=
11.84
+
1.173(
SL);
r2=
0.994;
SE=
4.9
Washington
(
TL)
37
45
28Male:
W=
0.0000000497TL2.83
(
r2=
0.925)

28Female:
W=
0.0000000305TL2.92
(
r2=
0.91)

28Both:
W=
0.0000000295TL2.92
(
r2=
0.951)

17*
Both:
W=
0.00606FL3.29
(
r2=
0.96)
4TL=
7.724
+
1.221(
SL);
r2=
0.998;
SE=
5.2
1
91
Alaska
(
FL)
13
54
15*
Both:
W=
0.0000115TL3.054
4TL=
1.595
+
1.017(
FL);
r2=
1.0;
SE=
2.1
Alaska
(
FL)
22
54
13*
Both:
W=
0.000014TL3.027
(
r2=
0.996)
4SL=
­
5.596
+
0.817(
TL);
r2=
0.998;
SE=
4.2
Alaska
(
NA)
29
47.5;
49.1
8*
Male:
W=
0.00005254TL2.806
(
r2=
0.929)
4SL=
­
5.427
+
0.834(
FL);
r2=
0.998;
SE=
4.2
Washington
(
TL)
18
NG
8*
Female:
W=
0.000007156TL3.15
(
r2=
0.970)
4FL=
­
1.421
+
0.983(
TL);
r2=
1.0;
SE=
2.1
Washington
(
TL)
19
NG
8*
Both:
W=
0.00000581TL3.187
(
r2=
0.980)
4FL=
7.194
+
1.197(
SL);
r2=
0.998;
SE=
5.0
Washington
(
TL)
17
61
(
3.064
)

10Male:
W=
0.000025FL2.922
Washington
(
FL)­
hook
&
line33
49.47
10Female:
W=
0.0000117TL3.126
Washington
(
FL)­
trawl33
50.94
12*
Male:
W
(
kg)=
0.000025FL2.922
Washington
(
TL)
37
NG
12*
Female:
W
(
kg)=
0.0000117FL3.126
12*
Both:
W
(
kg)=
0.0000184FL3.0
14*
Male:
W
=
0.00000115TL3.05
(
r2=
0.955)

14*
Female:
W
=
0.0000013TL3.03
(
r2=
0.955)
Oregon
(
FL)
16
49.7
(
1.5)

14*
Both:
W=
0.000000819TL3.11
(
r2=
0.955)
Washington/
Oregon
(?
L30
NG
23*
Both:
W
(
kg)=
0.018FL2.992
Washington/
Oregon
(?
L)
35
49.25
Oregon
(
FL)
4,21
50.3
&
52.03
(
C.
I.=
49.1­
51.5
&

50.5­
53.6)

Page
918
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1167
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2220e
­
Scorpaenidae
102
103
104
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106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
K
L
M
N
O
P
Q
R
S
T
U
31.9
31.8
0.092
0.079
0.08
­
3.6
­
4.2
­
4.2
NG
NG
NG
28.66
(
16.9)
NG
0.133
(
0.016)
0.101
(
0.018)
NG
­
2.074
(
0.519)
­
2.772
(
0.601)
NG
0.764
0.726
NG
25.25
(
0.43)
NG
0.063
(
0.082)
0.176
(
0.021)
NG
­
10.329
(
8.66)
­
3.358
(
0.708)
NG
NG
NG
NG
44
43
0.12
0.1
0.11
­
1
­
1.4
­
1.2
NG
NG
NG
44
to
46
NG
0.08
to
0.12
0.07
to
0.1
NG
­
2.3
to
­
1
­
3.2
to
­
1.4
NG
NG
NG
NG
47
(
1.523)
47
(
9.176)
0.1
(
0.01345)
0.092
(
0.012)
0.089
(
0.0053)
­
1.1
(
0.854)
­
1.5
(
0.883)
­
1.8
(
0.27)
NG
NG
NG
NG
54.7
NG
NG
0.0681
NG
NG
­
3.6
NG
NG
NG
54
NG
0.075
0.041
NG
­
5.9
­
9.8
NG
NG
NG
NG
54
68
0.075
0.041
0.032
­
5.9
­
9.8
­
8.5
NG
NG
NG
60
55
0.12
0.09
0.12
­
0.9
­
1.3
­
0.9
NG
NG
NG
55
to
60
NG
0.11
to
0.12
0.09
to
0.12
NG
­
1.5
to
­
0.9
­
1.3
to
­
0.67
NG
NG
NG
NG
51.3;
53.4
NG
0.24;
0.22
0.19;
0.2
NG
NG
NG
NG
NG
NG
NG
NG
64.2
NG
NG
0.1205
NG
NG
­
1.64
NG
NG
NG
NG
68
NG
NG
0.076
NG
NG
­
4.3
NG
NG
NG
75
(
3.592)
61
(
1.8)
0.13
(
0.0208)
0.08
(
0.0866)
0.14
(
0.0101)
­
1
(
0.5167)
­
2.9
(
3.492)
0.65
(
0.131)
NG
NG
NG
52.93
NG
0.132
0.132
NG
­
4.32
­
3.22
NG
NG
NG
NG
53.57
NG
0.143
0.176
NG
­
5.04
­
2.47
NG
NG
NG
NG
NG
68
NG
NG
0.76
NG
NG
­
4.3
NG
NG
NG
50.6
(
2.7)
NA
0.2373
(
0.0396)
0.2595
(
0.0724)
NG
0.139
(
0.5813)
0.769
(
0.8312)
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
NG
52.1
NG
0.177
0.183
NG
­
2.14
­
1.2
NG
NG
NG
NG
0.23
&
0.22
(
C.
I.=
0.20­
0.26
&
0.19­
0.25)
0.17
&
0.18
(
C.
I.=
0.14­
0.19
&
0.15­
0.2)
­
0.74
&
­
0.56
(
C.
I.=
­

0.99
to
­
0.49
&
­
0.77
to
­
0.35)

NG
­
0.46
&
­
0.44
(
C.
I.=
­

0.65
to
­
0.28
&
­
0.62
to
­
0.26)
NG
NG
NG
NG
57.8
&
58.78
(
C.
I.=
55.3­
60.4
&

56.4­
61.1)
NG
Page
919
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
V
W
X
Y
otoliths;
whole
No
No
NF
otoliths;
b
&
b
No
Interreader
comparisons
otoliths;
whole
No
Intrareader
comparisons
NF
otoliths;
whole
No
No
6
otoliths;
b
&
b
No
No
otoliths;
whole
No
No
otoliths;
whole
No
Intrareader
comparisons
otoliths;
whole
No
No
otoliths;
whole
NG
NG
otoliths;
whole
No
No
7
(
young
of
the
year)

otoliths;
b
&
b
No
No
10,15
(
juveniles)

otoliths;
b
&
b
No
NG
otoliths;
whole
No
Intrareader
comparisons
otoliths;
whole
No
No
otoliths;
whole
No
No
otoliths;
break
&
burn
No
Intrareader
comparisons
otoliths;
break
&
burn
No
Intrareader
comparisons
otoliths:
whole
NG
NG
otoliths;
whole
No
Intrareader
comparisons
otoliths;
whole/
section
OTC
NG
otoliths;
?
OTC
interreader/
departmental
comparison
otoliths;
section/
whole
No
Multiple
structure
comparison
Page
920
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
Z
AA
AB
AC
AD
AE
AF
7
64
7
Dioecious
Polygynous2
Viviparous3
No
NA
21
11
primitive
viviparity11
10,21,35,36
19
NF
Dioecious
Polygynous2
Viviparous3
No
1
0.7
10,35
>
32
2,3,6,7,9,11,13,18
Dioecious
Polygynous2
Viviparous3,7
No
1
0.99
15
55
Ovoviviparous10
21
15
29
38
33,34
76
18
>
20
2,3,6,7,8,9,18
Dioecious
Polygynous2
Viviparous3,7,11
No
1
0.91
1648
(
Males);
35
(
Females)
(
matrotrophic)
3,5,6
21,23
21
Ovoviviparous10
26,32
42
(
Males);
34
(
Females)

35
36
Page
921
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
AG
AH
AI
AJ
AK
AL
AM
32NG
(
NG)
NG
(
NG)
23
(
7)­
sex
unspecified
NG
NG
42<
22*
(
NG)
<
22
(
NG)
<
22
(
NG)
<
22
(
NG)
NA
NA
53
NG
(
NG)
NG
(
NG)
15.2
to
25.4
(
4
to
10)
15.2
to
25.4
(
4
to
10)
NG
NG
None1
3
22
(
7)
20
(
5)
22
(
7)
23
(
8)
otoliths;
whole
No
621*(
NG)
19
(
NG)
22.8
(
13)
20.8
(
10)
otoliths;
b
&
b
No
11NG*(
NG)
NG
(
NG)
21
(
NG)
21
(
NG)
NG
NG
11NG*(
NG)
NG
(
NG)
22
(
NG)
20
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
23
(
8)­
sex
unspecified
NG
NG
42
19*
(
NG)
18
(
NG)
21
(
NG)
19
(
NG)
NA
NA
53
NG
(
NG)
NG
(
NG)
22.9
to
27.9
(
7
to
10)
22.9
to
27.9
(
7
to
10)
NG
NG
None1
315
(
4)
17
(
5)
16
(
5)
18
(
5)
otoliths;
whole
No
513
(
NG)
14
(
NG)
13
(
NG)
14
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
18
(
5)­
sex
unspecified
NG
NG
53
NG
(
NG)
NG
(
NG)
15
(
4)
15
(
2)
NG
NG
None1
3,52
22
(
4)
26
(
6)
22
(
4)
26
(
6)
otoliths;
whole
No
21NG*(
NG)
NG
(
NG)
31
(
12)
32
(
14)
otoliths;
b
&
b
No
15,41NG
(
NG)
NG
(
2)
NG
(
NG)
NG
(
4)
otoltihs;
whole
No
18NG
(
6)
NG
(
5)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
25NG
(
NG)
NG
(
NG)
29.5
(
13)
28.1
(
11)
otoliths;
b
&
b
NG
32NG
(
NG)
NG
(
NG)
26
(
6)­
sex
unspecified
NG
NG
19
NG
(
NG)
NG
(
NG)
NG
(
NG)
29.3
(
11)
NG
NG
3,53
25
(
3)
30
(
5)
36
(
6)
41
(
7)
otoliths;
whole
No
21NG*
(
NG)
NG
(
NG)
40
to
42
(
10
to
13)
38
to
42
(
9
to
12)
otoliths;
b
&
b
No
12NG
(
4)
NG
(
4)
NG
(
5)
NG
(
6)
otoliths;
whole
No
None1
18NG
(
3)
NG
(
3)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
32NG
(
NG)
NG
(
NG)
41
(
7)­
sex
unspecified
NG
NG
37NG
(
NG)
NG
(
NG)
40
(
7)
35
(
6)
NG
NG
39
30
(
4)
30
(
4)
38.3
(
6.3)
42.2
(
7.9)
otoliths;?
OTC
53
NG
(
NG)
NG
(
NG)
45.7
(
9
to
13)
45.7
(
9
to
13)

Longer
anal
spine,
upper
jaw,
pectoral
fin
ray
and
orbit
width
per
SL
in
males3
Page
922
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
AN
AO
AP
AQ
AR
AS
AT
AU
AV
NG
c.
California4,45
*****

NA
s.
California13,76,77
*****

NG
California57
*****
*****
*****
*****

No
NF
Gulf
of
Alaska15
*****

interreader
comparisons
Alaska67,71,77
NG
British
Columbia15,70
*****

NG
Oregon16,19,66,70
*****
*****

NG
n.­
c.
California5,77
NA
California57
*****
*****

NG
No
NF
n.­
c.
California5,77
^^^^^
*****

NG
s.
California13,77
*****
*****

NG
California57
*****

NG
No
2,3
Gulf
of
Alaska20
No
Alaska67,71
No
British
Columbia15
*****

No
Washington7,12,42,53
*****

NG
Oregon66
NG
n.­
c.
California5
^^^^/****

NG
California57
*****
*****

No
2
Gulf
of
Alaska21
No
Alaska67,71
intrareader
comparisons
B.
C.
21,22,35
No
Oregon21,39,40
NG
n.­
c.
California5
^^^^^
*****

NG
c.
California4,51
interreader/
departmental
comparisons
California57,62,77
*****
*****
*****
*****

Page
923
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
AW
AX
AY
AZ
BA
BB
BC
BD
BE
*****
Once5
NF
NF
Multiple
2
3187
(
NA)
1,27
8,686
(
16.9
TL)
to
39,394
(
23.8
TL)

4
35,000
(
NG)

Mulitple?
14
NF
NF
Once7
NF
25
117,550
(
37
?
L)
to
1,200,000
(
49
?
L)

25
544,528
(
41)

Some
fraction
of
the
population
seem
to
not
spawn
annually20
Page
924
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
BF
BG
BH
BI
BJ
NF
NA
1
~
1
to
2
months
NF
NF
3F=
0.004219TL2.7994
(
r2=
0.599)
NA
1
~
1
to
2
months
NF
NF
3<
115
days
NA
1
~
1
to
2
months
NF
NF
12
F=
64,098?
L­
2,083,531
(
r2=
0.78)
NA
1
~
1
to
2
months
143
to
63
NF
6,10,1840
to
50
7
52.5
935
to
40
1530
to
35
1730
to
36
See
Richards
&
Schnute
(
1990)
for
additional
discussion
11
3120
to
180
days;

likely
110
days
Page
925
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
BK
BL
BM
BN
BO
BP
NF
No
external
eggs
NA
2,184.1
to
19.8
(
SL)
planktonic1
planktivore
204.1
(
TL)
to
NF
No
external
eggs
NA
254.7
(
SL)
to
planktonic1
planktivore
NF
No
external
eggs
NA
20,215.0
(
TL)
to
planktonic1
planktivore
235.4
(
SL)
to
6
0.83
No
external
eggs
NA
12,17,204.0
to
~
24
(
SL)
planktonic1,2
planktivore
15,245.5
to
20
(
SL)

233.8
(
SL)
to
263.6
(
SL)
to
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
Page
926
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
BQ
BR
BS
BT
BU
BV
NF
diurnal1
siphonophores;
19.8
(
SL)
to
maturity1
pelagic?;
NF3,9,22
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
pelagic;
NF3,9,46
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
NF
diurnal1
siphonophores;
33.1
(
SL)
to
maturity4
zooplanktivore
chaetognaths1
20
(
SL)
to
maturity5
24
(
SL)
to
maturity7
(
crustaceans,
such
calanoids,
mysids,
and
barnacle
cyprids;
fish
larvae;
juvenile
polychaetes)
1,6,8,10
pelagic;
rock
and
cobble;
rocky
shelf;

sponge
gardens,
such
as
boot
sponge;

high
relief
rock;
low­
relief
artificial
reefs
4,9,12,14,27,29,30,33,35,37,40,49
(
yp
shrimp
and
calanoid
copepods;
pelagic
fishes)
1,6,16,17
pelagic;
nearshore
sand­
rock
interface;

seagrass
beds;
kelp
canopy;
midwater;

high
rock;
artificial
reefs;
bays
and
estuaries;
tide
pools
2,3,4,9,10,13,15,17,24,29,30,34,44,50,51,54
Page
927
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
BW
BX
BY
BZ
CA
NF
NF
carnivore
(
crustaceans;
cephalopods)
20,48,76
NF
NF
planktivore
(
crustaceans)
16,33,48,76
NF
NF
carnivore
eelgrass
beds
at
night47
NF
diurnal4,5
carnivore
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise
1,2,3,5,6,8,9,12,14,16,19,22
(
fish,
including
juvenile
rockfishes
like
S.
mystinus
;

crustaceans;
polychaetes;
cephalopods;

chaetognaths;
jellyfish)
10,12,17,21,23,24,28,30,41,48,54,61,76
(
crustaceans;
small
fish,
including
rockfishes
anf
flatfishes;
bivalves;
polychaetes;
fish
eggs,
such
as
Ophiodon
elongatus's)
12,17,24,30,39,42,48,76
fishes,
including
rockfishes
(
such
as
yelloweye),
lingcod,

cabezon
and
salmon;
birds;

porpoise1,2,3,5,6,8,10,12
deeper
rock/
rocky
shelf;
high
relief
rock
and
artificial
reefs;
boulder
fields
5,12,19,23,34,35,36,39,41,43,52,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
rocky
shelf;
continental
slope/
basin;

boulder/
cobble­
mud
5,7,12,21,25,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
rocky
shelf;
high
relief;
mixed
bottoms
5,7,18,21,65
high­
relief
rocky
reefs;
in
and
around
kelp
beds;
boulder
fields;
midwater;
pelagic;

artificial
reefs
3,5,7,14,20,23,28,37,43,45,50,54,57,65
Page
928
of
1167
5­
2220e
­
Scorpaenidae
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
CB
CC
CD
CE
NF
NF
yelloweye
rockfish3
sharks;
dolphin;
seals1,4
possibly
river
otters12
NF
NF
NF
NF
dawn
&
dusk2
Puget
Sound;
Alaska
to
California7
diurnal5,10,14
possibly
river
otters12
NF
dusk2
diurnal5
dawn
&
dusk7,10
possibly
river
otters12
sharks;
dolphin;
seals;
salmon1,4,10
yelloweye
rockfish;
lingcod;

sharks;
salmon;
dolphin;
seals;

sea
lions;
pigeon
guillemots
1,2,4,5,6,7,10,11,17
Washington;
Oregon
(
based
on
genetics)
2
Oregon
to
northern
Washington;
Oregon
to
southern
California
(
based
on
tag
&
recapture)
2,13
B.
C.
to
alaska;
Washington
to
California
4
yelloweye
rockfish;
sharks;

dolphin;
seals1,3,4
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Scorpaenidae
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110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
CF
CG
CH
CI
CJ
CK
CL
CM
194.49
±
1.57
(
Alaska
to
CA)
2
2.05
±
1.04
11.3
(
0.1)
20.8
0.108
(
0.049)
0.111
(
0.051)
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1
1.21
11.4
(
0.1)
20
0.092
(
0.036)
0.046
(
0.02)
allozymes
NF
30.046
to
0.074
NF
NF
11.2
(
0.1)
20.8
0.046
(
0.024)
0.034
(
0.017)
allozymes
NF
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1167
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Scorpaenidae
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103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
CN
CO
CP
CQ
CR
CS
717
(
6)
17
(
6)
trawls
otoliths;
break­
and­
burn
No
intereader
comparisons
NF
NF
NF
NF
NF
NF
324.1
(
6)
29
(
9)
hook
&
line;
spearfishing
otoliths;
whole
No
No
15NG
(
3
to
4)
NG
(
8
to
9)
hook
&
line
otoliths;
whole
NG
NG
323.2
(
3)
33.7
(
5)
hook
&
line;
spearfishing
otoliths;
whole
No
No
1
30.5
(
4)
40
(
7)
hook
&
line
otoliths;
whole
No
intrareader
comparisons
2NG
(
NG)
45
to
55
(
9
to
11)
trawl
otoliths;
section/
whole
No
multiple
structure
comparison
10NG
(
4
to
6)
NG
(
NG)
jig
gear
otoliths;
break­
and­
burn
No
NG
11NG
(
6)
NG
(
NG)
NG
NG
NG
NG
12,13NG
(
4)
NG
(
4)
NG
NG
NG
NG
15NG
(
3
to
4)
NG
(
8
to
9)
hook
&
line
otoliths;
whole
NG
NG
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931
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Scorpaenidae
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111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
CT
CU
CV
CW
9
0.07­
Males;
0.07­
Females
0.28­
Males;
0.32­
Females
0.35­
Males;
0.39­
Females
Hoenig
estimation/
Z­
formula
9
0.04­
Males;
0.05­
Females
0.31­
Males;
0.34­
Females
0.35­
Males;
0.39­
Females
catch
curve/
Z­
formula
90.04­
Females
0.35­
Females
0.39­
Females
GSI/
Z­
formula
NF
NF
NF
NF
1
0.1253
0.0258
0.1511
tag
&
recapture
4NG
NG
0.49644
Hencke
survivorship/
ration
13NG
NG
0.06
to
0.09
NG
NG
NG
0.21
NG
21
0.115
0.112
0.227
Hencke
survivorship/
ration/
age
frequency
1
0.3081
0.02
0.3281
Chapman­
Robson/
tag
&
recap.

2NG
NG
0.276
catch
curve
3,7NG
NG
0.51
to
0.6
Chapman­
Robson
3,7NG
NG
0.4
to
0.54
catch
curve
4NG
NG
0.51395
Hencke
survivorship/
ration
10
0.2
to
0.4
NG
NG
catch
curve
11
0.409­
Males;
0.336
to
0.704­
Females
0.283­
Both
0.692­
Males;
0.619
to
0.987­
Females
tag/
recapture
14NG
NG
0.28
catch
curve
16
0.28
NG
NG
Reparam.
B­
H
Stock­
Recruitment
20
0.1­
Males;
0.12­
Females
NG
NG
Hoenig's
(
1983)
equation
20
0.19­
Males;
0.26­
Females
NG
NG
Beverton
&
Holt
(
1959)

20
0.17­
Males;
0.24­
Females
NG
NG
catch
curve
21
0.249
0.093
0.342
Hencke
survivorship/
ration/
age
frequency
Page
932
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1167
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Scorpaenidae
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106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
CX
CY
CZ
DA
6
to
64
NG
0.8­
Males;
0.82­
Females
NF
6
to
64
NG
0.89­
Males;
0.87­
Females
6
to
64
NG
0.9­
Females
NF
NF
NF
NF
6
to
37
0.0239
0.171
NF
9
to
16
NG
NG
NG
NG
NG
NG
NG
NG
9
to
16
0.156
0.493
3
to
14
0.0171
0.061
NF
7
to
14
NG
NG
12
to
17
NG
NG
12
to
17
NG
NG
4
to
12
NG
NG
NG
NG
NG
6
to
16
(
both
sexes)
NG
0.408­
Males;
0.287
to
0.457­
Females
15
to
25
NG
NG
NG
NG
NG
max.
age:
42
for
males;
34
for
females
NG
NG
NG
NG
NG
9
to
16
NG
NG
4
to
12
0.079
0.272
Page
933
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Scorpaenidae
102
103
104
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106
107
108
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110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
DB
DC
DD
DE
NF
NF
NF
infected
by
parasites
from
the
genus
Anisakidae
22
NF
NF
NF
5<
5
nm
Residential1,12
Yes4,5,8
72.8
km
associated
with
S.
caurinus
8
no
movement6
5<
5
nm
Transient6,7,19
No9
7320
km
Transient/
Residential12,13,18
parasitized
by
copepod
Clavellopsis
robusta
11
no
movement6
Mostly
residential22
14generally
<
10
mi
substantial
movement17
18up
to
22
km
22
up
to
345
miles
infected
by
parasites
from
the
families
Echinorhynchidae,

Polymorphidae,
Caligidae,
Chondracanthidae,
Hemiuridae,

Lepocreadiidae,
Opecoelidae,
Sanguinicolidae,

Microcotylidae,
and
Anisakidae22
may
contain
trematode
Derogenes
varicus
and
stomach
parasite
Lecithochirium
exodicum
6
infected
by
parasites
from
the
families
Polymorphidae,

Phyllobothriidae,
Tentaculariidae,
Bomolochidae,
Caligidae,

Chondracanthidae,
Lernaeopodidae,
Naobranchiidae,

Philichthyidae,
Acanthocolpidae,
Hemiuridae,

Lepocreadiidae,
Opecoelidae,
Sanguinicolidae,
Zoogonidae,

Aegiidae,
Capsalidae,
Microcotylidae,
Anisakidae,

Capillariidae,
Cucullanidae,
Cystidicolidae,
and
Sinuolineidae22
infected
by
parasites
from
the
families
Chondracanthidae,

Lernaeopodidae,
Bucephalidae,
Hemiuridae,

Lepocreadiidae,
Sanguinicolidae,
Piscicolidae,
Capsalidae,

Microcotylidae,
and
Myxobolidae22
Schooling
with
S.
mystinus
and
S.
serranoides
;
benthically
with
S.
chrysomelas
4,13
19most
no
movement,
but
some
>
5
km;
up
to
178
km
migrates
to
deeper
water
in
winter16
migrates
to
deeper
water
in
winter16
9,15
up
to
619
km,
but
mostly
no
movement
infected
by
parasites
from
the
families
Polymorphidae,

Phyllobothriidae,
Tentaculariidae,
Bomolochidae,

Lernaeopodidae,
Philichthyidae,
Bucephalidae,

Fellodistomatidae,
Hemiuridae,
Lepocreadiidae,

Opecoelidae,
Zoogonidae,
Capsalidae,
Microcotylidae,

Anisakidae,
Cystidicolidae,
and
Tetracapsulidae22
parasitized
by
Prosorhynchus
;
usually
heavily
parasitized15
infected
by
macroparasites
from
the
families
Microcotylidae,

Bothriocephalidae,
and
Anisakide14,20
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934
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Scorpaenidae
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111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
DF
DG
NF
NF
Yes15,23
NF
No7,23
No12
Yes19
Yes
5,7,9,11,20,21,22,24,26,29,35
No10
No8
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935
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Scorpaenidae
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153
154
155
156
157
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160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
A
B
C
D
E
S.
miniatus
1
1
<
1
to
436
S.
mystinus
1
1
<
1
to
549
S.
nebulosus
1
1
3
to
128
S.
nigrocinctus
1
1
18
to
298
S.
ovalis
1
1
30
to
366
china
rockfish;
china
cod;
blacks;
yellow
stripe;
cefalutano;
cerod;
pelican;
yellow
rockfish;
speckled
garrupa;
chinafish;

gopher1,3,4,5,6,9,10,12,13
vermillion
rockfish;
red
snapper;
red
rockfish;
red
rock
cod;
red;
Pacific
red
snapper;
bloodfish;
barracho(
n);
racha;

rasher;
genuine
red;
keeper1,3,4,5,6,7,12,13
blue
rockfish;
blue
bass;
blue­
fish;
reef
perch;
black
bass;
nervi;
neri;
black
snapper;
priestfish1,3,4,5,6,9,12,13
tiger
rockfish;
banded
rockfish;

blackbanded
rockfish;
barred
rockfish1,3,7,9
speckled
rockfish;
cinnamon;
bass;

bellina
cod;
J.
W.;
bank
perch;
zipola;

beccafico;
brownfish;
viuva1,3,4,5,6,7,9,12,13
Alaska
(
Zaikof
Bay,
Montague
I.,
Prince
William
Sound)
to
central
Baja
California
(
Islas
San
Bonito)
1,8
eastern
Gulf
of
alaska
(
Sitka)
to
northern
Baja
California
(
Punta
Baja)
1,3,5
northern
Gulf
of
Alaska
(
Kachemak
Bay)
to
southern
CA
(
Redondo
Beach
and
San
Miguel
I.)
1
northern
Gulf
of
Alaska
(
The
Triplets)
to
southern
CA
(
Tanner­
Cortes
Banks)
1
northern
WA
to
northern
Baja
California
(
Cabo
Colonet)
1
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Scorpaenidae
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161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
F
G
H
I
J
26Males:
W=
0.038?
L2.8
(
r2>
0.9)
c.
California
(
TL)
11
55.9
26Females:
W=
0.018?
L2.98
(
r2>
0.9)

28Male:
W=
0.0000000115TL3.05
(
r2=
0.955)

28Female:
W=
0.000000013TL3.03
(
r2=
0.98)

28Both:
W=
0.0000000819TL3.11
(
r2=
0.978)

1Both:
W=
0.02157TL2.92339
(
r2=
0.967)
4TL=
9.629
+
1.229(
SL);
r2=
0.988;
SE=
9.8
1
63
c.
California
(
TL)
11
57.5
7*
Both:
W=
0.000000059TL2.9165
SE=
0.302
4TL=
­
7.857
+
1.054(
FL);
r2=
0.998;
SE=
4.6
c.
to
n.
California
(
TL)
3
NG
8*
Male:
W=
0.00001971TL2.985
(
r2=
0.994)
4SL=
­
3.095
+
0.804(
TL);
r2=
0.988;
SE=
7.9
California
(
SL)
14
NG
8*
Female:
W=
0.00001967TL2.992
(
r2=
0.992)
4SL=
­
9.445
+
0.847(
FL);
r2=
0.988;
SE=
7.8
8*
Both:
W=
0.00001458TL3.041
(
r2=
0.996)
4FL=
8.665
+
0.946(
TL);
r2=
0.998;
SE=
4.4
4FL=
16.44
+
1.168(
SL);
r2=
0.994;
SE=
9.2
6*
Male:
W
(
in
lbs)=
0.000000033TL2.99
(
SE=
0.05)
4TL=
2.93
+
1.238(
SL);
r2=
0.996;
SE=
5.7
1
53
Oregon
(
FL)
16
48.4
(
2.2)

6*
Female:
W
(
in
lbs)
=
0.000000102TL2.81
(
SE=
0.2)
4TL=
2.495
+
1.039(
FL);
r2=
1.0;
SE=
2.3
California
(
TL)
31
NG
*
Both6:
W
(
in
lbs.)=
0.00000048TL2.54
(
SE=
0.2)
4SL=
­
1.192
+
0.804(
TL);
r2=
0.996;
SE=
4.6
c.
to
n.
California
(
FL)
39
:
from
deep
habitats
33.17
8*
Male:
W=
0.00002934TL2.889
(
r2=
0.925)
4SL=
0.644
+
0.836(
FL);
r2=
0.996;
SE=
4.2
c.
to
n.
California
(
FL)
39
:
from
shallow
habitats
32.31
8*
Female:
W=
0.00003408TL2.874
(
r2=
0.914)
4FL=
­
2.164
+
0.962(
TL);
r2=
1.0;
SE=
2.2
c.
to
n.
California
(
FL)
39
:
from
both
habitats
32.94
8*
Both:
W=
0.000009774TL3.09
(
r2=
0.978)
4FL=
0.352
+
1.192(
SL);
r2=
0.998;
SE=
5.0
s.
California
(
SL)
27
NG
12*
Male:
W
(
in
kg)=
0.0000533FL2.709
3SL=
0.8127(
TL)

12*
Female:
W
(
in
kg)=
0.0000943FL2.596
3SL=
0.8506(
FL)

12*
Both:
W
(
in
kg)=
0.0000494FL2.752
20Both:
W=
0.0000204(
SL)
3.059
8*
Male:
W=
0.000008793TL3.153
(
r2=
0.941)
4TL=
4.294
+
1.196(
SL);
r2=
0.99;
SE=
4.0
1
43
c.
California
(
TL)
11
37.5
8*
Female:
W=
0.000006644TL3.206
(
r2=
0.976)
4TL=
0.854
+
1.001(
FL);
r2=
1.0;
SE=
1.4
8*
Both:
W=
0.000007789TL3.177
(
r2=
0.982)
4SL=
­
0.731
+
0.828(
TL);
r2=
0.99;
SE=
3.3
4SL=
­
1.852
+
0.835(
FL);
r2=
0.986;
SE=
3.9
4FL=
­
0.487
+
0.998(
TL);
r2=
1.0;
SE=
1.4
4FL=
6.934
+
1.181(
SL);
r2=
0.986;
SE=
4.6
*
Both:
W=
0.00000809TL3.147
NF
1
61
NF
NF
1Male:
W=
0.00621TL3.21742
(
r2=
0.978)
4TL=
0.55
+
1.225(
SL);
r2=
0.994;
SE=
4.4
1
56
s.
California
(
TL)
10
35.86
(
2.6)

1Female:
W=
0.00643TL3.1371
(
r2=
0.973)
4TL=
9.914
+
1.033(
FL);
r2=
0.998;
SE=
2.0
4SL=
1.329
+
0.811(
TL);
r2=
0.994;
SE=
3.6
4SL=
5.13
+
0.836(
FL);
r2=
0.992;
SE=
3.9
4FL=
­
3.311
+
0.967(
TL);
r2=
0.998;
SE=
1.9
4FL=
­
3.554
+
1.187(
SL);
r2=
0.992;
SE=
4.7
Page
937
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­
Scorpaenidae
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166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
K
L
M
N
O
P
Q
R
S
T
U
66.9
69.1
0.1373
0.1119
0.0953
­
1.4
­
1.1
­
1.5
NG
NG
NG
62.4
59.8
0.2044
0.1396
0.1595
­
0.2
­
1.2
­
0.9
NG
NG
NG
NG
68.8
(
11.4)
NG
NG
0.09841
(
0.0031)
NG
NG
­
0.69
(
0.0494)
NG
NG
NG
NG
56.4
NG
NG
0.09841
NG
NG
­
0.69
NG
NG
NG
47.2
(
1.2)
NG
0.2296
(
0.0391)
0.3072
(
0.0387)
NG
­
0.0301
(
0.3876)
0.5239
(
0.2592)
NG
NG
NG
NG
NG
42.7
NG
NG
NG
NG
NG
NG
NG
NG
NG
43.07
NG
0.195
0.107
NG
­
0.94
­
1.94
NG
NG
NG
NG
39.33
NG
0.194
0.166
NG
­
0.99
­
1.14
NG
NG
NG
NG
40.02
NG
0.195
0.149
NG
­
0.95
­
1.34
NG
NG
NG
NG
NG
32.8;
33.6
NG
NG
0.143;
0.132
NG
NG
­
1.55;
­
1.74
NG
NG
NG
37.3
38.2
0.194
0.1924
0.1806
­
0.2
0.2
0.1
NG
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
49.99
(
2.08)
NG
0.06
(
0.018)
0.053
(
0.007)
NG
­
10.247
(
3.16)
­
4.936
(
1.031)
NG
NG
NG
NG
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938
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Scorpaenidae
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153
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161
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163
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165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
V
W
X
Y
otoliths;
whole
No
Interreader
comparisons
otoliths;
whole
No
Interreader
comparisons;
comparisons
with
operculum
NF
scales
No
No
otoliths;
whole
No
No
otoliths;
whole
No
Intrareader
comparisons
5,12,13
length
frequency
analysis/
Ford­
Walford
plot
No
No
16­
juvenile
and
adult
otoliths;
break
and
burn
Marginal
increment
analysis
Intrareader
comparisons
otoliths;
break
and
burn
Marginal
increment
analysis
Intrareader
comparisons
otoliths;
break
and
burn
Marginal
increment
analysis
Intrareader
comparisons
scales
No
Intrascale
comparison
otoliths;
whole
No
Interreader
comparisons
NF
NF
NF
NF
NF
otoliths;
section
No
Intrareader
comparison
NF
Page
939
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1167
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Scorpaenidae
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166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
Z
AA
AB
AC
AD
AE
AF
3>
20
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
0.94
10,11,25,32
25
Ovoviviparous10
21
30
33
43
35
22
2,33
17
(
Males);
24
(
Females)
4,14
Dioecious
Polygynous2
Viviparous3,7
No
1
0.86
21,35
24
Ovoviviparous10
24,32
23
primitive
viviparity11
2844
(
Males);
41
(
Females)

21
26
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
1.05
Ovoviviparous10
NF
NF
Dioecious
Polygynous2
Viviparous3,7
No
NF
Ovoviviparous10
primitive
viviparity11
10,35>
37
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
0.8
21,36
37
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Scorpaenidae
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166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
AG
AH
AI
AJ
AK
AL
AM
None1
3,52
35
(
5)
37
(
5)
38
(
5)
37
(
5)
otoliths;
whole
No
532
(
NG)
31
(
NG)
35
(
NG)
37
(
NG)
NG
NG
236.5
(
NG)
36.5
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
14NG
(
NG)
22.9
(
3)
33
(
6)
33
(
6)
scales
No
32NG
(
NG)
NG
(
NG)
37
(
5)­
sex
unspecified
NG
NG
50
31.8
(
5
to
6)­
sex
unspecified
38.1
(
8
to
9)­
sex
unspecified
NG
NG
53
NG
(
NG)
NG
(
NG)
35.6
(
5
to
6)
35.6
(
5
to
6)
NG
NG
3,52
22
(
4)
22
(
5)
27
(
5)
29
(
6)
otoliths;
whole
No
221.
9
(
NG)
19.6
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
719
(
3)
20
(
3)
26
(
7)
27
(
5.5)
scales
No
None1
12NG*
(
3)
NG
(
2)
NG
(
3)
NG
(
4)
otoliths;
whole
No
1323
(
3)
22
(
5)
6
(
26.2)
28.2
(
6)
scales
No
20,52
NG
(
3)
NG
(
5)
NG
(
4)
NG
(
NG)
scales
No
32NG
(
NG)
NG
(
NG)
29
(
6)­
sex
unspecified
NG
NG
38,53
NG
(
3)
NG
(
3)
26
(
5)
28
(
6)
NG
NG
4720.3
to
22.9
(
4)
25.4
(
5)
NG
(
NG)
NG
(
NG)
NG
NG
54NG
(
4
to
5)
NG
(
3)
54NG
(
5
to
6)
NG
(
5)
otoliths;
break
and
burn
Marginal
increment
analysis
None1
3,53
26
(
3)
28
(
4)
27
(
4)
28
(
4)
otoliths;
whole
No
234.2
(
NG)
26.2
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
27
(
4
to
7)­
sex
unspecified
NG
NG
None1
NF
NF
NF
NF
NF
NF
None1
328
(
4)
28
(
4)
28
(
4)
28
(
4)
otoliths;
whole
No
523
(
NG)
24
(
NG)
24
(
NG)
25
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
28
(
4
to
10)­
sex
unspecified
NG
NG
longer
anal
and
dorsal
ray
in
females;
longer
orbit
per
SL
width
in
males3
Page
941
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1167
5­
2220e
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Scorpaenidae
152
153
154
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156
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158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
AN
AO
AP
AQ
AR
AS
AT
AU
AV
No
1
c.­
n.
California5,17,28,64,77
NG
c.
California4,51
NG
s.
California13,18,64,76,77
No
California27,57,70,74
*****
*****
*****
*****
*****

NG
NG
NG
No
NF
c.­
n.
California5,23,24,28,32
^^^^^
*****

NG
c.
California4,23,33,34,51,76
*****
*****

compared
to
otoliths
s.
California25
intrareader
comparisons
California27,35,57,63,70,72,77
*****

intrareader
comparisons
No
NG
NG
NG
intrareader
comparisons
No
NF
Alaska44,67,71,77
*****

NG
c.­
n.
California5,26
^^^^/****

NG
c.
California4
California57,70
*****
*****
*****
*****

NF
NF
Alaska44,67,77
Oregon66
British
Columbia15
*****

Washington12,70
No
NF
c.­
n.
California5
NG
s.
California13,18,77
*****
*****

NG
California57
*****
*****
*****
*****

Page
942
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1167
5­
2220e
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Scorpaenidae
152
153
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156
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160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
AW
AX
AY
AZ
BA
BB
BC
BD
BE
*****
Once2,22
1347
(
122­
618)
1,26,27
158,915
(
46.0
TL)
to
2,683,768
(
68.0
TL)

Multiple10
3
182
(
NA)
5,2463,000
(
31.8
TL)
to
1,6000,000
(
54.6
TL)

*****
3,5
1,100,000
(
up
to
5,600,000)
26
3,300
(
31.5
SL)
to
1,625,600
(
55.0
SL)

*****
*****
*****
Once3,8,22
NF
7,13,26,27
50,000
(
25
TL)
to
300000
(
32.5
TL)

Multiple16,22
8,953,300
(
25.6
TL)
to
233,700
(
33
TL)

12524,000
(
40.5
TL)

Once17
NF
NF
NF
NF
NF
Multiple
1,2
1116
(
88­
152)
3,27
61,217
(
33.1
TL)
to
160,320
(
39.3
TL)

Page
943
of
1167
5­
2220e
­
Scorpaenidae
152
153
154
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156
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160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
BF
BG
BH
BI
BJ
3F=
0.0000012TL5.0226
(
r2=
0.877)
NA
1
~
1
to
2
months
925
to
30
NF
4F=
a
TL5.686
NF
NA
1
~
1
to
2
months
1,2042
to
64
NF
380
to
130
days
9,1530
to
35
4
129
19
76.2
NF
NA
1
~
1
to
2
months
NF
NF
NF
NA
1
~
1
to
2
months
NF
NF
NF
NA
1
~
1
to
2
months
NF
NF
Page
944
of
1167
5­
2220e
­
Scorpaenidae
152
153
154
155
156
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158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
BK
BL
BM
BN
BO
BP
NF
No
external
eggs
NA
204.3
(
SL)
to
planktonic1
planktivore
5
0.12
to
0.16
No
external
eggs
NA
7,17,20,263.8
(
NL)
to
21
(
SL)
planktonic1,3,8
planktivore
10,160.3
to
0.35
23,34
3.8
(
SL)
to
27,33,34
3.5
to
NF
No
external
eggs
NA
235.7
(
SL)
to
planktonic1
planktivore
NF
No
external
eggs
NA
23~
5.0
(
SL)
to
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
5
0.66
to
0.96
No
external
eggs
NA
17,185.0
(
SL)
to
planktonic1
planktivore
254.7
(
SL)
to
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
Page
945
of
1167
5­
2220e
­
Scorpaenidae
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153
154
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156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
BQ
BR
BS
BT
BU
BV
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6,24
NF
diurnal1
siphonophores;
7>
21
(
SL)
to
maturity
zooplanktivore
chaetognaths1
in
creviecs
at
night
47
NF
diurnal1
siphonophores;
NA
pelagic;
NF3,9
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
pelagic;
shallow,
rocky
reefs3,9,20
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
pelagic;
rocky
shelf
4,9
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
pelagic;
nearshore
sand­
rock
interface;

rocky
shelf;
demersal
2,3,4,9,13,24,53
pelagic;
nearshore
sand­
rock
interface;

kelp
canopy;
rocky
shelf;
high
rock;

artificial
reefs
2,3,4,9,13,24,26,28,31,38,47,50,52,53,54
(
larvaceans;
crustaceans,
such
as
harpacticoids
and
barnacle
cyprids;
hydroids;

jellyfish;
polychaetes;
tunicates)
1,2,3,5,6,8,10,23
Page
946
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2220e
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Scorpaenidae
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161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
BW
BX
BY
BZ
CA
NF
NF
planktivore/
carnivore
NF
diurnal2,4,5,10
omnivore/
zooplanktivore62,74
NF
NF
rock
and
cobble;
rocky
shelf
5,7,12,23,65
carnivore
(
ophiuroids;
crustaceans;
fish;
mollusks)
12,23,30,48,60,76
NF
NF
carnivore
(
crustaceans;
fish,
including
juvenile
rockfish)
12,30,76
NF
NF
rocky
shelf;
continental
slope/
planktivore/
carnivore
basin;
high
relief
5,18,65
(
zooplankton;
small
fish)
33,48,76
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12,14,16,23,27
olive,
black,
kelp,
and
gopher,

and
black­&­
yellow
rockfish24,25
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
rocky
substrates
of
shelf
and
continental
slope/
basin;
high
relief;
within
crevices
5,7,12,23,35,39,52,65
within
and
around
the
kelp
canopy;
high
relief
rocky
and
artificial
reefs
5,7,14,17,24,27,28,32,37,45,49,50,51,64,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12,18
rocky
shelf
and
boulder
fields;
in
crevices;

continental
slope/
basin;
sandy/
soft
bottoms;

kelp
forest
bottom
5,7,17,24,27,37,55,61,62,64,65
(
cephalopods;
small
fish,
such
as
surfperch;

euphasiids;
polychaetes;
pyrosomes)

6,9,23,37,48,56,64,70,75,76
(
jellyfish;
tunicates;
thaliaceans;
algae;
ctenophores;

hydrozoans;
gastropods;
polychaetes;
small
crustaceans,
like
mysids;
small
fish;
chaetognaths)

4,5,7,9,11,14,18,21,23,27,37,46,48,55,61,63,66,72,75,76
Page
947
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1167
5­
2220e
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Scorpaenidae
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162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
CB
CC
CD
CE
NF
NF
sharks;
dolphin;
seals1,4
NF
possibly
river
otters12
Brandt's
cormorant16
plankton
picker1
dawn4
NF
browser9
diurnal;
nocturnal5
dawn
&
dusk7
possibly
river
otters12
NF
NF
NF
possibly
river
otters12
lingcod15
picker2
NF
NF
possibly
river
otters12
NF
NF
NF
possibly
river
otters12
diurnal,
but
occasional
nighttime
feeding
bouts12
lingcod;
rockfishes;
sharks;

dolphin;
seals;
sea
lions1,4,7
sharks;
salmon;
dolphin;
seals1,4,10
sharks;
dolphin;
seals1,4
sharks;
dolphin;
seals1,4
Page
948
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166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
CF
CG
CH
CI
CJ
CK
CL
CM
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
11.1
(
0.1)
12.5
0.042
(
0.027)
0.026
(
0.016)
allozymes
NF
NF
NF
11.2
(
0.1)
16.7
0.054
(
0.028)
0.017
(
0.012)
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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949
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Scorpaenidae
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153
154
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160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
CN
CO
CP
CQ
CR
CS
NF
NF
NF
NF
NF
NF
118
(
2)
>
30
(
5)
hook
&
line
otoliths;
whole
No
intrareader
comparisons
45
(
1)
20
to
30
(
3
to
4)
hook
&
line
scales
No
intrareader
comparisons
5NG
(
NG)
(
5
to
7)
NG
NG
NG
NG
19NG
(
2
to
4)
NG
NG
otoliths;
break
and
burn
Marginal
increment
analysis
intrareader
comparisons
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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1167
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Scorpaenidae
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162
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164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
CT
CU
CV
CW
NF
NF
NF
NF
2NG
NG
0.26
catch
curve
8,23
0.006
(
0.001
to
0.008)
NG
NG
catch
curve
17NG
NG
1.2
(
doubtful)
tag­
recapture;
Ricker
formula
17NG
NG
0.26
age
composition;
Ricker
formula
23
0.14
NG
0.14
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
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1167
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Scorpaenidae
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164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
CX
CY
CZ
DA
NF
NF
NF
NF
5
and
older
NG
NG
NF
Young­
of­
the­
year
NG
NG
NG
NG
NG
2
to
19
NG
NG
juveniles
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
952
of
1167
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Scorpaenidae
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161
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164
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168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
DB
DC
DD
DE
5<
5
nm
Residential8,17
NF
11<
2
km
Transient9
parasitized
by
copepod
family
Neobrachiella19
school
with
olive
rockfish;
kelp
bass;
blacksmith;
halfmoon
3,25
3<
2
km
Residential1,5,10,15
Possibly1
cleaned
by
Oxyjulis
californica
and
Phanerodon
atripes9
4
<
5­
6
miles
Yes;
(<
25
m
radius)

infested
by
parasites
from
the
genus
Lironeca
24
5<
5
nm
9<
10
km
decrease
in
abundance
as
Macrosytis
beds
are
removed18
infested
by
roundworms
from
genera
Anisakis
20
5<
5
nm
Residential4,6,15
NF
seen
in
same
crevices
as
S.
rubberimus
2,25
NF
NF
NF
associated
with
S.
maliger
8
infested
by
roundworms
from
genera
Anisakis
20,22
NF
Transient11
NF
infected
by
ectoparasites
from
the
families
Caligidae
and
Lerneopodidae;
also
monogenetic
trematodes9
16<
15
miles,
but
most
<
1
mile
infected
by
ectoparasites
from
the
families
Bomolochidae,

Caligidae
and
Cymothoidae9
infected
by
parasties
from
the
families
Bomolochidae,

Caligidae,
Hemiuridaea,
Lepcreadiidae,
and
Cymothoidae22
y
p
Caligidae,
Naobranchiidae,
Philichthyidae,
Hemiuridae,

Opecoelidae,
Sanguinicolidae,
Zoogonidae,
Microcotylidae,

Anisakidae,
and
Capillariidae22
infected
by
parasites
from
the
families
Hemiuridae,

Lepocreadiidae,
Opecoelidae,
Capsalidae,
and
Anisakidae22
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
20
infected
by
parasites
from
the
families
Bothriocephalidae,

Naobranchiidae,
Microcotylidae,
Anisakidae,
Myxiddidae,

and
Myxobolidae22
Page
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Scorpaenidae
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161
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163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
DF
DG
Yes13,33
NF
No
23
Yes1,4,5,6,8,11,23,25,29,30,31,32,33
No9,10,11
No14,23
Yes10,11
No9,19,35
Yes7,14
Yes15,23
NF
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Scorpaenidae
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211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
A
B
C
D
E
S.
pinniger
1
1
<
1
to
275
S.
proriger
1
1
12
to
425
S.
rastrelliger
1
1
<
1
to
46
S.
rosaceus
1
1
7
to
262
S.
rosenblatti
1
1
61
to
396
greenblotched
rockfish;
bosco;
warthog;

starry
eye1,4,6
rosy
rockfish;
strawberry;
rinkydinky;

avocado
rockfish;
dude;
scacciatale;

corsair;
scratch­
tail;
schitzy1,4,5,6,7,12
redstripe
rockfish;
little
red
rockfish;
red
rockfish
of
Alaska1,3,9
canary
rockfish;
faintail;
red
snapper;
red
rock
cod;
red;
Pacific
red
snapper;
pinky;

swallowtail;
codalarga;
filione;
orange
rockfish;
yellow
snapper;
bosco;
orange
rockfish;
1,3,4,5,6,7,9,12,13
grass
rockfish;
grass
bass;
greenie;

pepper
bass;
rock
bass;
kelp
rock
cod;

scomoda;
green
garrupa;
schmo1,3,4,5,6,9,12
northern
CA
(
Pt.
Delgada)
to
central
Baja
California
(
Ranger
Bank)
1
southeastern
Bering
Sea
(
Pribilof
I.)
and
Aleutians
Is.(
Amchitka
I.)
to
southern
Baja
California3
OR
(
Yaquina
Bay)
to
central
California
(
Bahia
Playa
Maria)
1
WA
(
Puget
Sound)
to
central
Baja
California
(
Bahia
Tortugas)
1
western
Gulf
of
Alaska
(
south
of
Shelik
of
Strait)

to
northern
Baja
California
(
Cabo
Colonet)
1
Page
955
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211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
F
G
H
I
J
7*
Both:
W=
0.000000023TL3.0556
SE=
0.271
4TL=
11.48
+
1.239(
SL);
r2=
0.994;
SE=
8.0
1,4
76
British
Columbia
(
FL)
1
54.1
&
52.8
8*
Male:
W=
0.00001873TL2.951
(
r2=
0.990)
4TL=
­
4.107
+
1.07(
FL);
r2=
1.0;
SE=
2.8
Washingon
(
TL)
19
NG
8*
Female:
W=
0.00002182TL2.94
(
r2=
0.990)
4SL=
­
7.447
+
0.803(
TL);
r2=
0.994;
SE=
6.4
Washingon
(
FL)
32
52.8
8*
Both:
W=
0.000006883TL3.147
(
r2=
0.992)
4SL=
­
9.326
+
0.855(
FL);
r2=
0.996;
SE=
6.4
Washington
(
TL)
37
NG
12*
Male:
W
(
in
kg)=
0.0000564FL2.707
4FL=
4.180
+
0.934(
TL);
r2=
1.0;
SE=
2.6
Oregon
(
FL)
32
51.9
12*
Female:
W
(
in
kg)=
0.0000222FL2.707
4FL=
12.88
+
1.164(
SL);
r2=
0.996;
SE=
7.4
12*
Both:
W
(
in
kg)=
0.0000127FL3.12
14*
Both:
W
=
0.000000112TL3.39
(
r2=
0.854)

17*
Both:
W
=
0.03437FL2.82
(
r2=
0.93)
Oregon
to
Washington
(
FL)
7
52.1
&
55.95
22*
Male:
W=
0.1259FL2.49
Oregon
to
Washington
(
FL)
32
52.1
22*
Female:
W=
0.0245FL2.91
Oregon
to
Washington
(?
L)
36
51.9
22*
Both:
W=
0.2818FL2.28
California
to
Canada
(
FL)
5
55.72
22*
Male:
W=
0.123FL2.49
southern
Oregon
and
California
(
FL)
34
53.5
(
23.0)

22*
Female:
W=
0.7943FL2.01
c.
California
(
TL)
11
46.5
22*
Both:
W=
0.055FL2.70
22*
Male:
W=
0.0123FL3.09
22*
Female:
W=
0.0132FL3.07
22*
Both:
W=
0.0126FL3.08
24*
Male:
W=
0.0000102FL3.09
24*
Female:
W=
0.000009898FL3.09
27Male:
W=
0.000085?
L2.596
27Female:
W=
0.000065?
L3.665
28Both:
W=
0.00000000112TL3.39
(
r2=
0.854)

29
*
Male:
W=
0.014046FL3.07
29
*
Female:
W=
0.01286FL3.1
4Male:
W=
0.000008FL3.0894
(
r2=
0.965)
NF
1
61
British
Columbia
(
FL)
1
34
4Female:
W=
0.000009FL3.0652
(
r2=
0.972)
British
Columbia
(
FL)
2
33
17*
Both:
W
=
0.01931FL2.89
(
r2=
0.92)
CA
to
WA
(
FL)
9
29.52
(
8.2)

2Both:
W=
0.045TL2.77
(
r2=
0.98)
1TL/
SL=
1.219
1
56
California
(
TL)
6
NG
8*
Male:
W=
0.0001595TL2.661
(
r2=
0.861)
1SL/
TL=
0.822
c.
California
(
TL)
11
37.7
8*
Female:
W=
0.00004092TL2.894
(
r2=
0.933)

8*
Both:
W=
0.00000731TL3.178
(
r2=
0.994)

21Both:
W=
0.0000112SL3.21
1Both:
W=
0.0052TL3.38573
(
r2=
0.965)
4TL=
3.917
+
1.199(
SL);
r2=
0.992;
SE=
2.9
1
36
c.
California
(
TL)
11
30.2
8*
Male:
W=
0.00001198TL3.048
(
r2=
0.945)
4TL=
1.409
+
1.015(
FL);
r2=
0.998;
SE=
1.2
California
(
SL)
14
NG
8*
Female:
W=
0.000009049TL3.108
(
r2=
0.955)
4SL=
­
1.794
+
0.828(
TL);
r2=
0.992;
SE=
2.4
8*
Both:
W=
0.00000989TL3.088
(
r2=
0.964)
4SL=
0.225
+
0.837(
FL);
r2=
0.994;
SE=
2.3
11Both:
W=
0.000002295SL3.52
(
SL=
mm)
4FL=
­
1.085
+
0.984(
TL);
r2=
0.998;
SE=
1.2
4FL=
1.399
+
1.187(
SL);
r2=
0.994;
SE=
2.7
1Both:
W=
0.01103TL3.10572
(
r2=
0.996)
4TL=
9.567
+
1.182(
SL);
r2=
0.998;
SE=
6.7
1
48
s.
California
(
TL)
10
56.11
(
1.77)

4TL=
­
0.453
+
1.03(
FL);
r2=
1.0;
SE=
2.0
4SL=
­
7.374
+
0.844(
TL);
r2=
0.998;
SE=
3.1
Oregon
(
FL)
4
53.6
&
53.3
(
C.
I.=
52.4­
54.8
&

52.1­
54.5)

Page
956
of
1167
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Scorpaenidae
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214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
K
L
M
N
O
P
Q
R
S
T
U
55.3
&
62.1
53.8
&
53.3
0.114
&
0.137
0.209
&
0.095
0.139
­
3.98
&
­
0.045
2.0
&
­
1.05
­
2.71
&
­
0.24
NG
NG
NG
NG
46
NG
NG
0.071
NG
NG
­
22
NG
NG
NG
61.6
53.9
0.162
0.111
0.168
­
1.66
­
2.56
­
1.16
NG
NG
NG
NG
46
NG
NG
0.071
NG
NG
­
22.0
NG
NG
NG
58.2
53.4
0.167
0.143
0.182
­
1.84
­
1.46
­
0.71
NG
NG
NG
57.7
&
63.4
NG
0.19
&
0.17
0.16
&
0.13
NG
0.17
&
0.02
0.14
&
0.46
NG
NG
NG
NG
58.6
53.5
0.168
0.137
0.177
­
1.66
­
1.62
­
0.87
NG
NG
NG
56.8
NG
0.212
0.192
NG
­
0.064
0.169
NG
NG
NG
NG
66.11
NG
0.178
0.118
NG
0.595
­
0.24
NG
NG
NG
NG
62.2
(
41.4)
NG
0.196
(
0.037)
0.134
(
0.025)
NG
­
0.153
­
0.845
NG
NG
NG
NG
53.9
71.5
0.2614
0.2297
0.1123
­
0.3
0
­
0.6
NG
NG
NG
41.3
38.3
0.178
0.148
0.166
­
1.71
­
1.02
­
0.9
NG
NG
NG
42
41.7
0.32
0.15
0.12
1.2
­
0.3
­
1.6
NG
NG
NG
38.28
(
11.1)
NG
0.22
(
0.04)
0.158
(
0.021)
NG
­
0.723
(
0.28)
­
0.79
(
0.367)
NG
0.604
0.733
NG
NG
51.3
NG
NG
0.11
NG
NG
­
2.41
NG
NG
NG
36.9
37.8
0.2522
0.2552
0.2307
­
0.3
­
0.4
­
0.7
NG
NG
NG
32.9
32.5
0.1557
0.1241
0.1235
­
0.1
­
0.7
­
0.8
NG
NG
NG
NG
20.2
NG
NG
0.17416
NG
NG
­
1.13
NG
NG
NG
57.99
(
2.58)
NG
0.058
(
0.004)
0.053
(
0.006)
NG
­
2.103
(
0.23)
­
2.468
(
0.429)
NG
NG
NG
NG
0.19
&
0.18
(
C.
I.
=
0.17­
0.21
&
0.16­
0.2)
0.54
&
0.9
(
C.
I.=
­

0.03
to
1.11
&
0.49­

1.3)

0.15
&
0.18
(
C.
I.=
0.12­
0.17
&
0.15­
0.2)
NG
0.68
&
0.54
(
C.
I.=

0.39­
0.97
&
0.25­

0.83)
NG
NG
NG
NG
60.95
&
57.4
(
C=

58.1­
63.8
&
55.9­

60.0)
NG
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221
222
223
224
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229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
V
W
X
Y
otoliths;
section/
b&
b
No
?
1,2,3,11,17
otoliths;
whole
No
No
otoliths;
section/
break
&
burn
No
Interreader/
interlaboratory
comparisons
otolithd;
whole
NG
NG
otoliths;
section/
break
&
burn
No
Interreader/
interlaboratory
comparisons
otoliths;
section/
whole
No
Interreader
comparisons
otoliths;
section/
break
&
burn
No
Interreader/
interlaboratory
comparisons
otoliths;
?/
used
only
ages
>
5
yrs.
old
NG
Interreader
comparisons
otoliths;
whole
No
No
otoliths;
break
&
burn
No
Intrareader/
RACEBASE
comparisons
otoliths;
whole
No
Interreader
comparisons
otoliths;
section/
b&
b
No
?
NF
otoliths;
whole
No
No
otoliths;
b
&
b
No
Interreader
comparisons
otoliths;
section
No
Intrareader
comparisons
9
(
juvenile
growth)

otoliths;
whole
Edge
Analysis
Interreader
comparisons
otoliths;
whole
No
Interreader
comparisons
NF
otoliths;
whole
No
No
otoliths;
section
EA
at
<
6
yrs
Intrareader
comparisons
NF
otoliths;
section/
whole
No
Multiple
structure
comparison
Page
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Scorpaenidae
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228
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231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
Z
AA
AB
AC
AD
AE
AF
5,35
60­
75
1,2,5,7,8,15,16,17,19
Dioecious
Polygynous2
Viviparous3,7
No
1
0.94
18,36
69
Ovoviviparous8
5Males<
Females
21,32
60
27
67
(
Males);
69
(
Females)

7
34
1,6,7
Dioecious
Polygynous2
Viviparous3,7
No
NF
36
40
17,36>
23
NF
Dioecious
Polygynous2
Viviparous3,7
No
3
Male=
Female
21
20
17,35
>
13
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
1
21
14
Ovoviviparous10
33
18
10,36
>
50
NF
Dioecious
Polygynous2
Viviparous3
No
1
0.98
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222
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226
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228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
AG
AH
AI
AJ
AK
AL
AM
None1
328
(
4)
27
(
9)
40
(
9)
44
(
9)
otoliths;
whole
No
218.7
(
3)
29.6
(
NG)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
8NG*(
NG)
NG
(
NG)
39.5
(
NG)
49.2
(
NG)
NG
NG
11NG*
(
NG)
NG
(
NG)
41
(
NG)
48
(
NG)
NG
NG
12NG*
(
11)
NG
(
11)
39
(
12)
42.6
(
10)
otoliths;
whole
No
14NG
(
NG)
24
(
3)
35.6
(
5
to
6)
35.6
(
5
to
6)
scales
No
32NG
(
NG)
NG
(
NG)
44
(
9)­
sex
unspecified
NG
NG
3428
(
NG)
27
(
NG)
40
(
NG)
44
(
NG)
NA
NA
40
NG
(
NG)
NG
(
NG)
NG
(
NG)
NG
(
9.3)
otoliths;
?
NG
42
NG*
(
NG)
43
(
NG)
NG
(
NG)
50
(
NG)
NA
NA
43
NG*
(
NG)
NG
(
NG)
7
(
39)
47
(
8)
otoliths;
break­
and­
burn
NG
48
33
(
5)­
sex
unspecified
40.6
(
7)­
sex
unspecified
NG
NG
53
NG
(
NG)
NG
(
NG)
41
to
43
(
7
to
9)
41
to
43
(
7
to
9)
NG
NG
None1
621*
(
NG)
22
(
NG)
24.3
(
7)
26.2
(
6.5)
otoliths;
b
&
b
No
11NG*
(
NG)
NG
(
NG)
28
(
NG)
29
(
NG)
NG
NG
42
<
20*
(
NG)
28
(
NG)
>
21
28
(
NG)
NA
NA
None1
422
(
2)
22
(
NG)
24.5
(
3.5)
24
(
3.7)
otoliths;
section
No
235.9
(
8)
32.4
(
5)
NG
(
NG)
NG
(
NG)
otoliths;
whole
Edge
Analysis
4520.3
(
3
to
4)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NG
NG
None1
316
(
4)
15
(
4)
20
(
6)
20
(
6)
otoliths;
whole
No
5,53
14
(
NG)
12
(
NG)
15
(
7)
15
(
7)
NG
NG
221.8
(
NG)
24.5
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
20
(
6)­
sex
unspecified
NG
NG
None1
5,53
23
(
NG)
16
(
NG)
30
(
12)
28
(
12)
NG
NG
Page
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1167
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Scorpaenidae
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218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
AN
AO
AP
AQ
AR
AS
AT
AU
AV
NG
1
Alaska67,71,77
interreader
comparisons
British
Columbia15,35
*****

NG
Washington54
*****

NG
Oregon5,54,66,68
*****

intrareader
comparisons
Oregon
to
Washington65,70
No
c.­
n.
California5,17,77
NG
c.
California4
NA
California57,60,73
*****

interreader
comparisons
NA
NG
NG
NG
interreader
comparisons
NF
Alaska67,71
NG
British
Columbia15,70
NA
Oregon5,19,66
*****

c.­
n.
California5
intrareader
comparisons
NF
c.
California4,34
interreader
comparisons
s.
California6
*****

NG
California57
*****

No
NF
c.­
n.
California5,77
^^^^^
*****

NG
c.
California4,38
NG
s.
California13,14,18,77
*****

NG
California57,70
*****
*****
*****

NG
NF
s.
California13,14,77
*****

Page
961
of
1167
5­
2220e
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Scorpaenidae
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213
214
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216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Once13
1470
(
161­
815)
4,11260,000
(
48.3
TL)
to
1,9000,000
(
66.0
TL)

18,19820,000
(
49
FL)
to
1,320,000
(
56.9
FL)

*****
2,27
369,000
(
27
?
L)
to
1,113,000
(
54
?
L)

*****
^^^/***

^^^^^
Once5
NF
NF
*****
NF
2,2680,000
(
26
TL)
to
760,000
(
46.5
TL)

Once1
1234
(
143­
318)
1,27
12,601
(
15.2
TL)
to
95,267
(
23.5
TL)

Multiple
2
3291
(
NA)

Multiple
2
3136
(
NA)
1,27
30,600
(
32.5
TL)
to
655,050
(
46.7
TL)

Page
962
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Scorpaenidae
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218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
BF
BG
BH
BI
BJ
4F=
a
TL4.021
NA
1
~
1
to
2
months
859.4
(
SL)
1
13,998
to
29,107
7,8F=
64221.3FL(
cm)­
2330029
3<
116
days
9,1525
to
30
NF
NA
1
~
1
to
2
months
NF
NF
F1=
5.267TL3.006
(
r2=
0.95)
NA
1
~
1
to
2
months
NF
NF
F3=
0.00074TL3.6497
(
r2=
0.785)
NA
1
~
1
to
2
months
NF
NF
F3=
0.
000022TL4.3753
(
r2=
0.73)
NA
1
~
1
to
2
months
339
to
49
NF
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963
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1167
5­
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Scorpaenidae
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220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
BK
BL
BM
BN
BO
BP
130.84
to
1.45
No
external
eggs
NA
2,173.6
to
18.4
(
SL)
planktonic1,4
planktivore
18,204.0
to
18.4
(
SL)
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
303.6
(
SL)
to
NF
No
external
eggs
NA
205.4
(
TL)
to
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
NF
No
external
eggs
NA
3,7,264.6
to
27.7
(
SL)
planktonic1
planktivore
174.3
to
27
(
SL)
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
5
0.56
to
0.7
No
external
eggs
NA
253.9
(
SL)
to
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
NF
No
external
eggs
NA
NF
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
Page
964
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Scorpaenidae
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212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
BQ
BR
BS
BT
BU
BV
NF
diurnal1
siphonophores;
18.6
(
SL)
to
maturity1,7
zooplanktivore
chaetognaths1
NF
diurnal1
siphonophores;
NA
pelagic;
nearshore
sand­
rock
interface2,9
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
727
(
SL)
to
maturity
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
pelagic?;
NF3,9,22
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
pelagic?;
NF3,9,22
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids
and
copepods;
tunicates;
fish)
1,6,11,25
pelagic;
rock;
low­
growing
algae;

tidepools;
artificial
reefs3,4,9,20,26,51
pelagic;
sand,
mud,
and
gravel;
low
rock
and
cobble;
kelp
bed;
demersal;
artificial
reefs
3,8,9,13,24,29,30,39,50,54
(
crustaceans,
such
as
harpacticoids,
barnacle
cyprids,
and
euphasiid
eggs
and
larvae;

juvenile
polycahetes)
1,6,8,10,21
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965
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213
214
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217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
BW
BX
BY
BZ
CA
NF
diurnal5
planktivore/
carnivore
NF
NF
rocky
shelf;
continental
slope/
basin5
carnivore
(
small
fish;
cephalopods)
33
NF
NF
carnivore
NF
NF
rocky/
mixed
rock
substrates
5,7,18,65
carnovore
(
demersal
crustaceans;
tunicates)
33,48,76
picker1
diurnal11
carnivore
(
crustaceans;
tunicates;
cephalopods;
fish)
33,37,76
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
rocky/
non­
rocky
shelf,
canyon
and
continental
slope/
basin;
high
&
low
relief;

soft­
bottoms
5,18,21,25,31,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
(
fish,
such
as
juvenile
surfperch;
crustaceans;

cephalopods
and
gastropods)
3,9,19,23,36,43,48,60,63,75,76
low
relief
rock,
often
in
crevices;
vegetation;

kelp
forests;
artificial
reefs
5,6,7,15,17,27,32,38,53,57,64,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12,16,21
(
euphasiids
and
other
crustaceans;
small
fish,
such
as
Sebastes
jordani,
myctophids
and
stomiatiods)

6,10,23,30,40,45,48,52,58,67,76
aggegrate
around
reefs
and
pinnicles;
rocky
shelf;
larger
fish
found
at
deeper
depths
5,7,23,25,42,54,59,65
Page
966
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Scorpaenidae
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217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
CB
CC
CD
CE
NF
diurnal15
single
Pacific
stock3,12
possible
northern
range
and
southern
range
stocks
11
possibly
river
otters12
NF
NF
NF
possibly
river
otters12
NF
nocturnal16
NF
NF
NF
ambusher8
nocturnal13
NF
northern/
central
CA
(
north
of
SF);
southern
CA
(
south
of
SF)
1,9
sharks;
dolphin;
seals1,4
sharks;
salmon;
dolphin;
seals1,4,10
yelloweye
rockfish;

sharks;
salmon;
dolphin;

seals1,3,4,10
sharks;
dolphin;
seals1,4
yelloweye
rockfish;
lingcod;

salmon;
sharks;
dolphin;

seals1,4,5,8,10
Page
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Scorpaenidae
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217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
CF
CG
CH
CI
CJ
CK
CL
CM
NF
NF
11.1
(
0.1)
8.3
0.01
(
0.007)
0.01
(
0.008)
allozymes
NF
2
0.022
NG
allozymes
NF
11.1
(
0.1)
8.3
0.012
(
0.009)
0.012
(
0.009)
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
968
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Scorpaenidae
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217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
CN
CO
CP
CQ
CR
CS
633.3
FL(
6)
NG
(
NG)
trawls
otoliths;
whole
No
No
2
19
(
2)
NG
(
NG)
trawls
otoliths;
section/
whole
No
multiple
structure
comparison
720
(
4)
NG
(
7)
trawls
otoliths;
break
&
burn
No
between
readers
11NG
(
1)
NG
(
NG)
NG
NG
NG
NG
20
to
34
FL
(
4
to
5)­
Males
NG
(
NG)
trawls
otoliths;
section/
break
&
burn
No
interreader/
intelaboratory
comparison
26
to
34
FL
(
4
to
5)­
Females
NG
(
NG)
trawls
otoliths;
section/
break
&
burn
No
interreader/
intelaboratory
comparison
24
to
36
FL
(
4
to
7)­
Males
NG
(
NG)
trawls
otoliths;
section/
break
&
burn
No
interreader/
intelaboratory
comparison
26
to
36
FL
(
4
to
6)­
Females
NG
(
NG)
trawls
otoliths;
section/
break
&
burn
No
interreader/
intelaboratory
comparison
14
to
22
(
3)­
Males
NG
(
NG)
trawls
otoliths;
section/
break
&
burn
No
interreader/
intelaboratory
comparison
14
to
26
(
3
to
4)­
Females
NG
(
NG)
trawls
otoliths;
section/
break
&
burn
No
interreader/
intelaboratory
comparison
14
NG
(
3)
NG
(
3)
otoliths;
?
NG
interreader
comparisons
15NG
(
3
to
4)
NG
(
8
to
9)
hook
&
line
otoliths;
whole
NG
NG
16
36
(
6)
NG
trawls
otoliths;
break
&
burn
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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Scorpaenidae
202
203
204
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206
207
208
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210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
CT
CU
CV
CW
2NG
NG
0.262
catch
curve
3,7NG
NG
0.36
to
0.4
Chapman­
Robson
3,7NG
NG
0.15
to
0.16
catch
curve
5NG
NG
0.03
catch
curve
5NG
NG
0.04
catch
curve
5NG
NG
0.05
catch
curve
5NG
NG
0.04
catch
curve
5NG
NG
0.02
catch
curve
6NG
NG
0.89;
0.178
catch
curve
120.06;
0.6
to
0.2
0.5
0.56;
0.56
to
0.7
catch
curve
0.08
NG
NA
Reparam.
B­
H
Stock­
Recruitment
12NG
NG
0.096;
0.145
Hoenig's
(
1983)
equation
5NG
NG
0.1
catch
curve
9
0.13­
Males;
0.16­
Females
0.82­
Males;
0.46­
Females
0.95­
Males;
0.62­
Females
Hoenig
estimation/
Wetherall
9
0.1­
Males;
0.17­
Females
0.85­
Males;
0.45­
Females
0.95­
Males;
0.62­
Females
catch
curve/
Wetherall
9
0.13­
Females
0.49
Females
0.62­
Females
GSI/
Wetherall
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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1167
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Scorpaenidae
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210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
CX
CY
CZ
DA
8
to
18
NG
NG
NF
15
to
23
NG
NG
15
to
23
NG
NG
16
to
76
NG
NG
16
to
48
NG
NG
15
to
51
NG
NG
18
to
54
NG
NG
21
to
57
NG
NG
Males:
12­
34;
Females:
12­
60
NG
NG
Males
(
all
ages)/
Females
(
1­
10);
Females
11
to
>
25
0.2
(
overall)
0.71
to
0.89
NG
NG
NG
Males
47yrs.;
Females
31
yrs.
NG
NG
11
to
32
NG
NG
NF
4
to
34
NG
0.86­
Males;
0.74­
Females
4
to
34
NG
0.89­
Males;
0.73­
Females
4
to
34
NG
0.79­
Females
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
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971
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1167
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Scorpaenidae
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217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
DB
DC
DD
DE
parasitized
by
copepod
Clavellopsis
robusta
11
5to
380
nm
Transient11,22
NF
low
incidence
of
parasitism15
no
movement12,3
Residential12
NF
Residential11
Yes7
5<
5
nm
Residential1
NF
no
movement6,12
Residential8
NF
associated
with
S.
elongatus
7
NF
NF
NF
infested
by
roundworms
from
genera
Anisakis
20,22
infested
by
parasties
from
the
families
Anisakidae
and
Ceratomyxidae22
infested
by
parasites
from
the
families
Echinorhynchidae,

Bothriocephalidae,
Tentaculariidae,
Chondracanthidae,

Lernaeopodidae,
Naobranchiidae,
Pennellidae,

Philichthyidae,
Hemiuridae,
Opecoelidae,
Sanguinicolidae,

Syncoelidae,
Capsalidae,
Microcotylidae,
Anisakidae,
and
Cystidicolidae22
infested
by
parasites
from
the
families
Lacistorhynchidae
and
Anisakidae22
significant
movements,
up
to
236
km
infested
by
parasited
from
the
families
Bothriocephalidae,

Phyllobothriidae,
Tentaculariidae,
Bomolochidae,
Caligidae,

Chondracanthidae,
Lernaeopodidae,
Naobranchiidae,

Philichthyidae,
Bucephalidae,
Hemiuridae,
Lepcreadiidae,

Opecoelidae,
Sanguinicolidae,
Syncoelidae,
Casalidae,

Anisakidae,
and
Caratomyxidae22
infected
by
macroparasites
from
the
families
Microcotylidae,

Opecoelidae,
Anisakide,
and
Polymorphidae14
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
20
Page
972
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5­
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Scorpaenidae
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208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
DF
DG
Yes5
NF
NF
NF
No3,23
NF
No15
NF
NF
NF
Page
973
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Scorpaenidae
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253
254
255
256
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258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
A
B
C
D
E
S.
ruberrimus
1
1
15
to
549
S.
rubrivinctus
1
1
30
to
418
S.
saxicola
1
1
10
to
547
S.
semicinctus
1
half­
banded
rockfish;
inspector1,4,7,9
1
15
to
402
S.
serranoides
1
1
<
1
to
172
flag
rockfish;
barberpole;
red
bandit;

convict
fish;
shoflies;
Spanish
flag;
stripe
cod;
Hollywood1,3,4,5,6,7,12
stripetail
rockfish;
occhio­
grande;
popeye
rockfish;
bigeye
rockfish;
oliveback
rockfish1,4,5,7,9,12
northern
California
(
Redding)
to
central
Baja
California
(
Islas
San
Benito)
1
yelloweye
rockfish;
red
snapper;

turkeyred
rockfish;
golden
eye;
golden;
tambor;

drum;
pot
belly;
vecchia;
yellowbelly,

rasphead;
turkey
rock1,3,4,5,6,9,12
northern
WA
to
central
Baja
California
(
Bahia
de
Sebastian
Vizcaino)
1
east
Gulf
of
Alaska
(
Yakutat
Bay)
to
central
Baja
California
(
Bahia
de
Sebastian
Vizcaino)
1
nothern
California
(
San
Francisco)
to
northern
Baja
California
(
San
Quintin)
1,3
Aleutian
Is.
(
Umnak
and
Unalaska
Is.)
to
northern
Baja
California
(
Ensenada)
1
olive
rockfish;
Johnny
bass;
Jonathan;

yellowtail
rock
cod;
kelp
salmon;
kelp
bass;

kelp
yellowtail;
sugarfish1,3,4,5,6,7,9,12
Page
974
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Scorpaenidae
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266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
F
G
H
I
J
4SL=
­
8.023
+
0.7(
FL);
r2=
0.994;
SE=
2.9
4FL=
0.692
+
0.97(
TL);
r2=
1.0;
SE=
2.0
4FL=
9.938
+
1.147(
SL);
r2=
0.998;
SE=
3.3
*
8Male:
W=
0.0005893TL2.424
(
r2=
0.978)
4TL=
5.856
+
1.202(
SL);
r2=
0.992;
SE=
9.5
1
91
Alaska
(
FL)
13
69
8*
Female:
W=
0.00000378TL3.252
(
r2=
0.984)
4TL=
­
0.758
+
1.018(
FL);
r2=
1.0;
SE=
3.6
Alaska
(
FL)
22
69
to
72
*
Both8:
W=
0.000004313TL3.228
(
r2=
0.984)
4SL=
­
1.717
+
0.826(
TL);
r2=
0.992;
SE=
7.8
Oregon
(
FL)
16
64.7
(
2.2)

12*
Male:
W
(
in
kg)=
0.0000211FL2.994
4SL=
­
2.664
+
0.841(
FL);
r2=
0.992;
SE=
7.6
c.
California
(
TL)
11
75.6
12*
Female:
W
(
in
kg)=
0.0000086FL3.223
4FL=
1.296
+
0.981(
TL);
r2=
1.0;
SE=
3.6
12*
Both:
W
(
in
kg)=
0.0000127FL3.12
4FL=
6.665
+
1.181(
SL);
r2=
0.992;
SE=
9.0
14*
Male:
W
=
0.00000059TL3.18
(
r2=
0.974)

14*
Female:
W
=
0.0000004o8TL3.25
(
r2=
0.958)

14*
Both:
W
=
0.000000463TL3.22
(
r2=
0.960)

17*
Both:
W
=
0.00681FL3.26
(
r2=
0.96)

28Male:
W=
0.0000000059TL3.18
(
r2=
0.974)

28Female:
W=
0.00000000408TL3.25
(
r2=
0.958)

28Both:
W=
0.00000000463TL3.22
(
r2=
0.96)

NA
NF
1
51
NF
NF
1Male:
W=
0.03759TL2.61664
(
r2=
0.950)
4TL=
3.226
+
1.242(
SL);
r2=
0.998;
SE=
2.5
1
41
c.
to
n.
California
(
TL)
3
NG
1Female:
W=
0.02479TL2.80487
(
r2=
0.990)
4TL=
­
0.669
+
1.038(
FL);
r2=
0.998;
SE=
2.0
s.
California
(
TL)
10
17.38
(
5.81)

7*
Both:
W=
0.000000016TL3.1201
SE=
0.05
4SL=
­
2.252
+
0.804(
TL);
r2=
0.998;
SE=
2.0
California
(
SL)
14
NG
4SL=
­
2.315
+
0.831(
FL);
r2=
0.998;
SE=
2.1
4FL=
0.921
+
0.963(
TL);
r2=
0.998;
SE=
1.9
4FL=
3.234
+
1.2(
SL);
r2=
0.998;
SE=
2.5
1Male:
W=
0.0127TL3.01568
(
r2=
0.969)
4TL=
8.179
+
1.17(
SL);
r2=
0.958;
SE=
3.6
1
25
s.
California
(
TL)
10
14.62
(
0.15)

1Female:
W=
0.0152TL2.93761
(
r2=
0.978)
4TL=
­
0.422
+
1.05(
FL);
r2=
0.996;
SE=
1.2
4SL=
­
1.752
+
0.82(
TL);
r2=
0.958;
SE=
3.0
4SL=
­
0.343
+
0.849(
FL);
r2=
0.956;
SE=
3.1
4FL=
1.01
+
0.949(
TL);
r2=
0.996;
SE=
1.1
4FL=
6.486
+
1.128(
SL);
r2=
0.956;
SE=
3.56
3Male:
W=
0.0152TL2.964
(
r2=
0.968)
4TL=
8.292
+
1.209(
SL);
r2=
0.99;
SE=
7.3
1
61
c.
California
(
TL)
8
43.3
(
0.45)

3Female:
W=
0.0111TL3.063
(
r2=
0.972)
4TL=
1.419
+
1.029(
FL);
r2=
0.998;
SE=
2.6
c.
California
(
TL)
11
44
16Both:
W=
0.00978TL3.09
4SL=
­
3.542
+
0.819(
TL);
r2=
0.99;
SE=
6.0
c.
California
(
TL)
20
42.65
8*
Male:
W=
0.00001675TL2.966
(
r2=
0.964)
4SL=
­
0.672
+
0.837(
FL);
r2=
0.99;
SE=
5.7
8*
Female:
W=
0.00003015TL2.874
(
r2=
0.941)
4FL=
­
0.862
+
0.971(
TL);
r2=
0.998;
SE=
2.5
8*
Both:
W=
0.0000
631TL3.136
(
r2=
0.990)
4FL=
4.422
+
1.184(
SL);
r2=
0.99;
SE=
6.8
21Both:
W=
0.0000078SL3.18
Page
975
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267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
K
L
M
N
O
P
Q
R
S
T
U
71
69
0.09
0.08
0.09
2.5
­
7
3.1
NG
NG
NG
69
to
71
NG
0.07
to
0.09
0.07
to
0.08
NG
0.9
to
2.5
­
7
to
1.7
NG
NG
NG
NG
69.5
(
4.8)
NG
0.275
(
0.071)
0.138
(
0.044)
NG
4.87
(
0.862)
2.295
(
1.572)
NG
NG
NG
NG
64.6
66.6
0.089
0.1153
0.1117
0
­
0.2
0
NG
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NG
32.71
(
4.2)
NG
NG
0.1474
(
0.0048)
NG
NG
­
0.605
(
0.054)
NG
NG
NG
33.05
(
1.68)
NG
0.186
(
0.194)
0.064
(
0.007)
NG
­
3.782
(
2.526)
­
4.634
(
0.433)
NG
NG
NG
NG
NG
26.8
NG
NG
0.14738
NG
NG
­
0.6
NG
NG
NG
18.14
(
0.22)
NG
0.15
(
0.037)
0.22
(
0.03)
NG
­
1.458
(
0.182)
­
1.368
(
0.176)
NG
NG
NG
NG
51.9
(
0.93)
NG
0.27
(
0.02)
0.18
(
0.01)
NG
­
1.03
(
0.19)
­
1.57
(
0.23)
NG
NG
NG
NG
53.9
52.4
0.261
0.1667
0.1879
­
1.3
­
1.5
­
1
NG
NG
NG
50.19
NG
0.1991
0.2926
NG
­
0.8649
­
1.373
NG
NG
NG
NG
Page
976
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2220e
­
Scorpaenidae
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266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
V
W
X
Y
otoliths;
whole
No
No
4
otoliths;
break­
and­
burn
No
No
otoliths;
whole
No
Intrareader
comparisons
otoliths;
whole
No
Interreader
comparisons
NF
NF
NF
NF
scales
No
No
NF
otoliths;
section
Edge
Analysis;
<
6
yrs
Intrareader
comparisons
otoliths;
whole
No
No
otoliths;
whole
No
Intrareader
comparisons
NF
otoliths;
whole
No
Intrareader
comparisons
NA
otoliths;
whole
Edge
Analysis
Interreader
comparisons
otoliths;
whole
No
Intrareader
comparisons
Page
977
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Scorpaenidae
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267
268
269
270
271
272
273
274
275
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277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
Z
AA
AB
AC
AD
AE
AF
17,33,36
118
2,6,7,9,10
Dioecious
Polygynous2
Viviparous3
No
1
0.98
10,12,32,35
114
Ovoviviparous10
21
69
34
98
17,35>
18
NF
Dioecious
Polygynous2
Viviparous3,7
No
NF
21
18
3
17
7
Dioecious
Polygynous2
Viviparous3,7
No
1
0.69
21,36
38
Ovoviviparous10
35
6
(
Males);
22
(
Females)

35,36
15
NF
Dioecious
Polygynous2
Viviparous3
No
1
0.84
6,10,21,22,35
25
NF
Dioecious
Polygynous2
Viviparous3,7
No
1
0.93
Page
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Scorpaenidae
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265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
AG
AH
AI
AJ
AK
AL
AM
None1
336
(
6)
36
(
6)
40
(
7)
40
(
7)
otoliths;
whole
No
247.1
(
11)
40.8
(
10)
NG
(
NG)
NG
(
NG)
otoliths;
whole
No
21NG*(
NG)
NG
(
NG)
52
to
60
(
15
to
20)
50
to
52
(
15
to
20)
otoliths;
b
&
b
No
12NG*
(
9)
NG
(
10)
NG
(
12)
NG
(
11)
otoliths;
whole
No
32NG
(
NG)
NG
(
NG)
40
(
7)­
sex
unspecified
NG
NG
42
39*
(
NG)
36
(
NG)
45
(
NG)
41
(
NG)
NA
NA
19,52
NG
(
NG)
NG
(
NG)
NG
(
15)
NG
(
18)
NG
NG
None1
3,53
29
(
5)
29
(
5)
30
(
5)
34
(
8)
otoliths;
whole
No
23NG
(
NG)
NG
(
NG)
38
(
NG)
38
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
34
(
8)­
sex
unspecified
NG
NG
None1
315
(
3)
17
(
2)
16
(
3)
17
(
2)
otoliths;
whole
No
59
(
NG)
9
(
NG)
10
(
NG)
10
(
NG)
NG
NG
11NG*
(
NG)
NG
(
NG)
<
17
(
NG)
<
20
(
NG)
NG
NG
1412.7
(
2)
12.7
(
2)
14.6
(
NG)
17.1
(
4)
scales
No
32NG
(
NG)
NG
(
NG)
17
(
2)­
sex
unspecified
NG
NG
None1
510
(
NG)
10
(
NG)
11
(
NG)
11
(
NG)
NG
NG
3,53
32
(
4)
32
(
4)
33
(
5)
35
(
5)
otoliths;
whole
No
9,17,4928.1
(
3)
31.1
(
3)
32
(
5)
34
(
4)
otoliths;
whole
No
None1
226.9
(
NG)
28.5
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
32NG
(
NG)
NG
(
NG)
35
(
5)­
sex
unspecified
NG
NG
36NG
(
NG)
NG
(
NG)
33
to
36
(
4
to
5)­
sex
unspecified
NG
NG
5118
to
22
(
NG)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NG
NG
longer
pectoral
fin
ray
and
orbit
width
per
SL
in
males3
Page
979
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1167
5­
2220e
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Scorpaenidae
252
253
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259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
AN
AO
AP
AQ
AR
AS
AT
AU
AV
No
3
Alaska20,44,46,67,71,77
interreader
comparisons
British
Columbia15
No
Washington7,8,70
intrareader
comparisons
Oregon5,16,66
NG
n.­
c.
California5
^^^^^
*****

NA
c.
California4
NG
California28,57
*****
*****
*****

No
NF
Washington8
NG
Oregon16,46
NG
n.­
c.
California5,46
s.
California46
California57
No
1
British
Columbia15,35,70
NG
Oregon5,35,66,70
NG
n.­
c.
California5,17,33,77
*****

No
c.
California45
*****
*****
*****

NG
s.
California13,77
California57
*****
*****
*****

NG
NF
s.
California13,77
*****

No
NF
n.­
c.
California5
^^^^^
*****

intrareader
comparisons
c.
California4,37
*****

NG
s.
California29
*****

NG
California27,57,61,77
*****

NG
NG
Page
980
of
1167
5­
2220e
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Scorpaenidae
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261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Once1
1300
(
271­
324)
26
2.,
700,000
(
NG)

Multiple4
4
303.4
(
NA)

*****
*****
NF
NF
NF
*****
Once2,23
1415
(
169­
931)
1,27
1,245
(
9.5
TL)
to
131,382
(
28.0
TL)

3215
(
NA)
4,9,1115,000
(
17.8
TL)
to
200,000
(
31.8
TL)

^^^^^

*****

*****
Once2,23
3199
(
NA)
1,27
3,409
(
11.7
TL)
to
30,858
(
18.5
TL)

Once1,9,12
2130
(
NA)
5,1030,000
(
32.6
TL)
to
490,000
(
46.8
TL)

1330
(
NA)

Page
981
of
1167
5­
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Scorpaenidae
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263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
BF
BG
BH
BI
BJ
NF
NA
1
~
1
to
2
months
NF
NF
NF
NA
1
~
1
to
2
months
NF
NF
F3=
0.000548TL3.7925
(
r2=
0.939)
NA
1
~
1
to
2
months
9,11~
30
NF
3
118
days
15
27
to
53
F3=
0.000289TL3.9157
(
r2=
0.827)
NA
1
~
1
to
2
months
9
34
NF
F5=
0.006TL34.619
(
r2=
0.74)
NA
1
~
1
to
2
months
141
to
66
NF
930
to
40
16
25
Page
982
of
1167
5­
2220e
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Scorpaenidae
252
253
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261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
BK
BL
BM
BN
BO
BP
5
0.52
to
0.56
No
external
eggs
NA
204.2
(
TL)
to
planktonic1
planktivore
215.0
(
TL)
to
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
234.2
(
SL)
to
NF
No
external
eggs
NA
NF
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
NF
No
external
eggs
NA
43.3
(
NL)
to
20
(
SL)
planktonic1
planktivore
83.7
(
SL)
to
(
nauplii
and
invertebrate
eggs;
copepods)
1,2,3
204.7
to
214.3
(
TL)
to
NF
No
external
eggs
NA
255.5
(
SL)
to
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
NF
No
external
eggs
NA
174.8
(
SL)
to
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3,4
Page
983
of
1167
5­
2220e
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Scorpaenidae
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260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
BQ
BR
BS
BT
BU
BV
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
20
(
SL)
to
maturity2
zooplanktivore
chaetognaths1
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids
and
calenoid
copepods)
1,6,11,15
pelagic;
soft
substrate;
low
relief
rock;

algal
beds;
sand­
silt
interface3,9,13,18
pelagic;
nearshore
sand­
rock
interface;

algal
beds2,9,13,46
(
crustaceans,
such
as
calanoids,
zoea,
and
barnacle
cyprids;
juvenile
fishes;
polychaetes;

octopi
and
squid)
1,6,7,8,10,13,22
pelagic;
shallow,
broken­
rock;
sponge
gardens;
high­
relief
within
crevices
3,9,11,12,33,49
pelagic;
drifting
kelp
mats;
artificial
reefs
3,9,20,26,41,54
pelagic;
kelp
canopy;
rocky
shelf;
drifting
kelp;
surfgrass
beds;
artificial
reefs
3,4,6,9,22,24,26,28,32,41,48,54
Page
984
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1167
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Scorpaenidae
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261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
BW
BX
BY
BZ
CA
NF
NF
carnivore
NA
rocky
substrates;
artificial
reefs
5,32,65
carnivore
(
crustaceans;
cephalopods;
fish)
20,33,48,76
picker1
diurnal7,11
planktivore
NF
NF
NF
browser2
dawn1
(
meso)
carnivore8
large
juveniles­
nocturnal2,6,7,8,12,13
small
juveniles­
diurnal5,8
albacore26
non­
rocky
shelf;
continental
slope/
basin;

sandy/
soft
bottoms;
mud­
cobble
5,7,17,18,21,25,31,46,65
(
crustacenas,
such
as
euphasiids
and
calenoid
copepods)
6,33,37,48,76
(
fish,
including
many
adult
and
juvenile
rockfishes;

crustaceans;
gastropods;
lingcod
eggs)
12,24,30,48,76
soft
to
mixed
bottoms;
bank
edges;

cobblemud7,18,25,62,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
rocky
shelf;
continental
slope/
basin;
broken
rock
and
boulder
fields;
high,
complex
pinnacles;
crevices
and
cracks
5,7,9,12,16,23,25,35,52,54,65
fishes,
including
rockfishes
(
such
as
adult
yelloweye)

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,10,12
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
(
fish,
especially
juvenile
rockfishes
like
S.
mystinus
;

small
crustaceans;
polychaetes;
cephalopods;

tunicates)
7,8,9,18,19,20,21,23,25,41,48,53,61,63,69,75,76
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
rocky
shelf
and
shelf;
kelp
canopy
in
midwater
column;
artificial
reefs
5,7,13,14,17,27,28,32,37,38,45,49,50,51,60,64,65
Page
985
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1167
5­
2220e
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Scorpaenidae
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261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
CB
CC
CD
CE
NF
NF
none
within
British
Columbian
waters10
possibly
river
otters12
NF
NF
sharks;
dolphin;
seals1,4
NF
picker8
nocturnal10,13
NF
possibly
river
otters12
NF
NF
NF
NF
diurnal;
nocturnal5
sharks;
dolphin;
seals1,4
NF
nocturnal7,8,10,13
sharks;
dolphin;
seals1,4
sharks;
salmon;
dolphin;
seals1,4,10
sharks;
salmon;
dolphin;
seals1,4,10
Page
986
of
1167
5­
2220e
­
Scorpaenidae
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
CF
CG
CH
CI
CJ
CK
CL
CM
NF
NF
NF
NF
NG
0.41
to
0.89
microsatellites
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
11.0
(
0.0)
4.2
0.009
(
0.009)
0.009
(
0.009)
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
11.3
(
0.1)
20.8
0.065
(
0.039)
0.048
(
0.025)
allozymes
NF
Page
987
of
1167
5­
2220e
­
Scorpaenidae
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
CN
CO
CP
CQ
CR
CS
135
(
7)
>
50
(
11)
hook
&
line
otoliths;
whole
No
intrareader
comparisons
9NG
(
13)
NG
(
30)
hook
&
line
otoliths;
break­
and­
burn
No
intrareader
comparisons
15NG
(
9)
NG
(
NG)
hook
&
line
otoliths;
whole
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1825.4
(
NG)
NG
NG
NG
NG
NG
Page
988
of
1167
5­
2220e
­
Scorpaenidae
252
253
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256
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260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
CT
CU
CV
CW
2NG
NG
0.33
catch
curve
13
0.02
NG
0.0174
±
0.0053
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
989
of
1167
5­
2220e
­
Scorpaenidae
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
CX
CY
CZ
DA
14
and
older
NG
NG
NF
NG
NG
NG
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Page
990
of
1167
5­
2220e
­
Scorpaenidae
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
DB
DC
DD
DE
seen
in
same
crevices
as
S.
nigrocinctus
2
no
movement6,12,15,19
Residential13,22
NF
may
contain
parasite
Hemiurus
levinseni
6
associated
with
S.
maliger
8
infested
by
roundworms
from
genera
Anisakis
20
NF
NF
NF
associated
with
Microstomus
pacificus
7
NF
NF
NF
infested
by
parasites
from
the
family
Microcotylidae
22
parasitized
by
copepod
family
Neobrachiella19
NF
NF
NF
associated
with
surfpearch;
boccacio1
6,12<
1
km
Transient/
Residential1,5,6
NF
schooling
with
S.
mystinus
4
9<
10
km
cleaned
by
Oxyjulis
californica
9
decrease
in
abundance
as
Macrosytis
beds
are
removed18
competes
with
Paralabraz
clathratus
for
food
and
shelter23
infested
by
parasties
from
the
families
Echinorhynchidae,

Polymorphidae,
Bothriocephalidae,
Phyllobothriidae,

Tentaculariidae,
Bomolochidae,
Caligidae,

Chondracanthidae,
Lernaeopodidae,
and
Philichthyidae,

Bucephalidae,
Fellodistomatidae,
Gorgoderidae,
Hemiuridae,

Lepocreadiidae,
Opecoelidae,
Zoogonidae,
Capsalidae,

Anisakidae,
Capillariidae,
Cucullanidae,
and
Cystidicolidae22
infested
by
parasites
from
the
families
Caligidae,

Lernaeopodidae,
Anisakidae,
and
Myxobolidae22
infested
by
parasites
from
the
families
Bomolochidae,

infected
by
ectoparasites
from
the
families
Bomolochidae
and
Caligidae;
also
gnathiid
larvae
and
monognentic
trematodes9
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
20
Page
991
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1167
5­
2220e
­
Scorpaenidae
252
253
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257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
DF
DG
No
23
NF
No15,23
NF
No18
NF
Yes13
NF
Yes3,4,5,11,23,32,35
No9,10,11
No10
Page
992
of
1167
5­
2220e
­
Scorpaenidae
302
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311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
A
B
C
D
E
S.
serriceps
1
1
<
1
to
64
Sebastolobus
alascanus
1
1
20
to
1524
Sebastolobus
altivelis
1
1
20
to
1756
1­
Robins
et
al.
1991
1­
Eschmeyer
1998
1­
Robins
et
al.
1991
1­
Love
et
al.
2000
1­
Love
et
al.
2000
2­
Chen
1986
2­
Miller
&
Lea
1972
2­
Love
et
al.
1987
3­
Love
1996
3­
Klingbeil
&
Knaggs
1976
4­
Eschmeyer
1983
4­
Shaw
1999
5­
Phillips
1939
5­
Orr
&
Blackburn
2000
6­
Leos
1999
6­
Erickson
et
al
1991
7­
Phillips
1957
7­
Love
&
Lea
1997
8­
Shaw
1999
8­
O'Connell
et
al
1992
9­
Gates
&
Frey
1974
10­
Hart
1973
11­
Turner
et
al.
1969
12­
Reilly
et
al.
1994
13­
Roedel
1948
treefish;
convict
bass;
lipstick
fish;
gopher
1,3,6,13
longspine
thornyhead;
channel
rockfish;

hardhead;
hooligan;
idiotfish
lobe­
finned
rockfish;
anglefin
rockfish
bonehead;

scorpion;
deep­
water
rockfish
crimson
thornhead1,3,4,7,9
Eastern
aleutian
Is.
to
southern
Baja
California
(
Cabo
San
Lucas)
1
shortspine
thornyhead;
channel
rockfish;

hardhead;
idiotfish;
fagiano;
scorpion;

bonehead;
spiny­
cheaked
rockfish;

hooligan;
lobe­
finned
rockfish;
gurnard;

gurnet;
channel
cod;
deep
sea
red
rock
cod1,3,4,5,7,9,10,13
Bering
Sea
(
Navarin
Canyon)
to
central
Baja
California
(
Isla
Cedros)
1
northern
California
(
San
Francisco)
to
central
Baja
California
(
Isla
Cedros)
1
Page
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Scorpaenidae
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311
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313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
F
G
H
I
J
21Both:
W=
0.0000182SL3.07
NF
1
41
NF
NF
18Both:
W(
kg)=
0.0000049TL3.264
5Fresh
FL=
3.82
+
1.00(
frozen
FL)
r2=
0.98
3
84.6
Alaska
(
SL)
38
NG
25*
Both:
W=
0.000002651TL3.264
(
r2=
0.99)
5Fresh
FL=
10.76
+
1.169
(
frozen
SL)
r2=
0.966
Alaska
(
FL)
38
NG
30Both:
dressed
W
=
0.00000484(
fresh
SL)
3.244
r2=
0.995Fresh
FL=
6.11
+
1.157
(
fresh
SL)
r2=
0.996
Alaska
(
SL)
38
NG
30Both:
dressed
W=
0.00000484(
fresh
FL)
3.264
r2=
0.9925Fresh
SL=
2.6
+
1.0128
(
frozen
SL)
r2=
0.976
Alaska
(
SL)
38
NG
30Both:
W(
kg)=
0.00000163(
fresh
SL)
3.2932
r2=
0.992
5Fresh
SL=
­
4.79
+
0.86262
(
fresh
FL)
r2=
0.998
Alaska
(
SL)
38
NG
30Both:
W(
kg)=
0.00000136(
fresh
FL)
3.3904
r2=
0.992
Oregon
(
FL)
28
NG
Jacobson
1991
NG
c.
California
(
TL)
23
NG
c.
California
(
TL)
25
NG
25*
Both:
W=
0.000001794TL3.352
(
r2=
0.99)
2TL=
13.51
+
1.182(
SL);
r2=
0.99;
SE=
2.82
1
39
Jacobson
1991
NG
2TL=
1.76
+
1.01(
FL);
r2=
0.998;
SE=
1.3
c.
California23
NG
2SL=
­
9.74
+
0.84(
TL);
r2=
0.99;
SE=
1.92
2SL=
­
0.852
+
0.847(
FL);
r2=
0.99;
SE=
1.96
2FL=
­
1.07
+
0.991(
TL);
r2=
0.998;
SE=
1.3
2FL=
12.02
+
1.17(
SL);
r2=
0.99;
SE=
2.3
1­
Love
et
al.
1990
1­
Lea
et
al
1999
1­
Love
et
al
2000
1­
Archibald
et
al
1981
2­
Love
&
Johnson
1998
2­
Karpov
&
Kwiecien
1988
2­
Shaw
1999
2­
Westrheim
&
Harling
1975
3­
Love
&
Westphal
1981
3­
MacGregor
1983
3­
Kline
1996
3­
Phillips
1964
4­
Shaw
1999
4­
Echeverria
&
Lenarz
1984
4­
Williams
et
al.
1999
4­
Six
&
Horton
1977
5­
Patten
1973
5­
Miller
1985
5­
Boehlert
&
Kappenman
1980
6­
Miller
et
al
1967
6­
Love
&
Johnson
1998
7­
Phillips
1964
7­
Wilson
&
Boehlert
1990
8­
Lea
et
al
1999
8­
Love
&
Westphal
1981
9­
Love
et
al
1987
9­
Shaw
1999
10­
Stein
&
Hassler
1989
10­
Love
et
al
1990
11­
Chen
1971
11­
Lea
et
al
1999
12­
McClure
1982
12­
Love
et
al
1987
13­
Barker
1979
13­
Rosenthal
et
al
1981
14­
Washington
et
al.
1978
14­
Chen
1971
15­
Moulton
1977
15­
Miller
&
Geibel
1973
16­
Love
1978
16­
McClure
1982
17­
Rosenthal
et
al
1982
17­
Barker
1979
18­
Rogers
et
al
1998
18­
Moulton
1977
19­
Zaitlin
1986
19­
Washington
et
al.
1978
20­
MacGregor
1983
20­
Love
1978
21­
Quast
1968b
21­
Six
1976
Page
994
of
1167
5­
2220e
­
Scorpaenidae
302
303
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309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
K
L
M
N
O
P
Q
R
S
T
U
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NG
48.36
NG
NG
0.025
NG
NG
­
6.94
NG
NG
NG
NG
57.82
NG
NG
0.024
NG
NG
­
7.75
NG
NG
NG
NG
34.53
NG
NG
0.05
NG
NG
­
3.77
NG
NG
NG
NG
32.2
NG
NG
0.06
NG
NG
­
2.74
NG
NG
NG
NG
67.91
NG
NG
0.016
NG
NG
­
8.11
NG
NG
NG
NG
72.8
NG
NG
0.023
NG
NG
NG
NG
NG
NG
NG
87.2
(
3.3)
NG
NG
0.0145
(
0.001)
NG
NG
­
6.05
(
0.78)
NG
NG
NG
NG
94.5
(
7.3)
NG
NG
0.0169
(
0.0024)
NG
NG
­
5.54
(
1.14)
NG
NG
0.896
NG
70
NG
NG
0.03
NG
NG
1.45
NG
NG
NG
NG
33.7
(
1.2)
NG
NG
0.0585
(
0.0057)
NG
NG
­
0.38
(
0.56)
NG
NG
NG
NG
30.1
(
3.8)
NG
NG
0.072
(
0.0028)
NG
NG
­
1.9
(
0.21)
NG
NG
0.933
Page
995
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1167
5­
2220e
­
Scorpaenidae
302
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312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
V
W
X
Y
NF
NF
NF
NF
otoliths:
break
and
burn
No
No
8
otoliths:
break
and
burn
No
No
otoliths:
break
and
burn
(
mean
ages
weighted
by
n0.5/
No
No
otoliths:
break
and
burn
(
mean
ages
weighted
by
1/
s2
No
No
otoliths;
whole
No
No
otoliths;
section/
break
and
burn
radiometric
Interreader
comparisons
otoliths;
section
radiometric
Inter/
intrareader
comparisons
None
NG
NG
Pending
Pending
Pending
8
otoliths;
section
radiometric
Inter/
intrareader
comparisons
1­
Phillips
1964
2­
Westrheim
&
Harling
1975
3­
Chen
1971
4­
O'Connell
&
Funk
1986
5­
Miller
&
Geibel
1973
6­
Richards
&
Cass
1986
7­
Yoklavich
&
Boehlert
1987
8­
Ianelli
et
al.
1994
9­
Laidig
and
Sakuma
1998
10­
Boehlert
&
Yoklavich
1983
11­
Boehlert
&
Yoklavich
1986
12­
Wales
1953
13­
Miller
et
al.
1967
14­
Miller
1985
15­
Dewees
&
Gothshall
1974
16­
Tenera
Environmental
Services
2000
17­
Boehlert
&
Yoklavich
1984b
14
(
also
includes
Gompertz,
logistic,
power,
and
exponential
equations)

Page
996
of
1167
5­
2220e
­
Scorpaenidae
302
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311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
Z
AA
AB
AC
AD
AE
AF
NF
NF
Dioecious
Polygynous2
Viviparous3,7
No
NF
9
>
80
7,12,20
Dioecious
Polygynous
Oviparous2,10,11
No
4
Male<
Female
13
29
(
ovuliparity)
3
14
100
19,20,35
62
30
56
9
45
NF
Dioecious
Polygynous
Oviparous2,11
No
NF
(
ovuliparity)
3
1­
Love
et
al
1987
1­
Archibald
et
al
1981
1­
Love
et
al
1987
1­
Love
et
al
1987
1­
Lenarz
&
Echeverria
1991
2­
Miller
&
Geibel
1973
2­
Washington
et
al
1978
2­
Wourms
1991
2­
Moser
1996c
2­
Love
et
al.
1987
3­
Phillips
1964
3­
Barker
1979
3­
Wourms
1991
3­
Love
&
Johnson
1998
4­
Larson
1991
4­
Miller
et
al
1967
4­
Dygert
&
Gunderson
1991
4­
Tokranov
&
Novikov
1997
5­
Wilson
&
Boehlert
1990
5­
Boehlert
1980
5­
Boehlert
&
Yoklavich
1984a
5­
STAT
1999
6­
Love
1978
6­
Rosenthal
et
al.
1981
6­
Yoklavich
&
Boehlert
1991
7­
Franklin
1999
7­
Westrheim
&
Harling
1975
7­
Eigenmann
1892
8­
Zaitlin
1986
8­
Six
1976
8­
STAT
1999
9­
Kline
1996
9­
Rosenthal
et
al
1982
9­
Coyer
1979
10­
Casillas
et
al.
1998
10­
O'Connell
&
Funk
1986
10­
Garrison
&
Miller
1982
11­
VenTresca
1992
11­
Richards
&
Cass
1986
11­
Love
1996
12­
O'Connell
&
Funk
1986
12­
Tokranov
&
Novikov
1997
12­
Larson
1977
13­
Tokranov
&
Novikov
1997
13­
Richards
&
Cass
1986
14­
Butler
et
al.
1995
14­
Gotshall
1969
15­
Richards
&
Cass
1986
15­
STAT
1999
16­
Worton
2000
16­
Williams
et
al
1999
17­
Bloeser
1999
17­
Sampson
&
Stewart
1994
18­
STAT
1999
18­
Gowen
1983
19­
Ianelli
et
al.
1994
19­
Barss
et
al.
1982
20­
Miller
1985
20­
Miller
1985
21­
Reilly
et
al.
1994
21­
Baxter
1999a
Page
997
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
AG
AH
AI
AJ
AK
AL
AM
None1
NF
NF
NF
NF
NF
NF
None1
22
21.5
(
NG)
21.5
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
2416.5
(
5)
16.5
(
5)
21
(
13)
21
(
13)
otoliths;
section
No
27NG
(
NG)
NG
(
NG)
NG
(
NG)
22
(
NG)
NG
NG
29NG
(
NG)
NG
(
NG)
NG
(
13)
NG
(
13)
otoliths;
section/
b
&
b
radiometric
31
NG
(
NG)
17
(
NG)
NG
(
NG)
22
(
NG)
NA
NA
42
<
16*
(
NG)
26
(
NG)
<
16
(
NG)
30
(
NG)
NA
NA
4413.5^
(
5)
NG
(
NG)
19.2
(
12)
19.2
(
12)
otoliths;
break
and
burn
No
53
NG
(
NG)
NG
(
NG)
23
(
12)
23
(
12)
NG
NG
None1,4
1625.2
(
NG)
27.6
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
22
21.5
(
NG)
21.5
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
23NG
(
NG)
17
(
NG)
NG
(
NG)
NG
(
NG)
NG
NG
24NG
(
NG)
NG
(
NG)
NG
(
NG)
19
(
14)
otoliths;
section
No
31
NG
(
NG)
19
(
NG)
NG
(
NG)
21.5
(
NG)
NA
NA
42
NG*
(
NG)
NG
(
NG)
21
(
NG)
NG
(
NG)
NA
NA
53
NG
(
NG)
NG
(
NG)
10
to
18
(
NG)
10
to
18
(
NG)
NA
NA
1­
Chen
1971,1986
1­
Love
et
al
1987
2­
Moser
1967
2­
Lea
et
al
1999
3­
Wyllie
Echeverria
1986
3­
Wyllie
Echeverria
1987
4­
Wakefield
&
Smith
1990
4­
Love
&
Johnson
1998
5­
Love
et
al.
1990
6­
Shaw
1999
7­
Miller
&
Geibel
1973
8­
Gunderson
et
al
1980
9­
Love
and
Westphal
1981
10­
Larson
1980
b
11­
Westrheim
1975
12­
McClure
1982
13­
Miller
et
al
1967
14­
Phillips
1964
15­
Washington
et
al.
1978
16­
Best
1964
17.
Love
1978
18­
Barker
1979
19­
Yamanaka
&
Kronlund
1997
20­
Wales
1953
21­
Rosenthal
et
al.
1982
Page
998
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
AN
AO
AP
AQ
AR
AS
AT
AU
AV
NF
NF
NF
NG
NF
Alaska69
NG
Oregon30,66
NG
northeast
Pacific41
interreader
comparisons
c.
California46
NA
California47,52,56,70,77
*****

NA
No
NG
NG
NF
Oregon30,66
NG
northeast
Pacific41
NG
n.
California31
NG
c.
California48
NA
California47,52,56,77
*****
*****

NA
NA
1­
Chen
1971
1­
Love
et
al
1987
2­
Barker
1979
2­
Phillips
1968
3­
Yamanaka
&
Kronlund
1997
3­
David
1939
4­
Lea
et
al
1999
5­
Wyllie
Echverria
1987
6­
Love
&
Johnson
1998
7­
Washington
et
al.
1978
8­
DeLacy
et
al
1964
9­
Hitz
&
DeLacy
1965
10­
Larson
1980
a,
b,
c
11­
Patten
1973
12­
Moulton
1977
13­
Love
et
al
1990
14­
Chen
1971
15­
Westrheim
1975
16­
Hitz
1962
17­
Phillips
1964
18­
Moser
1967
19­
Shaw
1999
20­
Rosenthal
et
al.
1981
21­
Dunn
&
Hitz
1969
Page
999
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Once1,17
1192
(
NA)
NF
Multiple11,16,23
NF
1441,000
(
NG)
to
400,000
(
NG)

27
20,000
(
NG)
to
450,000
(
NG)

Multiple11,18,23
NF
143,000
(
NG)
to
106,000
(
NG)

1­
MacGregor
1970
1­
MacGregor
1970
1­
MacGregor
1970
2­
Love
et
al
1990
2­
Love
et
al
1990
2­
Love
et
al
1990
3­
Miller
&
Giebel
1973
3­
Haldorson
&
Love
1991
3­
Love
&
Johnson
1998
4­
Washington
et
al.
1978
4­
Hart
1973
4­
Haldorson
&
Love
1991
5­
Shaw
1999
5­
Tenra
Environmental
Services
2000
5­
Phillips
1964
6­
DeLacy
et
at
1964
6­
Love
&
Westphal
1981
7­
Stein
et
al
1989
7­
DeLacy
et
al
1964
8­
Miller
et
al
1967
8­
Miller
&
Giebel
1973
9­
Love
&
Westphal
1981
9­
Miller
et
al
1967
10­
VenTresca
et
al.
1996
10­
Hart
1973
11­
Bloeser
1999
11­
Love
1978
12­
Love
1978
12­
Matarese
et
al
1989
13­
Guillemot
et
al.
1985
13­
Wales
1953
14­
Matthews
1990a
14­
Casillas
et
al.
1998
15­
Wyllie
Echeverria
1987
15
Wakefield
&
Smith
1990
16­
Casillas
et
al.
1998
16­
Romero
1988
17­
Larson
1992
(
LR)
17­
Zaitlin
1986
18­
Wakefield
&
Smith
1990
18­
STAT
1999
19­
Romero
1988
19­
Gunderson
et
al.
1980
20­
Wallace
&
Tagart
1994
20­
Wang
1981
21­
Baxter
1999a
21­
Adams
1992a
Page
1000
of
1167
5­
2220e
­
Scorpaenidae
302
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311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
BF
BG
BH
BI
BJ
NF
NA
1
~
1
to
2
months
NF
NF
13F=
19,007
+
378.99
(
dressed
W)
r2=
0.817
6~
2
to
3
weeks
1,2
~
1
to
2
months
5,1322
to
27
(
SL)
NF
13F=
18,399
+
318.03
(
total
W)
r2=
0.823
1250
(
TL)

13F=
0.05295
(
fresh
SL)
2.642
r2=
0.584
14~
25
(
SL)

13F=
0.02782
(
fresh
FL)
2.6788
r2=
0.858
13F=
­
227,852
=
98.156
(
dressed
W)
+
1,296
(
fresh
SL)
+
727.571
(
age)
r
2=
0.85
13F=
­
234,841
+
93.197
(
dressed
W)
+
1,398.51
(
fresh
SL)
r
2=
0.848
NF
NF
1,2
~
1
to
2
months
2,5,1342
to
56
(
SL)
NF
12,14
55
1­
Love
&
Johnson
1998
1­
Orton
1955
1­
Moser
1996c
1­
Lea
et
al.
1999
1­
STAT
1999
2­
DeLacy
et
al
1964
2­
Barnhart
1932
2­
Moser
1974
2­
Moser
et
al.
1977
3­
Love
et
al
1990
3­
David
1939
3­
Krigsman
2000
3­
Chen
1971
4­
Haldorson
and
Love
1991
4­
Wourms
1991
4­
Tenera
Environem4­
Moser
&
Butler
1981
5­
Love
&
Westphal
1981
5­
Moser
1974
5­
Moser
1996c
6­
Washington
et
al.
1978
6­
Moser
pers
com.
6­
Boehlert
&
Yoklavich
1983
7­
Matarese
et
al.
1989
7­
Love
1996
7­
Laroche
&
Richardson
1980
8­
Gunderson
et
al.
1980
8­
Richardson
&
Laroche
1979
9­
Romero
1988
9­
Anderson
1983
10­
Zaitlin
1986
10­
Stein
&
Hassler
1989
11­
Richards
&
Schnute
1990
11­
Johnson
1997
12­
Wallace
&
Tagart
1994
12­
Casillas
et
al.
1998
13­
Miller
1985
13­
Moser
1974
14­
Garrison
&
Miller
1982
14­
Wakefield
&
Smith
1990
15­
Krigsman
2000
16­
Watters
1992
17­
Houk
1992b
18­
Garrison
&
Miller
1982
19­
Miller
1960a
20­
Tenera
Environmental
Services
2000
Page
1001
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
BK
BL
BM
BN
BO
BP
5
0.7
to
1.0
No
external
eggs
NA
NF
planktonic1
planktivore
(
nauplii
and
invertebrate
eggs;
copepods)
2,3
1,8,16
1.2
to
1.4
planktonic2
NF
5,332.6
(
TL)
to
planktonic1,2
planktivore
71.1
X
1.2
6,17,322.6
to
22
(
SL)

17
1.3
28to
~
25.4
(
NA)

293.0
to
20
(
SL)

81.2
to
1.4
planktonic2
NF
52.6
(
TL)
to
planktonic1
planktivore
71.1
X
1.2
6,172.6
to
22
(
SL)

11
1.1
293.0
to
20
(
SL)

17
1.3
1­
Hart
1973
1­
Moser
1996c
1­
Tenera
Environmental
Services1­
Moser
1996c
1­
Morris
1956b
2­
David
1939
2­
Pearcy
1962
2­
Richardson
&
Laroche
1979
2­
Boehlert
&
Yoklavich
1984a
2­
Sumida
et
al.
1985
3­
Barnhart
1932
3­
Feder
et
al.
1974
3­
Laidig
&
Sakuma
1998
3­
Karpov
et
al.
1995
3­
Moser
&
Boehlert
1991
4­
Orton
1955
4­
Love
1996
4­
Laidig
et
al.
1996
4­
STAT
1999
4­
Love
1978
5­
MacGregor
1970
5­
Westrheim
1975
5­
Miller
1985
6­
Boehlert
&
Yoklavich
1984a
6­
Moser
et
al.
1977
6­
Baxter
1999a
7­
Moser
1996c
7­
Moreno
1993
7­
Holbrook,
Carr,
Schmitt,
&
Coyer
1990
8­
Pearcy
1962
8­
Morris
1956b
8­
Tenera
Environmental
Services
2000
9­
Washington
et
al.
1984
9­
Hart
1973
10­
Wales
1953
10
Hitz
&
DeLacy
1960
11­
Wakefield
&
Smith
1990
11­
Dygert
&
Gunderson
1991
12­
Zaitlin
1986
12­
Laroche
&
Richarson
1980
13­
Waldron
1968
13­
David
1939
14­
Wang
1981
14­
Orton
1955
15­
Breder
&
Rosen
1966
15­
Boehlert
&
Yoklavich
1984a
16­
Garrison
&
Miller
1982
16­
Moser
&
Butler
1981
17­
Morris
1956b
17­
Moser
1996c
18­
Feder
et
al.
1974
18­
Stahl­
Johnson
1985
19­
Washington
et
al.
1984
20­
Matarese
et
al
1989
21­
DeLacy
et
al.
1964
Page
1002
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
BQ
BR
BS
BT
BU
BV
NF
diurnal1
siphonophores;
NA
zooplanktivore
chaetognaths1
(
crustaceans,
such
as
barnacle
cyprids)
1,6
shelter
among
urchin­
filled
crevices52
NF
diurnal1
NF
3,7,1020
(
SL)
to
maturity
pelagic;
shallow
continetal
shelf19
NF
NF
diurnal1
NF
3,1020
(
SL)
to
maturity
pelagic;
shallow
continetal
shelf19
carnivores
722
(
SL)
to
maturity
1­
Love
et
al
1987
1­
Moser
&
Bohlert
1991
1­
Yoklavich
et
al.
1996
1­
Richardson
&
Laroche
1979
1­
Larson
1980
a
1­
Love,
Carr,
Haldorson
1991
2­
Laidig
et
al.
1996
2­
VenTrescae
t
al.
1996
2­
Miller
&
Geibel
1973
3­
Moser
et
al.
1977
3­
Love,
Carr,
Haldorson
1991
3­
Gotshall
&
Smith
1965
4­
Laroche
&
Richardson
1980
4­
Bloeser
1999
4­
Prince
&
Gotshall
1976
5­
Boehlert
&
Yoklavich
1984a
5­
Love
&
Westphal
1990
5­
Hobson
&
Chess
1988
6­
Moser
&
Butler
1981
6­
Limbaugh
1955
6­
Gaines
&
Roughgarden
1987
7­
Moser
1996c
7­
Hoelzer
1988
7­
Limbaugh
1955
8­
Washington
et
al.
1984
8­
Miller
&
Geibel
1973
8­
Singer
1985
9­
Anderson
1983
9­
Moser
&
Boehlert
1991
9­
Prince
1975
10­
Moser
1974
10­
Boehlert
&
Yoklavich
1984
10­
Singer
1982
11­
O'Connell
&
Carlisle
1993
11­
Casillas
et
al.
1998
12­
Richards
1986
12­
Haldorson
&
Richards
1986
13­
Anderson
1983
13­
Hobson
&
Chess
1976
14­
Matthews
1990a
14­
Smith
&
Brown
1983
15­
Stein
&
Hassler
1989
15­
Allen,
M.
J.
1982
16­
Patten
1973
16­
Murie
1995
17­
Rosenthal
et
al.
1982
17­
Hueckel
&
Stayton
1982
18­
Johnson
1997
18­
Coyer
1979
19­
Vetter
&
Lynn
1997
19­
Adams
1992
a
20­
Casillas
et
al.
1998
20­
Adams
1992
b
21­
Love
1992
21­
Adams
1992
c
deep,
silt­
covered
cobble
and
boulder
bottoms43
pelagic;
drifting
kelp
mats/
high
relief
rock;

artificial
reefs3,9,20,22,26,41,54
(
pelagic
crustaceans,
such
as
euphausiids)
11,14
Page
1003
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5­
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­
Scorpaenidae
302
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312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
BW
BX
BY
BZ
CA
NF
NF
(
meso)
carnivore8
(
crustaceans;
mollusks;
fish)
8,9,25,63,65,76
Least
terns27
NF
NF
carnivore
NF
diurnal?
9
carnivore
1­
Allen,
M.
J.
1982
1­
Hobson
&
Chess
1976
1­
Love,
Carr,
Haldorson
1991
1­
Love
et
al
1987
1­
Love
et
al
1987
2­
Ebeling,
Larson,
Ale2­
Love
&
Ebeling
1978
2­
Morejohn
et
al.
1978
2­
Larson
1980
a,
b,
c
2­
Larson
1980
a
3­
Coyer
1979
3­
Love
1978
3­
Baltz
1976
Thesis
3­
Dunn
&
Hitz
1969
3­
Carlisle
et
al.
1964
4­
Singer
1985
4­
Rosenthal
et
al
1988
4­
Miller
&
Geibel
1973
4­
Miller
&
Geibel
1973
5­
Singer
1982
5­
Miller
&
Geibel
1973
5­
Bloeser
1999
5­
Hobson,
Chess
&
Howard
1996
6­
Hobson
et
al
1981
6­
Roberts
1979
6­
Laidig
&
Sakuma
1998
6­
Phillips
1964
7­
Casillas
et
al.
1998
7­
Hoelzer
1987
7­
Starr
1998
7­
Love
&
Ebling
1978
8­
Hobson
&
Chess
1976
8­
Ainley
et
al.
1981
8­
Love
et
al.
1998
8­
Quast
1968a
9­
Smith
&
Brown
1983
9­
Stein
&
Hassler
1989
9­
Richards
1986
9­
Quast
1968d
10­
Ebling
&
Brey
1976
10­
Rosenthal
et
al
1982
10
­
Hart
1973
10­
Bloeser
1999
11­
Allen,
M.
J.
1982
11­
Van
Dykhuizen
1983
11­
Limbaugh
1955
11­
Gotshall
&
Smith
1965
12­
Hallacher
&
Roberts
1985
12­
Ainley
et
al
1993
12­
Carlson
&
Straty
1981
12­
Rosenthal
et
al
1988
13­
Love
1996
13­
Casillas
et
al.
1998
13­
Love
1978
13­
Prince
&
Gotshall
1976
14­
Houk
1992
a,
b
14­
Hallacher
1977
14­
Hobson
&
Chess
1988
15­
Haldorson
&
Richards
1996
15­
Love
&
Johnson
1998
15­
Larson
1991
16­
Follet
&
Ainley
1976
16­
O'Connell
&
Carlisle
1993
16­
Love
&
Westphal
1990
17­
Jacobson
&
Vetter
1996
17­
Allen
1985
17­
Moulton
1977
18­
Hubbs
et
al.
1970
18­
Love
et
al
1990
18­
Roberts
1979
19­
Eldridge
and
Staff
1994
19­
Matthews
1990
a
19­
Limbaugh
1955
20­
Adams
1992
b
20­
Stein
&
Hassler
1989
20­
Love
1978
21­
Adams
1992
c
21­
Cross
1987
21­
Hallacher
1977
(
crustaceans,
such
as
caridean
shrimp,
amphipods,

and
isopods;
polychaetes;
juvenile
thornyheads)

10,59,76
shallow
rocky,
high­
relief
substrates;

crevices;
artificial
reefs
5,7,8,10,11,27,32,51,64,65
(
ophiuroids;
crustaceans;
fish;
mollusks;

polychaetes)
33,35,76
adult
longspine
and
shortspine
thornyheads13,17
adult
longspine
and
shortspine
thornyheads13,17
continental
slope/
basin
in
sand
or
mud
5,21,22,30,65
soft
bottoms
of
continental
slope/
basin
5,21,22,26,65
fishes,
including
rockfishes,

lingcod,
cabezon
and
salmon;

birds;
porpoise1,2,3,5,6,8,12
Page
1004
of
1167
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2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
CB
CC
CD
CE
ambusher6,11
nocturnal5,8,12,21
sharks;
dolphin;
seals1,4
NF
ambusher4,5
NF
sablefish13,14
None8
ambusher4,5
NF
NF
1­
Love
&
Ebling
1978
1­
Prince
&
Gotshall
1976
1­
Morejohn
et
al.
1978
FB
169
1­
Vetter
et
al.
2000
2­
Rosenthal
et
al.
1988
2­
Moulton
1977
2­
Bloeser
1999
2­
Wallace
et
al.
1999
3­
Hobson
et
al
1981
3­
Patten
1973
3­
Rosenthal
et
al.
1988
3­
STAT
1999
4­
Casillas
et
al.
1998
4­
Roberts
1979
4­
Antonelis
&
Fiscus
1980
4­
Rocha­
Olivares
&
Vetter
1999;
Rocha­
Olivares
1998
5­
Jacobson
&
Vetter
1996
5­
Love
1978
5­
Rosenthal
et
al.
1982
5­
Stepien
1995
6­
Ebeling
&
Bray
1976
6­
Hallacher
1977
6­
Casillas
et
al.
1998
6­
Love
&
Larson
1978
8­
Allen,
M.
J.
1982
7­
Singer
1982
7­
Houk
1992
a,
b
7­
Seeb
1998
9­
Ebeling,
Larson,
Alevizon,
&
Bray
1980
8­
Hobson
et
al
1981
8­
Adams
1992
c
8­
Gharrett
et
al.
2000
10­
Coyer
1979
9­
Van
Dykhuizen
1983
9­
Jacobson
&
Vetter
1996
9­
Buonaccorsi
et
al.
2000
a,
b
11­
Hobson
1994
10­
Casillas
et
al.
1998
10­
Merkel
1957
10­
Yamanaka
et
al.
2000
12­
Larson
1977
11­
Hobson
&
Chess
1976
11­
Houk
1992a
11­
Boehlert
1980
12­
Ebling
&
Bray
1976
12­
Stevens
et
al.
1984
12­
Wishard
et
al.
1980
13­
Allen,
M.
J.
1982
13­
Allen,
M.
J.
1982
13­
Wallace
&
Tagart
1994
14­
Murie
1995
14­
Conway
1967
15­
Brodeur
&
Pearcy
1984
15­
Fitch
&
Laveneberg
1975
16­
Holbrook
&
Schmitt
1988a
16­
Hubbs
et
al.
1970
17­
Coyer
1979
17­
Love
1996
18­
Hallacher
&
Roberts
1985
19­
Hobson
1994
20­
Carr
&
Rhodes
unpub.

21­
Love
1996
sablefish;
shortspine
thornyheads6,9
Alaskan
Gyre;
California
Current;
Cortes
Bank5
Page
1005
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1167
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2220e
­
Scorpaenidae
302
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316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
CF
CG
CH
CI
CJ
CK
CL
CM
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
2
15
4
0.0536
0.049
(
0.048)
allozymes
NF
NF
NF
NF
2
10
4
0.0447
0.047
(
0.045)
allozymes
NF
1­
Rocha­
Olivares
&
Vetter
1999;
R1­
Rocha­
Olivares
&
Vetter
1999
1­
Seeb
1986
1­
Seeb
1986
1­
Seeb
1986
2­
Stepian
1995
2­
Rocha­
Olivares
1998
2­
Johnson
et
al
1973
2­
Siebenaller
1978
2­
Wishard
et
al
1980
3­
Seeb
1998
4­
Siebenaller
1978
5­
Yamanaka
et
al.
2000
shared
pair
of
haplotypes
from
northern
and
southern
California
individuals2
Page
1006
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
CN
CO
CP
CQ
CR
CS
NF
NF
NF
NA
NA
NA
17NG
NG
(
16)
trawl
otolith;
break
and
burn
No
No
NF
NF
NF
NF
NF
NF
1­
McClure
1982
2­
Six
1976
3­
Barker
1979
4­
Miller
et
al
1967
5­
Hart
1973
6­
Boehlert
1980
7­
Shaw
1999
8­
Love
et
al.
1987
9­
O'Connel
&
Funk
1986
10­
Worton
2000
11­
Dorn
2000
12­
Wallace
et
al.
1999
13­
Wallace
&
Tagart
1994
14­
Sampson
&
Stewart
1994
15­
Gowen
1983
16­
Barss
et
al.
1982
17­
Miller
1985
18­
Miller
1960b
19­
Laidig
et
al.
in
prep.
Page
1007
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
CT
CU
CV
CW
NF
NF
NF
NF
0.06
0.06
0.12
Hoenig's
equation
(
1983)

15
0.05
NG
NG
NG
19
0.05/
0.07/
0.09
NG
NG
Hoenig's
equation
(
1983)

22
0.07
NG
0.117­
Males;
0.171­
Females
catch
curve
19
0.08
to
0.12
NG
NG
NG
1­
Barker
1979
2­
McClure
1982
3­
Six
and
Horton
1977
4­
Washington
et
al.
1978
5­
Archibald
et
al.
1981
6­
Wilson
&
Boehlert
1990
7­
Six
1976
8­
Adams
and
Howard
1996
9­
Franklin
1999
10­
STAR
Panel
1999
11­
Wallace
et
al.
1999
12­
STAT
1999
13­
Yamanaka
&
Kronlund
1997
14­
Rosenthal
et
al
1982
15­
Butler
et
al.
1995
16­
Dorn
2000
17­
Gotshall
1969
18­
Walton
1979
19­
Ianelli
et
al.
1994
20­
Wallace
&
Tagart
1994
21­
Gowan
1983
Page
1008
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
CX
CY
CZ
DA
NF
NF
NF
NF
NG
NG
NG
NF
NG
NG
NG
NG
NG
NG
16
to
56
NG
NG
NG
NG
NG
NG
1­
Larson
1980
a
2­
Larson
1977
Page
1009
of
1167
5­
2220e
­
Scorpaenidae
302
303
304
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306
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310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
DB
DC
DD
DE
no
movement6,9
Residential2,5,15
Yes7
infested
by
roundworms
from
genera
Anisakis
20
infested
by
various
Myxosporida22
NF
NF
NF
infested
by
various
Myxosporida22
NF
NF
NF
1­
Carlisle
et
al.
1964
1­
Starr
(
pers.
com.)
1­
Miller
&
Geibel
1973
1­
Miller
&
Geibel
1973
2­
O'Connell
&
Carlisle
1993
2­
Love
et
al.
1987
2­
Love
et
al
1987
2­
Hallacher
1977
3­
Heilprin
1992
3­
Miller
&
Geibel
1973
3­
Palsson
1998
3­
Heilprin
1992
4­
Burge
&
Schultz
1973
4­
Miller
et
al
1967
4­
Moulton
1977
4­
Matthews
1990
b
5­
Larson
1980
a
5­
Lea
et
al.
1999
5­
Stephens
et
al.
1994
5­
Matthews
1990
c
6­
Lloyd
1938
6­
Love
1978
6­
Love
1978
6­
Hallacher
1984
7­
Allen,
M.
J.
1982
7­
Barker
1979
7­
Dunn
&
Hitz
1969
7­
Casillas
et
al.
1998
8­
Murie
et
al.
1994
8­
Hallacher
1977
8­
Hallacher
1977
8­
Matthews
et
al
1986
9­
Hobson
1971
9­
Hartmann
1987
9­
Love
1980
9­
Coombs
1979
10­
Larson
1977
10­
Matthews
1986
10­
Heilprin
1992
10­
Tenera
Environmental
Services
2000
11­
Fraser
1919
11­
Matthews
1985
11­
Casillas
et
al.
1998
12­
Reilly
et
al.
1994
12­
Love
1980
12­
Gascon
&
Miller
1981
13­
Houk
1992a
13­
Van
Dykhuizen
1983
13­
Coombs
1979
14­
Alvarado­
Villamar
&
Ruiz­
Campos
1992
14­
Culver
1986
14­
Buckley
&
Huekel
1985
15­
Liston
et
al.
1960
15­
Coombs
1979
15­
Karpov
et
al.
1995
16­
Hallacher
&
Roberts
1985
16­
Gotshall
1969
16­
Walton
1979
17­
Hallacher
1977
17­
Wallace
&
Tagart
1994
17­
Reilly
et
al.
1994
18­
Bodkin
1988
18­
Gowen
1983
18­
Wallace
&
Tagart
1994
19­
Dojiri
1981
19­
DeMott
1983
19­
Gowan
1983
20­
Dailey
et
al.
1981
20­
Tenera
Environmental
Services
2000
20­
Lenartz
(
pers.
comm.)
in
Gowen
1983
21­
Holbrook,
Carr,
Schmitt,
&
Coyer
1990
21­
Carr
&
Rhodes
unpub.
21­
Baxter
1999a
infested
by
parasites
from
the
families
Caligidae,

Opecoelidae,
Sanguinicolidae,
Gnathiid
larvae,
Capsalidae,

and
Anisakidae22
y
p
,

Caligidae,
Gnathiid
larvae,
Cymothoidae,
Anisakidae,

Ceratomyxidae,
Myxidiidae,
Myxobolidae,
and
Sinuolineidae22
infected
by
ectoparasites
from
the
families
Caligidae;
also
gnathiid
larvae
and
monognentic
trematodes9
Page
1010
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1167
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2220e
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Scorpaenidae
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313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
DF
DG
No15
Yes3,11,15
No17
NF
No15,17
NF
1­
Miller
&
Geibel
1973
1­
Love
et
al
1987
2­
Barnhart
1932
2­
Larson
1980
a
3­
Carlisle
et
al.
1964
3­
Larson
1980
b
4­
Miller
&
Geilbel
1973
4­
Larson
1980
c
5­
Starr
1998
5­
Larson
1977
6­
Hobson
&
Chess
1988
6­
Haaker
1978
7­
Moulton
1977
7­
Hart
1973
8­
Roberts
1979
8­
Hoelzer
1982,1987
9­
Hart
1973
9­
Love
1978
10­
Limabaugh
1955
10­
Hallacher
1977
11­
Hallacher
1977
11­
Love
1980
12­
Love
&
Johnson
1998
12­
Matthews
1990
b
13­
Love
et
al
1990
13­
Hallacher
1984
14­
Rosenthal
et
al.
1982
14­
Garrison
&
Miller
1982
15­
Casillas
et
al.
1998
15­
Feder
et
al.
1974
16­
Turner
et
al
1969
16­
Love
1996
17­
Jacobosn
&
Vetter
1996
18­
Allen,
M.
J.
1982
19­
Murie
et
al.
1994
20­
Gascon
&
Miller
1981
21­
Wallace
et
al.
1999
Page
1011
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
A
B
C
D
E
Page
1012
of
1167
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2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
F
G
H
I
J
22­
STAT
1999
22­
Rosenthal
et
al.
1982
23­
Wallace
et
al.
1999
23­
Kline
1996
24­
Williams
et
al.
1999
24­
Romero
1988
25­
Ianelli
et
al.
1994
25­
Rogers
et
al
1998
26­
Wallace
&
Tagart
1994
26­
Zaitlin
1986
27­
Sampson
&
Stewart
1994
27­
MacGregor
1983
28­
Gowen
1983
28­
Bulter
et
al.
1995
29­
Barss
et
al.
1982
29­
Worton
2000
30­
Miller
1985
30­
Culver
1986
31­
Karpov
et
al
1995
32­
STAT
1999
33­
Wallace
et
al.
1999
34­
Williams
et
al.
1999
35­
Wallace
&
Tagart
1994
36­
Sampson
&
Stewart
1994
37­
Gowen
1938
38­
Miller
1985
39­
Laidig
et
al.
in
prep.

Page
1013
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
K
L
M
N
O
P
Q
R
S
T
U
Page
1014
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
V
W
X
Y
Page
1015
of
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­
Scorpaenidae
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357
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361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
Z
AA
AB
AC
AD
AE
AF
22­
Watters
1992
23­
Houk
1992a
24­
Houk
1992b
25­
VenTresca
1992
26­
Wallace
&
Tagart
1994
27­
Sampson
&
Stewart
1994
28­
Laidig
et
al.
in
prep.

29­
Barker
1979
30­
Miller
1985
31­
Baxter
1999a
32­
CDF
&
G
1995
33­
Tenera
Environmental
Services
2000
34­
Yamanaka
&
Kronlund
1997
35­
Love
1996
36­
Laidig
et
al.
in
prep.
Page
1016
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
AG
AH
AI
AJ
AK
AL
AM
22­
Kline
1996
23­
Casillas
et
al.
1998
24­
Owen
&
Jacobson
1992
25­
Richards
&
Cass
1986
26­
Romero
1988
27­
Rogers
et
al
1998
28­
Zaitlin
1986
29­
Butler
et
al.
1995
30­
Coyer
1979
31­
Ianelli
et
al.
1994
32­
Reilly
et
al.
1994
33­
Adams
1992
a
34­
Adams
1992
b
35­
Adams
1992
c
36­
Watters
1992
37­
Houk
1992a
38­
Houk
1992b
39­
Wallace
&
Tagart
1994
40­
Sampson
&
Stewart
1994
41­
Gowan
1983
42­
Barss
1989
43­
Barss
et
al.
1982
44­
Miller
1985
45­
Fitch
&
Lavenberg
1975
46­
Baxter
1999a
47­
Miller
1960a
48­
Phillips
1960a
49­
Miller
1960b
50­
Phillips
1960c
51­
Feder
et
al.
1974
52­
Tenera
Environmental
Services
2000
53­
Love
1996
54­
Laidig
et
al.
in
prep.

55­
Larson
1977
Page
1017
of
1167
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Scorpaenidae
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381
382
383
384
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386
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388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
AN
AO
AP
AQ
AR
AS
AT
AU
AV
22­
Stein
&
Hassler
1989
23­
Wales
1953
24­
Miller
&
Geibel
1973
25­
Helvey
1982
26­
Burge
&
Schultz
1973
27­
Fitch
1958
28­
Eigenmann
1891
29­
Love
&
Westphal
1981
30­
Pearcy
1962
31­
Best
1964
32­
VenTresca
et
al.
1995
33­
Yoklavich
et
al.
1996
34­
Moreno
1993
35­
Hart
1973
36­
Hitz
&
DeLacy
1960
37­
Love
1978
38­
Hallacher
1977
39­
Boehlert
&
Yoklavich
1984a
40­
Laroche
&
Richardson
1980
41­
Moser
1996c
42­
Matthews
1990a
43­
Love
&
Larson
1978
44­
Rosenthal
et
al
1982
45­
Johnson
1997
46­
Casillas
et
al.
1998
47­
Moser
1974
48­
Wakefield
&
Smith
1990
49­
Romero
1988
50­
Zaitlin
1986
51­
Carr
1983
52­
Siebenaller
1978
53­
Buckley
&
Hueckel
1985
54­
STAT
1999
55­
Coyer
1979
56­
Ianelli
et
al.
1994
57­
Reilly
et
al.
1994
58­
Adams
1992
a
59­
Adams
1992
b
60­
Adams
1992
c
61­
Watters
1992
62­
Houk
1992a
63­
Houk
1992b
64­
VenTresca
1992
65­
Sampson
&
Stewart
1994
66­
Barss
1989
67­
O'Connelll
1987
68­
Barss
et
al.
1982
69­
Miller
1985
70­
Garrison
&
Miller
1982
71­
O'Connell
1987
72­
Miller
1960a
73­
Phillips
1960a
74­
Phillips
1960c
75­
Feder
et
al.
1974
76­
Tenera
Environmental
Services
2000
Page
1018
of
1167
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2220e
­
Scorpaenidae
355
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366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
AW
AX
AY
AZ
BA
BB
BC
BD
BE
22­
Tenera
Environemenal
Services
2000
22­
Adams
1992b
23­
Love
1996
23­
Adams
1992c
24­
VenTresca
1992
25­
Wallace
&
Tagart
1994
26­
Tenera
Environmental
Services
2000
27­
Love
1996
Page
1019
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
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359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
BF
BG
BH
BI
BJ
Page
1020
of
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2220e
­
Scorpaenidae
355
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366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
BK
BL
BM
BN
BO
BP
22­
Anderson
1983
23­
Kendall
1989
(
NMFS
89­
21)

24­
Stein
&
Hassler
1989
25­
Moser
1967
26­
Moreno
1990
27­
Casillas
et
al.
1998
28
Owen
&
Jacobson
1992
29­
Moser
1974
30­
Waldron
1968
31­
Wang
1981
32­
Miller
1985
33­
Garrison
&
Miller
1982
34­
Tenera
Environmental
Services
2000
Page
1021
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
BQ
BR
BS
BT
BU
BV
22­
Kendall
&
Lenarz
1986
22­
Watters
1992
23­
Turner
et
al
1969
(
FB
146)
23­
Houk
1992b
24­
Carr
1983,1991a
24­
VenTresca
1992
25­
Allen,
M.
J.
1982
25­
Love
1996
26­
Carlisle
et
al.
1964
27­
West
et
al.
1994
28­
Danner
et
al.
1994
29­
Gascon
&
Miller
1981
30­
Gascon
&
Miller
1982
31­
Bodkin
1988
32­
Larson
&
DeMartini
1984
33­
Richards
et
al.
1985
34­
Pearcy
&
Myers
1974
35­
Hueckel
&
Stayton
1982
36­
Holbrook,
Schmitt,
&
Ambrose
1990
37­
Buckley
&
Hueckel
1985
38­
Karpov
et
al.
1995
39­
STAT
1999
40­
Walton
1979
41­
Mitchell
&
Hunter
1970
42­
Coyer
1979
43­
Miller
1985
44­
Dewees
&
Gotshall
1974
45­
Baxter
1999a
46­
Nishimoto
2000
47­
Ebeling
&
Bray
1976
48­
DeMartini
1981
49­
Richards
et
al.
1986
50­
Yoshiyama
et
al.
1986
51­
Grossman
1982
52­
Feder
et
al.
1974
53­
Tenera
Environemental
Services
2000
54­
Love
1996
Page
1022
of
1167
5­
2220e
­
Scorpaenidae
355
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364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
BW
BX
BY
BZ
CA
22­
Houk
1992a
22­
Wakefield
II
1990
22­
Larson
1972,
1977
23­
Houk
1992b
23­
Rosenthal
et
al
1982
23­
Lea
et
al
1999
24­
Bodkin
1988
24­
Burge
&
Schultz
1973
24­
Washington
et
al
1978
25­
Hallacher
&
Roberts
1985
25­
Yoklavich
et
al
in
press
25­
Hobson
et
al
1981
26­
Miller
1960b
26­
Wilderbuer
1986
26­
Matthews
1990
a
27­
Love
1996
27­
Turner
et
al.
1969
27­
Wales
1953
28­
Bodkin
1986
28­
Stein
&
Hassler
1989
29­
Hobson
&
Chess
1976
29­
Patten
1973
30­
Wakefield
&
Smith
1990
30­
Rosenthal
et
al
1982
31­
Allen,
M.
J.
1982
31­
Van
Dykhuizen
1983
32­
Carlisle
et
al.
1964
32­
Burge
&
Schultz
1973
33­
Johnson
et
al.
1994
33­
Casillas
et
al.
1998
34­
West
et
al.
1994
34­
Hobson
&
Chess
1976
35­
Murie
et
al.
1994
35­
Smith
&
Brown
1983
36­
Richards
1987
36­
Mitchell
1953
37­
Bodkin
1988
37­
Allen,
M.
J.
1982
38­
Larson
&
DeMartini
1984
38­
Murie
1994
39­
Richards
et
al.
1985
39­
Murie
1995
40­
Hobson
&
Chess
1986
40­
Brodeur
&
Pearcy
1984
41­
Huekel
&
Stayton
1982
41­
Bodkin
1988
42­
STAT
1999
42­
Huekel
&
Stayton
1982
43­
Walton
1979
43­
Holbrook
&
Schmitt
1988a
44­
Coyer
1979
44­
Buckley
&
Hueckel
1985
45­
Ebeling,
Larson,
&
Alevizon
1980
45­
STAT
1999
46­
Mearns
et
al.
1980
46­
Eldridge
and
Staff
1994
47­
Borton
1982
47­
Coyer
1979
48­
Love,
Stephens,
et
al.
1996
48­
Reilly
et
al.
1994
49­
Stephens
et
al.
1984
49­
Carr
&
Rhodes
unpub.

50­
Hallacher
&
Roberts
1985
50­
Adams
1992
a
51­
Ebeling
&
Bray
1976
51­
Adams
1992
b
52­
Richards
et
al.
1986
52­
Adams
1993
c
53­
Norris
1963
53­
Watters
1992
54­
DeMott
1983
54­
Houk
1992a
55­
Crozier
1966
55­
Houk
1992b
56­
Crozier
1968
56­
VenTresca
1992
57­
Grossman
1982
57­
Diaz
&
Hammann
1987
58­
Love
et
al.
1986
58­
Barss
et
al.
1982
59­
Phillips
1960a
59­
Miller
1985
60­
Miller
1960b
60­
Fitch
&
Lavenberg
1975
61­
Phillips
1960c
61­
Hallacher
&
Roberts
1985
62­
Tenera
Environemental
Services
2000
62­
Ebeling
&
Bray
1976
63­
Carr
and
Rhodes
1974
63­
Holbrook
et
al.
1997
64­
Feder
et
al.
1974
64­
Crozier
1966
65­
Love
1996
65­
Winget
1968
66­
Larson
1977
66­
Miller
1960a
67­
Phillips
1960a
69­
Miller
1960b
70­
Phillips
1960c
71­
Tenera
Environemental
Services
2000
72­
Hobson
1994
73­
Carr
and
Rhodes
1974
74­
Bray
1978
75­
Feder
et
al.
1974
76­
Love
1996
Page
1023
of
1167
5­
2220e
­
Scorpaenidae
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359
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361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
CB
CC
CD
CE
22­
Larson
1977
Page
1024
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
CF
CG
CH
CI
CJ
CK
CL
CM
Page
1025
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
CN
CO
CP
CQ
CR
CS
Page
1026
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
CT
CU
CV
CW
22­
Miller
1985
23­
Tenera
Environmental
Services
2000
Page
1027
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
CX
CY
CZ
DA
Page
1028
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
DB
DC
DD
DE
22­
Love
&
Moser
1983
22­
Love
1996
22­
DeMott
1983
23­
Feder
et
al.
1974
23­
Tenera
Environmental
Services
2000
24­
Bennett
1993
25­
Love
1996
Page
1029
of
1167
5­
2220e
­
Scorpaenidae
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
DF
DG
22­
Walton
1979
23­
Reilly
et
al.
1994
24­
Houk
1992a
25­
Houk
1992b
26­
Wallace
&
Tagart
1994
27­
Gowen
1983
28­
Richards
et
al.
1986
29­
DeMott
1983
30­
Miller
1960a
31­
Hobson
1994
32­
Feder
et
al.
1974
33­
Tenera
Environemental
Services
2000
34­
Love
et
al
.
1987
35­
Love
1996
36­
Larson
1977
Page
1030
of
1167
5­
2220e
­
Scorpaenidae
410
411
A
B
C
D
E
Page
1031
of
1167
5­
2220e
­
Scorpaenidae
410
411
F
G
H
I
J
Page
1032
of
1167
5­
2220e
­
Scorpaenidae
410
411
K
L
M
N
O
P
Q
R
S
T
U
Page
1033
of
1167
5­
2220e
­
Scorpaenidae
410
411
V
W
X
Y
Page
1034
of
1167
5­
2220e
­
Scorpaenidae
410
411
Z
AA
AB
AC
AD
AE
AF
Page
1035
of
1167
5­
2220e
­
Scorpaenidae
410
411
AG
AH
AI
AJ
AK
AL
AM
Page
1036
of
1167
5­
2220e
­
Scorpaenidae
410
411
AN
AO
AP
AQ
AR
AS
AT
AU
AV
77­
Love
1996
78­
Larson
1977
Page
1037
of
1167
5­
2220e
­
Scorpaenidae
410
411
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Page
1038
of
1167
5­
2220e
­
Scorpaenidae
410
411
BF
BG
BH
BI
BJ
Page
1039
of
1167
5­
2220e
­
Scorpaenidae
410
411
BK
BL
BM
BN
BO
BP
Page
1040
of
1167
5­
2220e
­
Scorpaenidae
410
411
BQ
BR
BS
BT
BU
BV
Page
1041
of
1167
5­
2220e
­
Scorpaenidae
410
411
BW
BX
BY
BZ
CA
Page
1042
of
1167
5­
2220e
­
Scorpaenidae
410
411
CB
CC
CD
CE
Page
1043
of
1167
5­
2220e
­
Scorpaenidae
410
411
CF
CG
CH
CI
CJ
CK
CL
CM
Page
1044
of
1167
5­
2220e
­
Scorpaenidae
410
411
CN
CO
CP
CQ
CR
CS
Page
1045
of
1167
5­
2220e
­
Scorpaenidae
410
411
CT
CU
CV
CW
Page
1046
of
1167
5­
2220e
­
Scorpaenidae
410
411
CX
CY
CZ
DA
Page
1047
of
1167
5­
2220e
­
Scorpaenidae
410
411
DB
DC
DD
DE
Page
1048
of
1167
5­
2220e
­
Scorpaenidae
410
411
DF
DG
Page
1049
of
1167
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2220e
­
Scorpaenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Serranidae1
Paralabrax
clathratus
1
1
2
to
61
Paralabrax
maculatofasciatus
1
1<
1
to
61
Paralabrax
nebulifer
1
1shallow
to
183
1­
Robins
et
al.
1991
1­
Eschmeyer
1998
1­
Gates
&
Frey
1974
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Young
1963
2­
Allen
et
al.
1995
3­
Robins
et
al.
1991
4­
Cordes
1992
5­
Roedel
1948
kelp
bass;
rock
bass;
calico;
cabrilla;
sand
bass;
red
spotted
rock
bass;
bull
bass;
kelp
salmon;
lockee
cod1,2,3,4,5
TAXONOMY
RANGE
WA
(
Columbia
River)
to
s.
Baja
CA
(
Bahia
Magdalena)
1
spotted
sand
bass;
spotted
bass;
rock
bass;
cabrilla;
red
spotted
rock
bass;

spotted;
spotted
cabrilla;
pinta
cabrilla1,3,5
c.
CA
(
San
Francisco
Bay,
but
rare
north
of
Santa
Monica
Bay)
to
Mexico
(
Mazatlan)
1,2
barred
sand
bass;
rock
bass;
Johnny
verde;
kelp
bass;
sand
bass;
ground
bass;

sugar
bass;
cabrilla;
California
sand
bass;

California
rock
bass1,3,5
c.
CA
(
Santa
Cruz)
to
s.
Baja
CA
(
Bahia
Magdalena)
1
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32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
F
G
H
I
JK
Length
weight
equation
Max.
reported
Geographic
area
Linf
in
cm
Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
2Both:
W=
0.0000066SL3.26
(<
250
mm)
2TL=
1.41
+
1.2SL
1,2
72
s.
California
(
TL)
2
NG
NG
2Both:
W=
0.000123SL2.73
(>
250
mm)
2TL=
0.83
+
1.01FL
3Both:
W(
oz.)=
0.00376TL(
in.)
3.27
2FL=
2.38
+
1.1SL
4Both:
W=
0.0119TL(
in.)
3.119
(
r2=
0.99)

1Both:
W=
0.000026(
SL)
3.0187
(
r2=
0.9739)
NF
1
56
s.
California
(
SL)
1
NG
NG
4Both:
W=
0.1197TL(
in.)
2.712
(
r2=
0.95)
1TL=
7.27
+
1.21(
SL);
r2=
0.99;
SE=
9.12
1
65
s.
California
(
TL)
2
NG
NG
1TL=
­
2.03
+
1.05(
FL);
r2=
1.0;
SE=
3.04
1SL=
­
4.80
+
0.823(
TL);
r2=
0.99;
SE=
7.53
1SL=
­
6.52
+
0.861(
FL);
r2=
0.99;
SE=
7.62
1FL=
2.10
+
0.956(
TL);
r2=
1.0;
SE=
2.9
1FL=
8.99
+
1.154(
SL);
r2=
0.99;
SE=
8.83
2TL=
0.7
+
1.22SL
2TL=
0.46
+
1.05FL
2FL=
1.11
+
1.17SL
1­
Allen
et
al.
1995
1­
Karpov
&
Kwiecien
1988
1­
Love
et
al
2000
1­
Allen
et
al.
1995
2­
Quast
1968b
2­
Love,
Brooks,
Busatto,
Stephens,
&
Gregory
1996a
2­
Hartney
1996
2­
Love,
Brooks,
Busatto,
Stephens,
&
Gregory
1996a
3­
Young
1963
4­
DeMartini
et
al.
1994
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
Page
1051
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Serranidae
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
L
M
N
O
P
Q
R
S
T
U
V
W
X
Growth
Parameters
(
SE)
k
(
SE)
to
(
SE)

Both
Male
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
69.8
(
3.76)
NG
NG
0.06
(
0.008)
NG
NG
­
3.5
(
0.48)
NG
NG
NG
otoliths/
section
Edge
Analysis
NG
35.13
NG
NG
0.1077
NG
NG
­
6.99
NG
NG
0.67
otoliths/
section
Edge
Analysis
intra/
interreader
comparisons
66.2
(
5.08)
NG
NG
0.08
(
0.014)
NG
NG
2.63
(
0.63)
NG
NG
NG
otoliths/
section
Edge
Analysis
NG
AGE
&
GROWTH
r2
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1052
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Serranidae
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
Y
Z
AA
AB
AC
AD
Age
composition
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
1,4,5
1
31
NF
Dioecious2,3,4,5
NF
Oviparous1,2,3
2,3
(
YOY)
2,9
32
4,5,12
33
6
34
7,10>
30
NF
3>
20
NF
NF
Oviparous3,4
4,8,12
14
5
12
69
to
10
11
12
to
15
5
1
31
NF
Dioecious2
NF
Oviparous1,3
4,5,6,12
24
1­
Young
1963
1­
Emmett
et
al.
1991
1­
Hastings
1989
1­
Emmett
et
al.
1991
2­
Cordes
&
Allen
1997
2­
Read
1992
2­
Emmett
et
al.
1991
2­
Strategic
Assessment
Branch
1990
3­
Cordes
1992
3­
Ono
1992
3­
Strategic
Assessment
Branch
1990
3­
Watson
1996f
4­
Limbaugh
1955
4­
Allen
et
al.
1995
4­
Oda
et
al.
1993
4­
Love
1996
5­
DeMartini
et
al.
1994
5­
Love,
Brooks,
Busatto,
Stephens,
&
Gregory
1996a
5­
Smith
&
Young
1966
6­
Wisenbaker
2000
6­
Hovey
&
Allen
2000
Breeding
Behavior
Protogynous
hermaphrodite;
instances
of
prematurational
sex
change1,4
Both
hermaphroditic
and
dioecious
populations6
possible
protogynous
hermaphrodite4
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1053
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AE
AF
AG
AH
AI
AJ
AK
AL
Parental
Care
Male
vs.
Female
length
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
NF
NF
118
(
2
or
3)
18
(
2
or
3)
NG
NG
NG
215
?
L
(
NG)
16
(
NG)
NG
NG
NG
525*
(
NG)
25
(
NG)
NG
NG
NG
622.5
(
NG)
22.5
(
NG)
25
(
NG)
25
(
NG)
NG
718
(
2)
18
(
2)
22
(
2.5)
22.6
(
3)
otoliths;
section
8
17.8^
(
2)­
unspecified
sex
NG
(
NG)
NG
(
NG)
otoliths;
NG
18
to
23
(
2)­
unspecified
sex
NG
(
NG)
NG
(
NG)
tag
&
recapture/
size
modes
NF
1Male>
Female
4,1018^
(
1.4)
15
(<
1)
18
(
1.4)
15
(
1.5)
otoliths;
whole
NF
NF
NF
325.4
(
3
to
4)
25.4
(
3
to
4)
NG
NG
NG
525*
(
NG)
25
(
NG)
NG
NG
NG
719
(
1)
21
(
1)
21.9
(
2.5)
23.9
(
3)
otoliths;
section
1­
Allen
et
al.
1995
1­
Emmett
et
al.
1991
1­
Emmett
et
al.
1991
2­
Allen
et
al.
1995
2­
Strategic
Assessment
Branch
1990
3­
Quast
1968c
3­
Ono
1992
et
al.
1992
4­
Turner
et
al.
1969
4­
Allen
et
al.
1995
5­
Clark
1933
6­
Collyer
&
Young
1953
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
Male
snout
yellow
during
breeding
season1,3,4
50%
maturity
1st
maturity
males­
black
&
white
with
white
spots
below
dorsal
fins
and
white
chins;
females­
golden
hues
&
yellow
chins2
Sexual
dimorphism
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1054
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10
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
Fertilized
eggs
present
Spawning
females
present
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.

NG
NG
1
s.
California1,3,4,6,7,8,9,10,11,12,13,14,15,16,17
*****
*****
******
*****
*****

NG
NG
NG
NG
NG
NG
Edge
analysis
NG
NG
NG
NA
NA
Edge
analysis
Intra/
interreader
comp.
NF
s.
California2,5,16,17
NG
NG
NF
s.
California1,2,7,8,17
*****
*****

NG
NG
Edge
analysis
NG
1­
Young
1963
1­
Emmett
et
al
1991
2­
Eigenmann
1891
3­
Strategic
Assessment
Branch
1990
4­
Monaco
et
al.
1990
5­
Allen
et
al.
1995
6­
Quast
1968c
REPRODUCTION
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
record
Page
1055
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
AZ
BA
BB
BC
BD
BE
BF
Both
reproductive
states
present
Spawning
Frequency
Size
specific
fecundity
equation
Oct.
Nov.
Dec.
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)

Multiple
(
capable
of
daily)
1,2,3,4,5
NF
181,000
(
30.0
SL,
700
g)
1Batch
F=
0.0000027SL2.93
(
r2=
0.49;
SE=
0.43)

~
32
per
season6
Multiple
(
capable
of
daily)
3
1Batch
F=
0.0389SL2.71
(
r2=
0.33;
SE=
0.22)

Multiple
(
capable
of
daily)
3,4
116,500
(
148
g)
to
185,000
(
2,625
g)
1Batch
F=
0.00000017SL3.17
(
r2=
0.71;
SE=
0.34)

~
41
per
season6
1­
Strategic
Assessment
Branch
1990
1­
DeMartini
1987
1­
Oda
et
al.
1993
2­
Quast
1968c
3­
Oda
et
al.
1993
4­
DeMartini
1987
5­
Young
1963
6­
DeMartini
et
al.
1994
Fecundity
ds
Fecundity
range
in
#
eggs
(@
cm,

grams,
or
age)

Mean
(
range)
fecundity
in
eggs/
g
female
wt.

Page
1056
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BG
BH
BI
BJ
BK
BL
BM
Size
at
settlement
Size
Egg
duration
Larval
duration
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
1,2,3
36
hours
1,3,8
1
month
18
to
11
NF
1,90.94
to
0.97
planktonic2,3,4,5,7,10
NF
4,528
±
3
days
2,3
10.3
(
8.6
to
12.7)
2
0.04
730
±
2.7
days
4
11
4
0.79
to
0.9
(
in
70%
ethanol)

5
10
NF
2
25
to
31
days
NF
1
1,30.8
to
0.89
planktonic1,2,5,6,8,9
NF
4,621
±
3
days
4
0.79
to
0.9
(
in
70%
ethanol)

NF
426
±
2
days
NF
NF
1,30.94
to
0.97
planktonic1,2,5,10
NF
50.85
(
SD=
0.02)

1­
Cordes
&
Allen
1997
1­
Love,
Brooks,
Busatto,
1­
Carr
1994
1­
Allen
et
al.
1995
1­
Butler
et
al
1982
1­
Feder
et
al
1974
2­
Coyer
1979
2­
Hovey
&
Allen
2000
2­
Cordes
&
Allen
1997
2­
Read
1992
2­
Butler
et
al.
1982
3­
Cordes
1992
3­
Pondella
2000
3­
Cordes
1992
3­
Watson
1996f
3­
Emmet
et
al.
1991
4­
Wisenbaker
2000
4­
Butler
et
al.
1982
4­
Oda
et
al.
1993
4­
Strategic
Assessment
Branch
1990
5­
Cordes
&
Allen
1997
5­
Carr
1991b
5­
Gadomski
&
Caddell
1996
5­
Watson
1996f
6­
Tranah
&
Allen
1999
6­
Allen
et
al.
1995
Eggs
DISPERSAL
&
RECRUITMENT
4
0.79
to
0.9
(
in
70%
ethanol)

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1057
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17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BN
BO
BP
BQ
BR
BS
BT
Fish
Size
Fish
Size
(
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
(
SL
mm)

1,2,32.2
to
16.6
mm
pelagic1,2,3,4,5,6
carnivore
diurnal1
NF
1
16.6
to
maturity
(
pelagic
crustaceans)
1
1,32.2
to
12.3
pelagic3
NF
NF
diurnal1
NF
1
12.3
to
maturity
1,2,3
to
13.3
pelagic1,3,6
carnivore
NF
diurnal1
NF
1
13.3
to
maturity
(
pelagic
crustaceans)
1
1­
Butler
et
al.
1982
1­
Emmett
et
al.
1991
1­
Emmett
et
al.
1991
1­
Watson
&
Davis
Jr.
1989
1­
Butler
et
al.
1982
2­
Emmett
et
al.
1991
2­
Strategic
Assessment
Branch
1990
3­
Watson
1996f
3­
Watson
1996f
4­
Holbrook
et
al.
1990
5­
Carr
1991b
6­
Love
1996
Larvae
(
tintinnids;
bivalves
veligers;
copepods)
1
Food
habits
Page
1058
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9
10
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12
13
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15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BU
BV
BW
BX
BY
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
carnivore
foliage­
searcher2
diurnal
1,2,3,4
surfgrass
beds18
seagrass
beds;
lagoons4
carnivore
picker1
NF
comorants3
(
small
fish;
crustaceans;
mollusks)
2
carnivore
NF
NF
surfgrass
beds18
eelgrass
beds
21
1­
Emmet
et
al.
1991
1­
Emmet
et
al.
1991
1­
Ferry
et
al
1997
1­
Coyer
1979
1­
Emmett
et
al.
1991
2­
Strategic
Assessment
Branch
1990
2­
Ferry
et
al.
1997
2­
Coyer
1979
2­
Cordes
1992
2­
Carr
1994
3­
Hartney
1996
3­
Barraclough
1967
3­
Carr
1991b
3­
Fitch
&
Lavenberg
1975
4­
Rosales­
Casian
1997
4­
Valle
et
al.
1999
4­
Love
1996
4­
Carr
1991b
5­
Turner
et
al.
1969
5­
Coyer
1979
6­
Valle
et
al.
1999
6­
Robert
et
al.
1984
Life
Stage
Juveniles
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
giant
kelpfish;
kelp
bass;

possibly
island
kelpfish,

spotted
kelpfish,
scorpionfish,

treefish,
and
kelp
and
grass
rockfish4
(
crustaceans,
such
as
calenoid
and
harpacticoid
copepods
and
mysids;

larvaceans;
polychaetes;
coelenterates;

small
fish;
algae)
1,3,5,7,8,9
eelgrass
beds
and
boa
kelp;
kelp
canopy
(
YOY
essential
habitat);
algae
holdfasts;

surf
zone;
lagoons;
drifting
kelp
1,2,3,4,7,8,9,10,11,12,13,14,16,19,20
kelp
bass;
giant
sea
bass;

grouper1,2
bay
mouths;
eelgrass
beds;
estuaries
and
lagoons;
breakwaters;
kelp
beds1,4,5,6,7,15,17
kelp
bass;
barred
sand
bass;

giant
sea
bass;
grouper1
(
demersal
crustaceans;
cephalopods;

polychaetes;
ophiuroids;
fish,
such
as
anchovy
and
surfperch)
1,4,6,9
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
BZ
CA
CB
CC
Adults
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
(
meso)
carnivore20/
piscivore24
ambusher2,3
diurnal2,3,5,6
nocturnal?
6,9,10
diurnal/
crepuscular
8
shelter
in
holes
and
crevices
at
night
24
dawn/
dusk11
scavange
sea
urchins
form
sheephead
attacks
22
carnivore34
picker1
diurnal
4,7
ambusher3
carnivore
ambusher3
nocturnal2,10
1­
Feder
et
al
1974
1­
Feder
et
al
1974
1­
Ferry
et
al.
1997
1­
Ebling
&
Bray
1976
2­
Emmet
et
al.
1991
2­
Emmet
et
al.
1991
2­
Coyer
1979
2­
Emmett
et
al.
1991
3­
Strategic
Assessment
Branch
1990
3­
Strategic
Assessment
Branch
1990
3­
Hobson
1994
3­
Strategic
Assessment
Branch
1990
4­
Allen
1985
4
Ferry
et
al.
1997
4­
Ono
1992
et
al.
1992
5­
Allen
et
al.
1995
5­
Love
&
Ebeling
1978
5­
Hartney
1996
6­
Quast
1968c
6­
Allen
et
al.
1995
6­
Hobson
&
Chess
1986
rocky
bottoms;
kelp
holdfasts
and
beds;
eelgrass
and
surfgrass
beds;
not
greatly
influenced
by
removal
of
kelp
canopy;
mud/
sand
bottoms;

artificial
reefs;
areas
of
high
relief
2,3,4,6,10,11,12,13,14,15,16,17,18,19,22,23,25,26,27,28
(
fish,
such
as
other
kelp
bass,
pipefishes,
flatfishes,

pomacentrids
and
surfperches;
cephalopods;

crustaceans,
such
as
crabs,
euphasiids,
and
caridians;

polychaetes;
kelp;
tunicates;
hydrozoans)

2,3,5,7,8,9,10,12,13,15,17,19,20,21,23,27,28,29,30
sand
bottoms
near
rocks
or
eelgrass;
sand
depressions;
bays
and
estuaries;
surfgrass
1,4,5,7,8,9,19,23,27,28
(
demersal
crustaceans,
especially
crabs;
small
fishes,

including
kelpfish;
ophiuroids;
mollusks,
especially
bivalves;
polychaetes)
1,4,6,14,16,20,25,30
diurnal,
with
occasional
nighttime
feeding
bouts1
(
small
fishes,
such
as
surfperch;
crustaceans;

mollusks;
ophiuroids;
cephalopods)
1,9,10,11,12,18,26,27,30
demersal;
among
rocks
near
or
on
sand
or
sandy
flats;
kelp
and
surfgrass
beds;
rocky
and
artificial
reefs
1,2,4,10,11,12,14,18,19,20,21,23,25,26,27,28
Page
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9
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CD
CE
CF
CG
CH
CI
CJ
CK
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)

none1,2
NF
NF
NF
NF
1NG
0.019
3,4NG
0.012
moray
eels5
birds2
NF
NF
NF
NF
2NG
0.085
(
±
0.012)

2NG
0.09
(
±
0.011)

NF
NF
NF
NF
NF
NF
NF
NF
1­
Emmett
et
al.
1991
1­
Emmett
et
al.
1991
1­
Beckwitt
1983
2­
Hubbs
et
al.
1970
2­
Beckwitt
1983
2­
Stepien
1995
3­
Lowe
et
al.
2000
3­
Smith
1967
3­
Waples
&
Rosenblatt
1987
4­
Bray
&
Hixon
1978
4­
Tranah
&
Allen
1999
4­
Waples
1987
5­
Feder
et
al.
1974
5­
Stepian
1995
6­
Rosenblatt
&
Graham
199
southern
CA
mainland
and
Santa
Catalina
I.
3
kelp
bass;
giant
sea
bass;

sea
lions;
electric
ray1,3,4
GENETICS
Gulf
of
California;
southern
Pacific
Baja;
southern
California4
Gulf
of
California;
outer
Baja
coast
and
southern
California5
Nucleotide
diversity
(#)
Mean
heterozygosity
None
found
between
San
Diego
and
Gulf
of
California
using
allozymes6
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CL
CM
CN
CO
CP
CQ
CR
CS
CT
CU
CV
RECRUITMENT
INTO
FISHERY
Mortali
First
Recruitment
Full
Recruitment
Approach
Effective
population
size
Total
length
(
age)
Total
length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)

allozymes
NF
NF
NF
NF
NF
NF
NF
1NG
NG
0.544
allozymes
1
0.287
NG
NG
allozymes
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Young
1963
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
CW
CX
CY
CZ
DA
DB
DC
ty
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Interactions
/
associations
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
catch
curve
3
to
12
NG
NG
competes
with
barred
sand
bass1
1<
16
km
catch
curve
7
to
13
NG
NG
2<
36
km
3<
12
miles
4<
17
miles
6no
significant
movements
only
YOY
closely
associated
with
Macrocystis
8,9,10
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
competes
with
kelp
bass1
7<
65
km
8
15
to
78
km
high
levels
of
DDT
found
in
southern
Californian
fish6
1­
Emmett
et
al.
1991
1­
Limbaugh
1955
1­
Emmett
et
al.
1991
2­
Hobson
1971
2­
Strategic
Assessment
Branch
1990
3­
Cross
&
Hose
19????
3­
Quast
1968c
4­
Thomson
&
Lehner
1976
4­
Collyer
&
Young
1953
5­
Dailey
et
al.
1981
5­
Young
1963
6­
McCain
et
al.
1992
6­
Limbaugh
1955
infected
by
ectoparasite
from
the
families
Bomolochidae,
Caligidae,
and
Dichelesthiidae2
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
5
reproductive
success
decreased
by
contaminants
introduced
by
humans3
infected
by
parasites
from
the
families
Bomolochidae,

Caligidae,
Hartschekiidae,
Lernaeopodidae,

Pennellidae,
Opecoelidae,
Cymothoidae,
Anisakida,

and
Ophryoglenidae11
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants11
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants11
maybe
a
keystone
species
in
the
northern
Gulf
of
California4
infested
by
roundworms
from
genera
Anisakis
and
Phocanema
5
may
be
negatively
affected
by
chlorine
discharge
plums
from
power
plants11
infected
by
parasites
from
the
families
Bomolochidae,

Caligidae,
Hatschekiidae,
Lernaeopodidae,

Pennellidae,
Hemiuridae,
Monorchiidae,
Opecoelidae,

Cymothoidae,
Macroval
Vitrematidae,
Anisakidae,
and
Spiruridae11
infected
by
parasites
from
the
families
Bomolochidae,

Caligidae,
Hatschekiidae,
Lernaeopodidae,

Pseudocycnidae,
Hemiuridae,
Opecoelidae,

Capsalidae,
and
Diplectanidae11
MORTALITY
cleaned
by
Oxyjulis
californica
and
Brachyistius
frenatus
1,2
5,9up
to
282
miles,
but
most
<
0.1
miles
infected
by
ectoparasite
from
the
families
Bomolochidae
and
Caligidae2
Page
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12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
DD
DE
DF
DG
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
Residential
1,2,3,5,6,8,10,11,13
No5
Yes;
spawning
aggregations1,3,4
No1,2,4
1(
40
ha)
Rarely2
Possibly3
2(
700
m)
Mostly
solitary5
3(
1175
ft.)
solitary8
Residential
7,9
NF
NF
NF
Residential
1,4,5,12,13
NF
Yes3
NF
1­
Emmett
et
al.
1991
1­
Emmett
et
al.
1991
1­
Ebling
&
Bray
1976
1­
Strategic
Assessment
Branch
1990
2­
Miller
&
Geibel
1973
2­
Strategic
Assessment
Branch
1990
2­
Strategic
Assessment
Branch
1990
2­
Quast
1968c
3­
Quast
1968c
3­
Quast
1968c
3­
Turner
et
al.
1969
3­
Lowe
et
al.
2000
4­
Turner
et
al.
1969
4­
Lowe
et
al.
2000
4­
Love,
Brooks,
&
Ally
1996
4­
Limbaugh
1955
5­
Johnson
et
al.
1994
5­
Young
1963
5­
Coyer
1979
6­
Larson
&
DeMartini
1984
6­
Cordes
1992
Movement
Solitary,
but
aggragates
during
spawning
period6,7,9
BEHAVIOR
4(
29
to
24553
m2
[
90%
home
range])
Page
1064
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­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
A
B
C
D
E
Page
1065
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1167
5­
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Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
F
G
H
I
JK
Page
1066
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
L
M
N
O
P
Q
R
S
T
U
V
W
X
Page
1067
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
Y
Z
AA
AB
AC
AD
7­
Cordes
&
Allen
1997
8­
Tranah
&
Allen
1999
9­
Coyer
1963
10­
Cordes
1992
11­
Fitch
and
Lavenberg
1975
12­
Love
1996
Page
1068
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
AE
AF
AG
AH
AI
AJ
AK
AL
7­
Love,
Brooks,
Busatto,
Stephens,
&
Gregory
1996
8­
Coyer
1979
9­
Limbaugh
1955
10­
Love
1996
Page
1069
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
7­
Clark
1933
8­
Collyer
&
Young
1953
9­
Oda
et
al.
1993
10­
Smith
&
Young
1966
11­
Turner
et
al.
1969
12­
Cordes
&
Allen
1997
13­
Love,
Brooks,
&
Ally
1996
14­
Coyer
1979
15­
Cordes
1992
16­
Limbaugh
1955
17­
Love
1996
Page
1070
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
AZ
BA
BB
BC
BD
BE
BF
Page
1071
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
BG
BH
BI
BJ
BK
BL
BM
7­
Cordes
1992
7­
Coyer
1979
8­
Love
1996
8­
Limbaugh
1955
9­
Fitch
&
Laveneberg
1975
10­
Love
1996
Page
1072
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
BN
BO
BP
BQ
BR
BS
BT
Page
1073
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
BU
BV
BW
BX
BY
7­
Larson
&
DeMartini
1984
7­
Carr
1991b
8­
Holbrook,
Carr
et
al.
1990
8­
Hobson
1994
9­
Carr
1989
9­
Love
1996
10­
Carr
1994
11­
Mitchell
&
Hunter
1970
12­
Coyer
1979
13­
Cordes
1992
14­
Limbaugh
1955
15­
Phillips
et
al.
1997
16­
Pondella
&
Stephens
1994
17­
CDF
&
G
1982
18­
DeMartini
1981
19­
Carr
1991b
20­
Love
1996
21­
Hoffman
1986
Page
1074
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
BZ
CA
CB
CC
7­
Hovey
&
Allen
2000
7­
Quast
1968a
7­
Thomson
&
Lehner
1976
8­
Rodriguez­
Romero
et
al.
1998
8­
Young
1963
8­
Holbrook
&
Schmitt
1988
9­
Rosales­
Casian
1997
9­
Quast
1968d
9­
Coyer
1979
10­
Carlisle
et
al.
1964
10­
Carlisle
et
al.
1964
10­
Helvey
1985
11­
Turner
et
al.
1969
11­
Turner
et
al.
1969
11­
Love
1996
12­
Johnson
et
al.
1994
12­
Johnson
et
al.
1994
13­
Valle
et
al.
1999
13­
Hobson
&
Chess
1986
14­
Larson
&
DeMartini
1984
14­
Thomson
&
Lehner
1976
15­
Hobson
&
Chess
1986
15­
Holbrook
&
Schmitt
1988
16­
Carr
1989
16­
Galvan­
Magana
et
al.
2000
17­
Coyer
1979
17­
Coyer
1979
18­
Ebeling,
Larson,
&
Alevizon
1980
18­
Roberts
et
al.
1984
19­
Limbaugh
1955
19­
Cordes
1992
20­
Phillips
et
al.
1997
20­
Limbaugh
1955
21­
Love,
Stephens,
et
al.
1996
21­
Diaz
&
Hammann
1987
22­
Stephens
et
al.
1984
22­
Nelson
&
Vance
1979
23­
CDF
&
G
1982
23­
Limbaugh
1964
24­
Ebeling
&
Bray
1976
24­
Davis
&
Anderson
1989
25­
DeMartini
1981
25­
Fitch
&
Lavenberg
1975
26­
Winget
1968
26­
McCain
et
al.
1992
27­
Love
et
al.
1986
27­
Holbrook
et
al.
1997
28­
Love
1996
28­
Winget
1968
29­
Hobson
1994
30­
Love
1996
Page
1075
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
CD
CE
CF
CG
CH
CI
CJ
CK
Page
1076
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
CL
CM
CN
CO
CP
CQ
CR
CS
CT
CU
CV
Page
1077
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
CW
CX
CY
CZ
DA
DB
DC
7­
Hose
et
al.
1983
7­
Phillips
et
al.
1997
8­
Holbrook
et
al.
1990
8­
CDF
&
G
1982
9­
Larson
&
DeMartini
1984
9­
Love
1996
10­
Carr
1989
11­
Love
&
Moser
1983
Page
1078
of
1167
5­
2220e
­
Serranidae
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
DD
DE
DF
DG
7­
Thomson
&
Lehner
1976
7­
Limbaugh
1955
8­
Carr
1994
8­
Davis
&
Anderson
1989
9­
Tranah
&
Allen
1999
9­
Love
1996
10­
Coyer
1979
11­
Cordes
1992
12­
CDF
&
G
1982
13­
Helvey
1985
Page
1079
of
1167
5­
2220e
­
Serranidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
A
B
C
D
E
F
Length
weight
equation
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Sphyraenidae
Sphyraena
argentea
1
1surface
to
38
1Both:
W=
0.003962TL2.983
1­
Robins
et
al
1991
1­
Eschmeyer
1998
1­
Robins
et
al.
1991
1­
Love
et
al
2000
1­
Love
et
al
2000
1­
Walford
1932
2­
Gates
&
Frey
1974
TAXONOMY
RANGE
California
barracuda;
Pacific
barracuda;

scoots;
scooters;
snake;
log
barracuda;

cuda1,2
Alaska
(
Kodiak
I.)
to
s.
Baja
CA
(
Cabo
San
Lucas)
and
Gulf
of
CA1
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
Page
1080
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
G
H
I
J
K
L
M
N
O
P
Q
R
S
Growth
Parameters
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)
to
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
Both
Male
Female
Both
Male
NF
1
122
s.
California
(
TL)
1
NG
NG
102.25
(
1.71)
NG
NG
0.249
(
0.0136)
NG
NG
­
0.7689
(
0.941)
NG
1­
Love
et
al
2000
1­
Pinkas
1966
AGE
&
GROWTH
Page
1081
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
T
U
V
W
X
Y
Z
AA
Age
composition
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)
information
NG
NG
scales
NO
intrareader
comparison
1
1
12
1
2,3,4
11
1­
Walford
1932
1­
Strategic
Assessment
Branch
1990
1­
Pinkas
1966
2­
Ally
&
Miller
1992
3­
Walford
1932
4­
Fitch
&
Lavenberg
1971
5­
Pinkas
1960a
r2
Page
1082
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
AB
AC
AD
AE
AF
AG
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
M
vs.
F
length
Secondary
Characteristics
Dioecious1
NF
Oviparous1,2
NF
1Males<
Females
NF
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
1­
Fitch
&
Lavenberg
1971
2­
Garrison
&
Miller
1982
Breeding
Behavior
Sexual
dimorphism
Page
1083
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
AH
AI
AJ
AK
AL
AM
AN
AO
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.

1,2,3NG
(
2)
NG
(
2)
NG
(
2)
NG
(
2)
NF
NF
NF
NF
4,533
(>
1)
44
(>
1)
43
(~
2)
47
(~
2)
scales
Edge
Analysis
No
1­
Strategic
Assessment
Branch
1990
2­
Hart
1973
3­
Ally
&
Miller
1992
4­
Walford
1932
5­
Garrison
&
Miller
1982
1st
maturity
REP
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
50%
maturity
Page
1084
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1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
states
present
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.

California2,3,4,5,6,7,8,9,10,11
*****
*****
*****

s.
California1
1­
Eigenmann
1891
2­
Strategic
Assessment
Branch
1990
3­
Hart
1973
4­
Walford
1932
5­
Monaco
et
al.
1990
6­
Fitch
&
Lavenberg
1971
7­
Sandkop
&
Watson
1996
8­
Breder
&
Rosen
1966
9­
Garrison
&
Miller
1982
10­
Kramer
&
Smith
1973
11­
Pinkas
1960a
PRODUCTION
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
Page
1085
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
BC
BD
BE
BF
Spawning
Frequency
Size
specific
fecundity
equation
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)

Multiple1,2,3,4,5
NF
NF
250,000
(
2
yrs.)
to
400,000
(
6
yrs.)

651,000
(
NG)
to
484,000
(
NG)

1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
2­
Hart
1973
2­
Ally
&
Miller
1992
3­
Walford
1932
3­
Walford
1932
4­
Garrison
&
Miller
1982
4­
Walford
1932
5­
Pinkas
1960a
5­
Fitch
&
Lavenberg
1971
6­
Garrison
&
Miller
1982
7­
Pinkas
1960a
751,000
(
50
?
L;
2
yrs)
to
340,000
(
91
?
L:
7
yrs.)

Fecundity
1,3,4,5
42000
(
50
TL;
2yrs.)
to
484,000
(
94
TL;
6
yrs.)

Fecundity
range
in
#
eggs
(@
cm,
grams,
or
age)

Mean
(
range)
fecundity
in
eggs/
g
female
wt.
Page
1086
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
BG
BH
BI
BJ
BK
BL
BM
Size
at
settlement
Size
Egg
duration
Larval
duration
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
couple
of
days1,2
NF
NF
NF
1,21.0
to
1.6
planktonic1,2,3,4,5,6
NF
3,5,7
1.24
to
1.6
41.0
to
1.4
6
1.02
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
2­
Fitch
&
Lavenberg
1971
2­
Hart
1973
2­
Ally
&
Miller
1992
3­
Walford
1932
3­
Sandknop
&
Watson
1996
4­
Sandknop
&
Watson
1996
4­
Limbaugh
1955
5­
Fitch
&
Lavenberg
1971
5­
Garrison
&
Miller
1982
6­
Breder
&
Rosen
1966
6­
Feder
et
al.
1974
7­
Garrison
&
Miller
1982
DISPERSAL
&
RECRUITMENT
Eggs
Page
1087
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
BN
BO
BP
BQ
BR
BS
Fish
Size
(
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
12.2
to
12
pelagic1,2,3
carnivore
NF
NF
NF
22.3
to
30.8
(
pelagic
crustaceans)
1
32.5
to
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
2­
Sandknop
&
Watson
1996
3­
Ally
&
Miller
1992
3­
Garrison
&
Miller
1982
3­
Garrison
&
Miller
1982
Larvae
Food
habits
Page
1088
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
BT
BU
BV
BW
BX
BY
Fish
Size
Food
habits
(
SL
mm)
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
1
12
to
maturity
carnivore
NF
NF
NF
230.8
to
maturity
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
2­
Sandknop
&
Watson
1996
2­
Feder
et
al.
1974
(
larval
and
juvenile
fishes;
planktonic
crustaceans)
1
in
water
column
over
sandy
areas;

protected
bays
near
rocks1,2
Juveniles
Life
Stage
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Page
1089
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
BZ
CA
CB
CC
Adults
Food
habits
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
carnivore
ambusher1
NF
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
1­
Strategic
Assessment
Branch
1990
2­
Allen
1985
2­
Schultz
1983
3­
Carlisle
et
al.
1964
3­
Quast
1968d
4­
Turner
et
al.
1969
4­
Fitch
&
Lavenberg
1971
5­
Larson
&
DeMartini
1984
5­
Limbaugh
1955
6­
Limbaugh
1955
6­
Feder
et
al.
1974
7­
CDF
&
G
1982
8­
Follett
1976
9­
Love
et
al.
1996
10­
Feder
et
al.
1974
(
fish,
such
as
anchovy
and
sardine;
squid)

1,2,3,4,5,6
neritic
nearshore;
artificial
reefs;
kelp
beds;

bays;
over
soft
substrata
1,2,3,4,5,6,7,8,9,10
Page
1090
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
CD
CE
CF
CG
CH
CI
CJ
CK
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)

None1,2
NF
NF
NF
NF
NF
NF
1­
Strategic
Assessment
Branch
1990
1­
Ally
&
Miller
1992
2­
Hart
1973
2­
Pinkas
1966
3­
Schultz
1983
4­
Fitch
&
Lavenberg
1971
pinnipeds;
porpoises;
large
fish,
such
as
giant
sea
bass
1,2,3,4
Nucleotide
diversity
(#)
Mean
heterozygosity
GENETICS
Page
1091
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1167
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2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
CL
CM
CN
CO
CP
CQ
CR
CS
CT
CU
RECRUITMENT
INTO
FISHERY
Mortal
First
Recruitment
Full
Recruitment
Approach
Effective
population
size
Total
length
(
age)
Total
length
(
age)
Sampling
gear
Ageing
structure
/
Method
Validation
Verification
Natural
(
M)
Fishing
(
F)

NF
NF
1NG
(
2)
NG
(
5)
hook
&
line;
gill
net
scales
No
intrareader
comp.
1
0.18338
0.539
to
0.568
1
NG
NG
1
NG
NG
1
NG
NG
1­
Pinkas
1966
1­
Pinkas
1966
Page
1092
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
CV
CW
CX
CY
CZ
DA
DB
DC
DD
DE
ity
Estimates
Rate
of
Exploitation
Method
of
ge
range
use
Rate
of
Exploitation
Interactions
/
associations
Total
(
Z)
mortality
estimation
alculate
mort
exploitation
(
u)
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)

0.723
to
0.751
Beverton
and
Holt
5
to
9
NG
0.7558
NF
carry
parasitic
worms1
1to
340
miles
Transient1
NF
0.726
to
0.764
Chapman­
Robson
5
to
9
NG
NG
0.62
to
0.976
Chapman­
Robson
5
to
9
NG
NG
0.667
to
0.794
Chapman­
Robson
5
to
9
NG
NG
1­
Fitch
&
Lavenberg
1971
1­
Pinkas
1966
1­
Ally
&
Miller
1992
2­
Love
&
Moser
1983
infested
with
parastites
from
the
families
Dasyrhynchidae,
Trypanorhyncga,

Bomolochidae,
Caligidae,
and
Axinidae2
BEHAVIOR
Movement
MORTALITY
Page
1093
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
DF
DG
Schooling
Territoriality
Yes1,2,3,4,5
NF
1­
Strategic
Assessment
Branch
1990
2­
Fitch
&
Lavenberg
1971
3­
CDFG
1982
4­
Follett
1976
5­
Feder
et
al.
1974
Page
1094
of
1167
5­
2220e
­
Sphyraenidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Stichaeidae1,2
Cebidichthys
violaceus
1,2
1intertidal
to
24
Cebidichthyidae
(
for
Cebidichthys
only)
3
Xiphister
mucosus
1,2
1intertidal
to
18
X.
mucosum
3
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Robins
et
al.
1991
1­
Love
et
al.
2000
1­
Love
et
al.
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
&
Frey
1974
3­
Eschmeyer
et
al.
1983
3­
Marliave
&
DeMartini
1977
TAXONOMY
RANGE
rock
prickleback;
blenny­
eel;
rock
blenny;

rock­
eel;
clack
eel;
eel;
slick
eel
1,2
monkeyface
prickleback;
monkeyface­
eel;

bleeny
eel;
monkeyface
blenny;
eel;
giant
monkeyface­
eel;
California
monkeyfaceeel1,2
southern
Oregon
to
northern
Baja
California
(
Bahia
San
Quintin)
1
southeastern
Alaska
(
Port
San
Juan)
to
central
California
(
Pt.

Arguello),
possibly
southern
California
(
Santa
Cruz
I.)
1
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42
43
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45
46
47
48
49
50
51
52
53
54
F
G
H
I
JKLMN
Length
weight
equation
Growth
Paramete
Max.
reported
Geographic
area
Linf
in
cm
(
SE)
k
(
SE)

Length
conversion
equations
length
(
cm)
(
length
measurement)
Male
Female
Both
Male
Female
1*
Male:
W=
0.006797SL3.072
NF
1
76
northern
California
(
SL)
1
70
(
5.0)
62
(
2)
71
(
8)
0.12
(
0.02)
0.14
(
0.02)

1*
Female:
W=
0.0325SL2.654
northern
California
(
SL)
1
NG
NG
72
(
8)
NG
NG
1*
Both:
W=
0.01289SL2.9
2*
Both:
W=
0.00965SL2.971
NF
1
58.6?
NF
NF
NF
NF
NF
NF
1­
Marshall
&
Wyllie
Echeverria
1992
1­
Love
et
al.
2000
1­
Marshall
&
Wyllie
Echeverria
1992
2­
Edwards
1981
3­
Horn
&
Riegle
1981
Length
(
TL=
cm
&
FL,
SL=
mm)
to
weight
(
g)

equation.
*
denotes
TL
=
mm
or
FL,
SL=
cm
3*
Both:
W=
0.007SL2.95
(
based
on
sample
set
of
29
C.
violaceus
and
19
A.
purpurescens
)

3*
Both:
W=
0.0042SL2.95
(
based
on
sample
set
of
28
X.
mucosus
and
29
X.
atropurpureus
)
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22
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24
25
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
O
P
Q
R
S
T
U
V
W
X
Y
ers
to
(
SE)

Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.

0.1
(
0.04)
­
1.91
(
0.29)
­
1.95
(
0.28)
­
2.63
(
1.31)
NG
NG
NG
operculum
No
comparison
to
otoliths
0.1
(
0.03)
NG
NG
­
1.89
(
1.08)
NG
NG
NG
otoliths;
whole
No
comparison
to
opercula
2,4
3­
Juveniles
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Bray
et
al.
1986
2­
Fris
&
Horn
1993
3­
Helm
1990
4­
Tenera
Environmental
2000
AGE
&
GROWTH
r2
1­
have
relatively
slower
growth
rates
than
other
fishes
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21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
Z
AA
AB
AC
AD
AE
Age
composition
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
1,3,4
18
NF
Dioecious
NF
Oviparous
nest
guarding
(
females?)
1,7
2,4
11
NF
Dioecious
promiscuous?
1
Oviparous1,2,3
males
nest
guard2,3,4
1­
Marshall
&
Wyllie
Echeverria
1992
1­
Marliave
&
DeMartini
1977
1­
Garrison
&
Miller
1982
1­
Fitch
&
Lavenberg
1971
2­
Fitch
&
Lavenberg
1975
2­
Matarese
et
al.
1989
2­
Martin
1995
3­
Tenera
Environmental
Services
2000
3­
Love
1996
3­
Garrison
&
Miller
1982
4­
Love
1986
4­
Matarese
et
al.
1989
5­
Marliave
&
DeMartini
1977
6­
Gibson
1982
7­
Tenera
Environmental
Services
2000
Breeding
Behavior
males
nest
guard,
wrapping
their
bodies
around
the
egg
mass5,6
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
AF
AG
AH
AI
AJ
AK
AL
AM
AN
M
vs.
F
length
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
NF
NF
1,5
NG
(
3
to
4)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NG
NG
NG
2
28
to
32.5
(
NG)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NA
NA
NA
3,5,6
36
(
4)­
sex
unspecified
39
(
5)­
sex
unspecified
operculum
No
comparison
to
otoliths
1Males<
Females
NF
4,6
28
to
30
(
5)­
sex
unspecified
NG
(
NG)
NG
(
NG)
NG
NG
NG
1­
Fitch
&
Lavenberg
1975
1­
Fitch
&
Lavenberg
1971
2­
Ralston
&
Horn
1986
3­
Marshall
&
Wyllie
Echeverria
1992
4­
Garrison
&
Miller
1982
5­
Tenera
Environmental
Services
2000
6­
Love
1996
TL,
FL*,
or
SL^
in
cm
(
age
in
yrs)
at
1st
maturity
50%
maturity
Sexual
dimorphism
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29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
Fertilized
eggs
present
Spawning
females
present
Both
reproductive
states
present
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.

NF
northern
California1,3,4
*****
*****
*****

NF
California2,4
In
the
winterGibson
1982
1­
Marshall
&
Wyllie
Echeverria
1992
2­
Kelly
et
al.
1985
3­
Tenera
Environmental
Serviecs
2000
4­
Love
1996
REPRODUCTION
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
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20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BC
BD
BE
BF
BG
Spawning
Frequency
Size
specific
fecundity
equation
(
per
year)
(
TL=
cm,
SL=
mm;
*
denotes
TL=
mm;
SL=
cm)
Egg
duration
NF
1,5
NG
(
possibly
6,000
to
8,000)
NF
NF
Once1
2,4
7,276
±
2,553
(
2,000
to
15,000)
NF
NF
NF
3,6
NG
(
5,500
to
9,500)

1­
Garrison
&
Miller
1982
1­
Fitch
&
Lavenberg
1971
1­
Marshall
&
Wyllie
Echeverria
1992
2­
Marliave
&
DeMartini
1977
2­
Garrison
&
Miller
1982
2­
Tenera
Environemental
Services
2000
3­
Matarese
et
al.
1989
4­
Love
1996
4­
Marliave
&
DeMartini
1977
5­
Tenera
Environemental
Services
2000
6­
Love
1996
Fecundity
Fecundity
range
in
#
eggs
(@
cm,
grams,
or
age)

Mean
(
range)
fecundity
in
eggs/
g
female
wt.

Males­
multiple;
females­
once2
1,2,4
17,500
(
41
SL;
7
yrs.)
to
46,000
(
61
SL;
11yrs.)

Page
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31
32
33
34
35
36
37
38
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40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BH
BI
BJ
BK
BL
BM
BN
Size
at
settlement
Size
Fish
Size
Larval
duration
(
SL
in
mm)
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
(
mm)

NF
1
17
to
22
TL
NF
NF
NF
NF
2into
tidepools:
65
SL
NF
1
17
to
22
TL
NF
1
2.5
amphipods1
1
11
to
2
9.5
to
1­
Setran
&
Behrens
1993
1­
Matarese
et
al.
1989
1­
Fitch
&
Lavenberg
1971
1­
Marliave
&
DeMartini
1977
1­
Garrison
&
Miller
1982
2­
Helm
1990
2­
Garrison
&
Miller
1982
2­
Matarese
et
al.
1989
3­
Marliave
&
DeMartini
1977
4­
Tenera
Environmental
Services
2000
5­
Love
1996
DISPERSAL
&
RECRUITMENT
Eggs
demersal,
adhering
to
rocks1,4
demersal,
adhering
to
rocks2,3,5
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24
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28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BO
BP
BQ
BR
BS
BT
BU
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
(
SL
mm)
Habitat
NF
NF
NF
NF
NF
1,2
17
to
maturity
pelagic1
NF
NF
NF
NF
1,2
17
to
maturity
high
rocky
intertidal3,9,10
lower
intertidal
12
1­
Garrison
&
Miller
1982
1­
Setran
&
Behrens
1993
1­
Fitch
&
Lavenberg
1971
2­
Setran
1990
2­
Setran
&
Behrens
1993
3­
Barton
1982
4­
Horn
&
Riegle
1981
5­
Edwards
&
Cech
1990
6­
Setran
1990
7­
Urquhart
1984
8­
Yoshiyama
et
al.
1986
9­
Cross
1981
10­
Kelly
et
al.
1985
11­
Tenera
Environemental
Services
2000
12­
Wourms
and
Evans
1974
Li
young
juveniles:
gravel
bottoms;
older
juveniles
(
32
to
42
mm
TL):
pebble
bottoms2,6
kelp
beds;
high
rocky
intertidal;
tide
pools
1,3,5,7,8,10,11
(
juveniles
of
this
species
especially
adapted
for
high
intertidal)
4
HABITAT
ASSOCIATIO
Food
habits
Larvae
young
juveniles:
gravel
bottoms;
older
juveniles
(
32
to
42
mm
TL):
pebble
bottoms2,6
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22
23
24
25
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27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
BV
BW
BX
BY
BZ
Food
habits
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Habitat
1,2,5,6,7,9<
45
to
47
mm
SL/
4<
80
mm
TL:
carnivore
NF
during
flooding
tides1
possibly
Atedius
lateralis
1
prior
to
ebbing
tides2
crevice,
coiled
among
egg
mass
29
1,2,3,5,6,9>
45
to
47
mm
SL/
4>
80
mm
TL:
herbivore
(
green
and
red
macroalgae)
1,2,3,4,6,9
750
mm<
body
length<
70
mm:
omnivore
2,5<
47
mm
SL:
carnivore
NF
prior
to
ebbing
tides2
NF
spawning
individuals
found
subtidally24
deep
lower
intertidal
28
2,5,6>
47
mm
SL:
herbivore
(
red
and
green
macroalgae)
2,6
1­
Ralston
&
Horn
1986
1­
Ralston
&
Horn
1986
1­
Yoshiyama
1981
1­
Fitch
&
Lavenberg
1971
2­
Setrans
&
Behrens
1993
2­
Setran
&
Behrens
1993
2­
Love
1996
2­
Setrans
&
Behrens
1993
3­
Edwards
&
Horn
1982
3­
Helm
1990
3­
Marshall
&
Wyllie
Echeverria
1992
4­
Barton
1982
4­
Edwards
&
Horn
1982
5­
Horn
et
al.
1982
5­
Horn
&
Neighbors
1984
6­
Horn
et
al.
1986
6­
Barton
1982
7­
Montgomery
1977
7­
Horn
&
Riegle
1981
8­
Helm
1990
8­
Fris
&
Horn
1993
9­
Setrans
1990
9­
Boettcher
&
Targett
1993
10­
Urquhart
1984
10­
Edwards
&
Cech
1990
11­
Cross
1981
11­
Martin
1995
12­
Love
1996
12­
Fitch
&
Lavenberg
1975
13­
Burge
&
Schultz
1973
14­
Garrison
&
Miller
1982
15­
Martin
1993
16­
Norris
1963
17­
Setran
1990
18­
Riegle
1977
fe
Stage
switch
from
winter
carnivorous
to
summer
herbivorous
diet8
10<
45
to
60
mm
SL­
carnivores;
>
45
to
60mm
SL­

herbivorous.
When
herivorous,
eat
primarily
red
and
green
macroalgae
(
Ulva
&
Porphyra
)

(
zooplankton,
such
as
calanoids,
harpacticoid,

gammarids,
isopods,
juvenile
mysids,
and
juvenile
spionid
polychaetes)
1,2,4,7,8,9,12
great
egrets
and
great
blue
herons2
Piscivorous
birds,
such
as
great
egrets
and
redbreasted
merganser3
Juveniles
N
&
TROPHIC
INTERACTIONS
11omnivorous:
gradual
switch
from
primarily
carnivorous
to
herbivorous
diet
after
60
mm
TL
breakwaters
and
rocky
areas;
high
and
low
intertidal;
out
of
water,
under
rocks
and
algae1,2,3,4,5,6,7,8,10,11,13,15,16,17,18,19,20,21,22,24,25,27
switch
to
substrate­
dwelling,
larger
prey
as
body
size
and
mouth
gape
increases2
switch
to
substrate­
dwelling,
larger
prey
as
body
size
and
mouth
gape
increases2
(
zooplankton,
such
as
calanoids,
harpacticoid,

gammarids,
juvenile
mysids,
and
juvenile
spionid
polychaetes)
2,11,12
high
and
low
intertidal
(
mostly
low);
out
of
water,

under
rocks
and
algae;
cobble
beaches
2,6,7,9,11,12,14,15,17,18,20,21,22,23,24,26,27
(
small
crustaceans;
Ulva
and
other
green
algae;

red
algae)
7
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22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CA
CB
CC
CD
CE
CF
CG
Adults
Food
habits
Haplotype
diversity
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Substocks
(
Geographic
Area)

omnivore
browser1
NF
NF
NF
NF
(
algae;
small
crustaceans)
2
herbivore19,20,21,22,23,26,27
grass
rockfish;
cabezon3
herbivore17,18,21,23,25,26
browser1
ebbing
tides1
possibly
river
otters1
NF
NF
NF
grass
rockfish;
cabezone3
annual
macrophytes
preferred
over
perennials11,21
1­
Hart
1973
1­
Horn
1989
1­
Miller
&
Marshall
1987
1­
Stevens
et
al.
1984
2­
Fitch
&
Lavenberg
1971
2­
Helm
1990
3­
Bray
et
al.
1986
3­
Jones
1981
4­
Ralston
&
Horn
1986
5­
Setran
&
Behrens
1993
6­
Neighbors
&
Horn
1991
7­
Edwards
&
Horn
1982
8­
Horn
&
Neighbors
1984
9­
Horn
et
al.
1985
10­
Barton
1982
11­
Horn
et
al.
1982
12­
Horn
et
al.
1986
13­
Fris
&
Horn
1993
14­
Miller
&
Marshall
1987
15­
Fitch
&
Lavenberg
1975
16­
Burge
&
Schultz
1973
17­
Sturm
&
Horn
1998
18­
Montgomery
1977
(
red
and
green
macroalgae,
such
as
Rhodoglossum
,
Porphyra
,
Microcaldia
,

Gigartina
,
and
Ulva
)
1,5,6,10,12,15,16,21,24,25,28
(
macroalgae,
such
as
Rhodoglossum
,

Phyllospadix,
Porphyra
,
Iridaea
,
Microcaldia
,

Macrocystis
Gigartina
,
Spongomorpha
and
Ulva
)
3,4,5,6,7,8,9,12,13,14,16,20,21,22,24,27,28
preference
for
annual
red
and
green
over
perennial
red
8,11,20,21
Nucleotide
diversity
(#)

Piscivorous
birds,
such
as
great
egrets
and
red­
breasted
merganser2
rockweed
gunnels
have
also
been
found
in
stomachs16
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9
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12
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14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CH
CI
CJ
CK
CL
CM
CN
CO
CP
CQ
RECRUITMENT
INTO
FISHERY
Genetic
variance
Mean
number
of
%
Polymorphic
First
Recruitment
Full
Recruitment
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Total
length
(
age)
Total
length
(
age)
Sampling
gear
eing
structure
/
Meth
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
Mean
heterozygosity
GENETICS
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Stichaeidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
CR
CS
CT
CU
CV
CW
CX
CY
CZ
DA
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Jones
1981
MORTALITY
competitively
excludes
Aniplarchus
purpurescens
from
rocky
habitat1
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1107
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Stichaeidae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
DB
DC
DD
DE
DF
DG
Interactions
/
associations
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
Residential1
None3
No
No2
1(<
2.1
±
0.3
m2)
Yes3
home
to
specific
underrock
refuge2
NF
Residential2
None3
No
No1
Secondary
resident3
1­
Horn
&
Riegle
1981
1­
Helm
1990
1­
Ralston
&
Horn
1986
1­
Ralston
&
Horn
1986
1­
Marliave
&
DeMartini
1977
2­
Edwards
&
Cech
1990
2­
Martin
1995
2­
Helm
1990
2­
Helm
1990
3­
Martin
1993
3­
Cross
1981
3­
Barton
1973
3­
Tenera
Environmental
Services
2000
4­
Gibson
1982
5­
Norris
1963
6­
Riegle
1977
7­
Kelly
et
al.
1985
8­
Love
&
Moser
1983
Movement
BEHAVIOR
infested
by
parasites
from
the
families
Opecoelidae
and
Hemiuridae8
No
movement
on
large
scale;

small
scale
movements
from
undderrock
refuge
to
foraging
sites1
often
found
with
the
black
prickleback
and
rockweed
gunnel5
one
of
the
highest
survival
times
out
of
water
realtive
to
other
stichaeoids;

have
air­
breathing
capabilities1,3,4,6,7
infested
by
parasites
from
the
family
Opecoelidae8
low
increase
in
survival
time
out
of
water
as
body
size
increases,
relative
to
other
stichaeoids;
high
survival
out
of
water
when
<
10
cm
SL;
have
airbreathing
capabilities1,2,3,4
Page
1108
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Stichaeidae
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56
57
58
59
60
61
62
63
64
65
A
B
C
D
E
Page
1109
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Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
F
G
H
I
JKLMN
Page
1110
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1167
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2220e
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Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
O
P
Q
R
S
T
U
V
W
X
Y
Page
1111
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
Z
AA
AB
AC
AD
AE
Page
1112
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1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
AF
AG
AH
AI
AJ
AK
AL
AM
AN
Page
1113
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
Page
1114
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
BC
BD
BE
BF
BG
Page
1115
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
BH
BI
BJ
BK
BL
BM
BN
Page
1116
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
BO
BP
BQ
BR
BS
BT
BU
Page
1117
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
BV
BW
BX
BY
BZ
19­
Urquhart
1984
20­
Jones
1981
21­
Yoshiyama
et
al.
1986
22­
Grossman
1982
23­
Cross
1981
24­
Kelly
et
al.
1985
25­
Tenera
Environmental
Services
2000
26­
Chadwick
1976
27­
Love
1996
28­
Wourms
and
Evans
1974
29­
Schultz
and
DeLacy
1932
Page
1118
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
CA
CB
CC
CD
CE
CF
CG
19­
Sousa
1979
20­
Irelan
&
Horn
1991
21­
Horn
1989
22­
Helm
1990
23­
Setran
1990
24­
Jones
1981
25­
Cross
1981
26­
Hobson
1994
27­
Tenera
Environmental
Services
2000
28­
Love
1996
Page
1119
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
CH
CI
CJ
CK
CL
CM
CN
CO
CP
CQ
Page
1120
of
1167
5­
2220e
­
Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
CR
CS
CT
CU
CV
CW
CX
CY
CZ
DA
Page
1121
of
1167
5­
2220e
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Stichaeidae
55
56
57
58
59
60
61
62
63
64
65
DB
DC
DD
DE
DF
DG
Page
1122
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Stichaeidae
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5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
A
B
C
D
E
F
Total
Length
(
cm)

Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)
to
weight
(
g)
equation
Stromateidae1,2
Peprilus
simillmus
1,2
Pacific
butterfish
12­
182
m
NF
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Eschmeyer
and
Herald
1983
1­
Love
et
al
2000
1­
Love
et
al
2000
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
3­
Hart
1973
Pacific
pompano,
butterfish,

California
pompano,
California
butterfish,
pompano
1,2,3
TAXONOMY
RANGE
Queen
Charlotte
Sound,
British
Columbia
to
Bahia
Magdelena,
southern
Baja
California
and
Gulf
of
California1
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1123
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Stromateidae
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67
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
G
H
I
J
K
L
M
N
O
P
Q
Max.
reported
Geographic
area
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
Female
Both
Male
Female
Both
Male
Female
NF
128
cm
TL
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Love
et
al
2000
Linf
in
cm
(
SE)
AGE
and
GROWTH
k
(
SE)
to
(
SE)

Page
1124
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
R
S
T
U
V
W
X
Y
Z
r2
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Longevity
(
yrs.)

NF
NF
NF
NF
NF
NF
NF
Growth
Parameters
Page
1125
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Stromateidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AA
AB
AC
AD
AE
AF
AG
AH
AI
Age
composition
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
Secondary
Characteristics
Male
Female
NF
oviparous1,2
No
14
cm
14
cm
1­
Matarese
et
al
1980
2­
Love
1996
Total
lengt
1st
maturity
REPRODUCTION
Breeding
Behavior
Sexual
dimorphism
Page
1126
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Stromateidae
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5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AJ
AK
AL
AM
AN
AO
AP
AQ
AR
AS
Fertilized
eggs
present
Male
Female
Ageing
structure
/
Method
Validation
Verification
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
NG
NG
NG
NG
NG
southern
California5
California1,3
Pacific
coast2,4
1­
Matarese
et
al
1980
2­
Love
1996
3­
Fitch
and
Lavenberg
1971
4­
Fitch
and
Lavenberg
1975
5­
Goldberg
1980
h
(
age)
at
Fertilization
and
Spawning
Period
(^^^

50%
maturity
Page
1127
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Stromateidae
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Mean
Fecundity
t
Spawning
females
present
Both
reproductive
states
present
eggs/
g
Number
of
eggs
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
(
weight
range)
(
at
cm,
grams,
or
age)

Multiple
spawner1
NF
NF
1­
Goldberg
1980
^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
Fecundity
Page
1128
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5
67
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BF
BG
BH
BI
BJ
BK
Size
specific
fecundity
eq
Size
at
settlement
Size
(
length
or
weight
range)
Egg
duration
Larval
duration
&
location
Annual
cohort
success
(
diameter
in
mm)

NF
NF
NF
NF
NF
NF
DISPERSAL
&
RECRUITMENT
Page
1129
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Stromateidae
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8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BL
BM
BN
BO
BP
BQ
BR
Eggs
Habitat
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
pelagic1,2,3,4
NF
1,31.8­
20
mm
SL
most
found
nearshore1,3
NF
NF
NF
21.8­
28
mm
SL
nearshore
and
coastal
waters2
1­
Matarese
et
al
1980
1­
Matarese
et
al
1980
1­
Love
1996
2­
Moser
1996a
2­
Moser
1996a
2­
D'Vincent
et
al
1980
3­
Fitch
and
Lavenberg
1971
3­
D'Vincent
et
al
1980
3­
Moser
and
Pommeranz
1999
4­
Fitch
and
Lavenberg
1975
4­
Horn
1970
5­
Horn
1970
Food
habits
Larvae
nearshore
and
several
miles
offshore,
but
generally
in
large
protected
areas4
generally
spawned
a
few
miles
offshore
in
surface
waters5
occurs
between
the
surface
and
48
m4
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1130
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BS
BT
BU
BV
BW
BX
BY
Predators
Fish
Size
Habitat
Foraging
guild
Foraging
method
When
/
Frequenc
Predators
NF
327.9
mm
to
maturity
associated
with
soft
bottom
habitat1
NF
NF
NF
NF
3­
D'Vincent
et
al
1980
1­
D'Vincent
et
al
1980
Juveniles
Food
habits
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
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1131
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2220e
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Stromateidae
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67
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
BZ
CA
CB
CC
CD
Habitat
Foraging
guild
Foraging
method
When
/
Frequency
Predators
over
soft
bottom
on
continental
shelf
1,3
carnivore3
NF
NF
inhabits
shallow
water
nearshore2
(
small
crustaceans)
1,2
Caspian
terns
and
Fosters'
terns3
1­
Moser
1996a
1­
Fitch
and
Lavenberg
1971
1­
Love
1996
2­
Fitch
and
Lavenberg
1971
2­
Fitch
and
Lavenberg
1975
2­
Fitch
and
Lavenberg
1973
3­
D'Vincent
et
al
1980
3­
Horn
1970
3­
Harvey
1982
Adults
California
halibut,
barracuda,
kelp
bass
and
other
large
predators2
Food
habits
kelp
bass,
barracuda,
California
halibut,

California
sea
lions,
Dall's
porpoise,
common
murres,
tufted
puffins,
least
terns,
rhinoceros
auklets
and
Brandt's
cormorants1
Page
1132
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5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CE
CF
CG
CH
CI
CJ
CK
CL
Genetic
variance
Haplotype
diversity
Mean
number
of
%
Polymorphic
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)
loci
Expected
(
SE)
Observed
(
SE)
Approach
NF
NF
NF
NF
NF
NF
NF
NF
GENETICS
Nucleotide
diversity
(#)
Mean
heterozygosity
Page
1133
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Stromateidae
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9
10
11
12
13
14
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CM
CN
CO
CP
CQ
CR
CS
CT
RECRUITMENT
INTO
FISHERY
First
Recruitment
Full
Recruitment
Effective
population
size
Fishery
/
Total
Length
(
age)
Fishery
/
Total
Length
(
age)
Sampling
gear
eing
structure
/
Meth
Validation
Verification
Natural
(
M)

NF
NF
NF
NF
NF
NF
NF
NF
Page
1134
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1167
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2220e
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Stromateidae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
CU
CV
CW
CX
CY
CZ
DA
Mortality
Estimates
Rate
of
Exploitation
Method
of
Age
range
used
Rate
of
Exploitation
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)
ratio
(
F/
Z)
Competition
NF
NF
NF
NF
NF
NF
NF
MORTALITY
Page
1135
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1167
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2220e
­
Stromateidae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
DB
DC
DD
DE
DF
DG
DH
Interactions
/
associations
with
other
organisms
Tagging
studies
Resident
or
Transient
Homing
ability
(
Home
range)
Schooling
Territoriality
temporary
associations
with
jellyfish
(
Pelagia
)
1
NF
NF
NF
Yes1,2
1­
Horn
1970
1­
Love
1996
2­
Fitch
and
Lavenberg
1971
BEHAVIOR
Movement
Page
1136
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2220e
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Stromateidae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
DI
DJ
DK
DL
DM
DN
DO
Page
1137
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1167
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2220e
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Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
A
B
C
D
E
F
Page
1138
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1167
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2220e
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Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
G
H
I
J
K
L
M
N
O
P
Q
Page
1139
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
R
S
T
U
V
W
X
Y
Z
Page
1140
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1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
AA
AB
AC
AD
AE
AF
AG
AH
AI
Page
1141
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
AJ
AK
AL
AM
AN
AO
AP
AQ
AR
AS
Page
1142
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
BD
BE
Page
1143
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
BF
BG
BH
BI
BJ
BK
Page
1144
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
BL
BM
BN
BO
BP
BQ
BR
Page
1145
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
BS
BT
BU
BV
BW
BX
BY
Page
1146
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
BZ
CA
CB
CC
CD
Page
1147
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
CE
CF
CG
CH
CI
CJ
CK
CL
Page
1148
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
CM
CN
CO
CP
CQ
CR
CS
CT
Page
1149
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
CU
CV
CW
CX
CY
CZ
DA
Page
1150
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
DB
DC
DD
DE
DF
DG
DH
Page
1151
of
1167
5­
2220e
­
Stromateidae
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
DI
DJ
DK
DL
DM
DN
DO
Page
1152
of
1167
5­
2220e
­
Stromateidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
A
B
C
D
E
Family
name
Scientific
name
(
Synonym)
Common
name(
s)
Geographic
range
Observed
depth
range
(
m)

Syngnathidae
1,2
Syngnathus
leptorynchus
1,2
bay
pipefish
1
Prince
William
Sound,
Alaska
to
Bahia
Santa
Maria,
NF
southern
Baja
California
1
Syngnathidae
1,2
Syngnathus
californiensis
1,2
kelp
pipefish
Bodega
Bay,
northern
California
to
Bahia
Santa
Maria,
shallow
to
15
m
1,2
great
pipefish
1,2
southern
Baja
California;
northern
populations
from
Alaska
to
Monterey
Bay;
southern
populations
from
Morro
Bay
southward
1
Syngnathidae
1,2
Hippocampus
ingens
1,2
Pacific
seahorse
Pt.
Conception,
southern
California
to
Peru
and
usually
offshore;
10
m
or
caballito
del
mar
1,2
including
Islas
Galapagos;
old
(
doubtful)
record
from
deeper
1,2
San
Francisco
Bay
1
1­
Robins
et
al
1991
1­
Robins
et
al
1991
1­
Love
1996
1­
Love
et
al
2000
1­
Eschmeyer
and
Herald
1983
2­
Eschmeyer
1998
2­
Eschmeyer
1998
2­
Gates
and
Frey
1974
2­
Love
et
al
2000
TAXONOMY
RANGE
Page
1153
of
1167
5­
2220e
­
Syngnathidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
F
G
H
I
JKL
Total
Length
(
cm)
Max.
reported
Geographic
area
to
weight
(
g)
equation
Length
conversion
equations
length
(
cm)
(
length
measurment)
Male
Female
Both
1
Both:
W=
0.000001445SL2.65
NF
1
38.5
cm
TL
NF
NF
NF
NF
2
Both:
log
W=
0.000199526TL3.12
3Both:
W=
0.00000000164L3.95516
(
L=
SL
(
mm))
NF
1
19
cm
TL
NF
NF
NF
NF
NF
NF
1
24.7
cm
TL
NF
NF
NF
NF
1­
Mahan
1985
1­
Love
et
al
2000
2­
Bayer
1980
3­
Quast
1968b
Linf
in
cm
(
SE)

Page
1154
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1167
5­
2220e
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Syngnathidae
1234
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
M
N
O
P
Q
R
S
T
U
V
W
r2
Male
Female
Both
Male
Female
Both
Male
Female
Both
Ageing
structure
/
Method
Validation
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
k
(
SE)
to
(
SE)
Growth
Parameters
AGE
and
GROWTH
Page
1155
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1167
5­
2220e
­
Syngnathidae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
X
Y
Z
AA
AB
AC
AD
AE
AF
Age
composition
Verification
Other
refs.
Longevity
(
yrs.)
information
Sexuality
Mating
system
Mode
of
reproduction
Parental
Care
Male
:
Female
SL
ratio
NF
1
1
18­
24
months
NF
NF
NF
oviparous
1
NF
NF
2
1
year
NF
NF
NF
NF
NF
NF
oviparous
1
yes,
female
lays
NF
eggs
in
male's
pouch
1
NF
NF
NF
NF
NF
NF
oviparous
1
NF
NF
1­
Fritzsche
1980
1­
Mahan
1985
1­
Moser
1996a
1­
Feder
et
al
1974
2­
Fritzsche
1980
Sexual
Breeding
Behavior
Page
1156
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1167
5­
2220e
­
Syngnathidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
AG
AH
AI
AJ
AK
AL
AM
AN
Secondary
Characteristics
Male
Female
Male
Female
Ageing
structure
/
Method
Validation
Verification
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
dimorphism
Total
length
(
age)
at
REPRODUCTION
1st
maturity
50%
maturity
Page
1157
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1167
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2220e
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Syngnathidae
1234
5
67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
AZ
BA
BB
BC
Fertilized
eggs
present
Both
Parturition
Other
refs.
Geographic
Spawning
Area
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
Spawning
Frequency
NF
Yaquina
Bay,
Oregon
3
NF
California
1
NF
southern
California
2
NF
NF
NF
NF
1­
Moyle
1976
2­
Feder
et
al
1974
3­
Bayer
1980
Fertilization
and
Spawning
Period
(^^^^^=
peak
fertilization;*****=
peak
spawn;
#####
=
doubtful
spawning
records
Page
1158
of
1167
5­
2220e
­
Syngnathidae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
BD
BE
BF
BG
BH
Mean
Fecundity
Size
specific
eggs/
g
Number
of
eggs
fecundity
eq
(
weight
range)
(
at
cm,
grams,
or
age)
(
length
or
weight
range)
Egg
duration
Larval
duration
NF
NF
NF
2­
3
week
incubation
1
NF
NF
NF
NF
28­
10
days
NF
NF
NF
NF
NF
NF
1­
Moyle
1976
2­
Fitch
and
Lavenberg
1975
Fecundity
DISPERSA
Page
1159
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5­
2220e
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Syngnathidae
1234
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67
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9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
BI
BJ
BK
BL
BM
BN
BO
Eggs
Size
at
settlement
Size
&
location
Annual
cohort
success
(
diameter
in
mm)
Habitat
Predators
Fish
Size
Habitat
NF
NF
1
1.3­
1.5
mm
eggs
brooded
within
specialized
NF
NF
NF
structure
on
adult1
NF
NF
1
1.1­
1.3
mm
eggs
brooded
within
specialized
NF
NF
NF
structure
on
adult
male1,2
NF
NF
1
<
4.0
mm
eggs
brooded
within
specialized
NF
NF
NF
structure
on
adult
1
1­
Ahlstrom
et
al
1984
1­
Ahlstrom
et
al
1984
2­
Fitch
and
Lavenberg
1975
3­
Limbaugh
1955
eggs
are
laid
by
female
in
pouch
of
male,
eggs
hatch
and
are
retained
inside
pouch
until
release
as
minature
adults3
AL
&
RECRUITMENT
Page
1160
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2220e
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Syngnathidae
1234
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67
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
BP
BQ
BR
BS
BT
BU
BV
Foraging
guild
Foraging
method
When
/
Frequency
Predators
Fish
Size
Habitat
Foraging
guild
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
amoung
seaweed
and
kelp
beds1NF
NF
NF
NF
NF
1
approx.
40
mm
NF
NF
1­
Ahlstrom
et
al
1984
1­
Feder
et
al
1974
2­
Limbaugh
1955
Food
habits
HABITAT
ASSOCIATION
&
TROPHIC
INTERACTIONS
Life
Stage
Larvae
Juveniles
Page
1161
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33
34
35
BW
BX
BY
BZ
CA
Foraging
method
When
/
Frequency
Predators
Habitat
Foraging
guild
NF
NF
common
in
eelgrass
of
bays
and
carnivore
sloughs
1,2
(
small
crustaceans)
1
NF
NF
kelp
beds
1
carnivore
(
mysids
and
small
amphipods)
2,5
(
mysids,
amphipods,
and
other
shrimp)
3
microcarnivore4
kelp
column
and
canopy5
NF
NF
NF
usually
offshore
1
carnivore
found
in
bays
attached
to
eelgrass4
1­
Love
and
Ebeling
1978
1­
Eschmeyer
and
Herald
1983
1­
Moyle
1976
2­
Moyle
1976
2­
Feder
et
al
1974
3­
Feder
et
al
1974
3­
Quast
1968d
4­
Jones
et
al
1988
4­
Quast
1968b
5­
Quast
1968b
5­
Limbaugh
1955
6­
Limbaugh
1955
(
tiny
planktonic
crustaceans;
such
as
mysids,

small
shrimp,
and
amphipods)
4
Food
habits
Fo
found
in
sand­
bottom
kelp
holdfast,

midkelp,
and
rocky­
bottom
kelp­
bed
habitat6
blue
rockfish,
olive
rockfish,

and
kelp
bass1
blue
rockfish,
olive
rockfish,

and
kelp
bass1
amoung
seaweeds,
in
canopy
and
mid­
kelp
habitat;
also
in
short
coralline
algae,
Torrey's
surgrass,
and
eelgrass
beds,
sometimes
drifting
in
kelp
many
miles
from
shore3
Adu
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31
32
33
34
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CB
CC
CD
CE
CF
CG
CH
Genetic
Haplotype
diversity
Mean
number
of
Foraging
method
When
/
Frequency
Predators
Substocks
(
Geographic
Area)
alleles
per
locus
(
SE)

suck
up
prey
with
tubular
mouth
1
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
sucks
food
into
mouth2
NF
yellowfin
tuna
1
NF
NF
NF
tunas
and
gulls2
1­
Moyle
1976
1­
Fritzsche
1980
2­
Fitch
and
Lavenberg
1975
2­
Fitch
and
Lavenberg
1975
ood
habits
ults
GEN
Nucleotide
diversity
(#)

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32
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CI
CJ
CK
CL
CM
CN
CO
CP
RECRUITMENT
INTO
FISHERY
variance
%
Polymorphic
First
Recruitment
Full
Recruitment
loci
Expected
(
SE)
Observed
(
SE)
Approach
Effective
population
size
Fishery
/
Total
Length
(
age)
Fishery
/
Total
Length
(
age)
Sampling
gear
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
ETICS
Mean
heterozygosity
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27
28
29
30
31
32
33
34
35
CQ
CR
CS
CT
CU
CV
CW
CX
CY
Mortality
Estimates
Rate
of
Ex
Method
of
Age
range
used
Rate
of
eing
structure
/
Meth
Validation
Verification
Natural
(
M)
Fishing
(
F)
Total
(
Z)
mortality
estimation
to
calculate
mortality
exploitation
(
u)

NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
MORTALITY
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CZ
DA
DB
DC
DD
DE
xploitation
Exploitation
Interactions
/
associations
ratio
(
F/
Z)
Competition
with
other
organisms
Tagging
studies
Resident
or
Transient
ng
ability
(
Home
NF
NF
early
juvenile
survivorship
is
limited
by
presence
oNF
in
summer
pipefish
may
emigrate
NF
adult
conspecifics
and
predators
1
from
the
lower
estuary
to
the
upper
parasites:
7
families,
12
spp.
(
see
reference)
2
estuary
(
Yaquina
Bay)
1
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
NF
1­
Behrents
1987
1­
Bayer
1980
2­
Love
and
Moser
1983
BEHAVIOR
Movement
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DF
DG
Schooling
Territoriality
NF
NF
NF
NF
NF
NF
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