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
MR.
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
FINAL
REPORT
31
AUGUST
2000
2
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.
General
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
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
FINAL
REPORT
31
AUGUST
2000
3
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
socioeconomic
variables
to
enable
CDFG
to
evaluate
the
trends
in
this
increasing
fishery.

WORK
ACCOMPLISHED
The
results
from
this
project
will
be
a
major
step
toward
characterizing
the
fish
assemblages
in
California's
nearshore
ocean
waters
and
providing
essential
information
for
measuring
the
impacts
of
human
activities,
measuring
economic
costs
and
benefits
to
the
state
from
commercial
and
recreational
fishing,
and
evaluating
alternative
management
strategies
for
marine
fisheries
through
the
FMP
process.
These
results
are
expected
to
enhance
the
quality,
availability,
and
consistency
of
life
history
data
used
in
CDFG
management
decisions,
and
to
make
these
data
available
to
the
public
and
other
government
or
private
academic
institutions.
This
research
will
complement
a
separate
state­
funded
nearshore
marine
Geographic
Information
System
(
GIS)
project
to
start
mapping
nearshore
sedimentary
and
rocky
habitats
in
0­
30
meters
off
California
and
classify
these
bathymetric
habitats
and
their
species
associations.

A.
ANALYSIS
AND
SUMMARY
OF
EXISTING
LIFE
HISTORY
PARAMETERS
The
most
recent
analyses
of
the
knowledge
of
the
life
histories
of
and
status
of
fisheries
for
species
taken
in
the
nearshore
fishery
was
accomplished
by
Leet
et
al.
(
1992)
for
California
and
by
Starr
et
al.
(
1998)
and
Lea
et
al.
(
1999)
for
central
California.
Much
information
about
life
histories
of
these
species
is
scattered
about
the
published
literature,
as
well
as
contained
in
state
and
federal
agency
reports
that
have
not
been
published
in
refereed
journals
or
made
publicly
available.
The
MLMA
has
specific
size
limits
for
a
select
group
of
fish
species
targeted
by
the
live
fish
fishery,
yet
an
examination
of
the
published
literature
on
these
species
does
not
provide
age
estimates
and
reproductive
maturity
information
that
supports
all
of
these
size
limits.
Therefore,
a
review
and
compilation
of
the
existing
information
is
imperative.
FINAL
REPORT
31
AUGUST
2000
4
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);
specieshabitat
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.
FINAL
REPORT
31
AUGUST
2000
5
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.

B.
IDENTIFICATION
OF
GAPS
IN
THE
EXISTING
KNOWLEDGE
OF
LIFE
HISTORY
PARAMETERS
FOR
NEARSHORE
FISH
SPECIES
The
database
serves
not
only
as
an
information
source,
but
also
serves
to
identify
where
information
is
lacking.
To
clarify
and
identify
true
gaps
in
the
literature,
three
abbreviations
were
used
where
information
was
not
provided:
1)
NA,
not
applicable;
2)
NG,
not
given;
or
3)
NF,
not
found.
Not
applicable
(
NA)
refers
to
a
life
history
parameter
that
is
not
valid
for
a
species
(
e.
g.
egg
stage
not
applicable
for
embiotocids).
Not
given
(
NG)
refers
to
a
study
where
similar
life
history
parameters
are
provided
within
a
section
of
the
survey,
but
not
for
the
specified
parameter
(
e.
g.
an
age
and
growth
study
where
von
Bertalanffy
growth
parameters
are
provided
for
males
and
females
but
not
for
the
sexes
combined;
or
a
maturity
study
with
age
at
first
maturity
but
not
age
at
50%
maturity).
Not
found
(
NF)
refers
to
a
parameter
where
no
information
was
found;
either
it
does
not
exist,
or
we
did
not
find
the
information.
Therefore
an
empty
space
or
cell
does
not
mean
a
lack
of
information,
rather
a
space
for
formatting.

There
are
many
gaps
in
the
existing
life
history
literature
of
California's
nearshore
fishes.
In
general,
the
following
life
history
information
is
lacking
for
most
species:
1)
age
validation
and
growth
studies,
2)
fecundity
estimates,
3)
measures
of
genetic
variance,
4)
size
and
age
at
settlement,
5)
mortality
estimates,
6)
recruitment
to
the
fishery,
and
7)
movement.

Large
gaps
in
the
existing
knowledge
of
life
history
parameters
occur
primarily
for
elasmobranchs,
and
for
the
families
Cottidae,
Hexagrammidae,
Muraenidae,
Anoplopomatidae,
Sphyraenidae,
Pleuronectidae,
Bothidae,
Polyprionidae,
Carangidae,
Anarhichadidae,
Scorpaenidae
(
several
species),
and
Embiotocidae
(
especially
age
validation).
Several
specific
gaps
are
identified
below
in
Part
C.

C.
ADDITIONAL
STUDIES
ON
THE
LIFE
HISTORY
OF
NEARSHORE
FISHES
Following
Goals
A
and
B,
we
are
pursuing
our
third
goal,
much
of
which
will
comprise
the
bulk
of
the
funding
we
have
requested
from
PSMFC
for
2000­
2001,
which
will
be
to
continue
some
of
the
research
that
we
initiated
in
the
first
year's
project
and
to
add
several
other
interesting
and
important
projects.
FINAL
REPORT
31
AUGUST
2000
6
1)
Age
and
Growth,
Longevity,
and
Age
at
First
Maturity
of
Selected
MLMA/
NFMA
Species
The
first
objective
will
be
to
initiate
age
and
growth,
reproduction,
and
age­
at­
maturity
studies
on
some
of
the
species
for
which
there
is
uncertainty
about
the
relationship
between
MLMA
size
limits
and
verified
or
validated
estimates
of
age.
Our
literature
review
this
year
indicated
that
some
of
the
size
limits
did
not
agree
well
with
existing
knowledge
of
the
age
at
first
maturity
for
some
species.
To
determine
how
vulnerable
some
of
the
nearshore
fish
stocks
are
to
exploitation,
an
idea
of
their
longevity,
based
on
these
proposed
validated
age
and
growth
studies
and
coupled
with
better
estimates
of
natural
and
fishing
mortality
and
fecundity
of
older/
larger
age
classes,
will
result
in
better
information
for
nearshore
fishery
management.

One
of
the
species
that
requires
the
most
urgent
attention
is
the
cabezon
(
Scorpaenichthys
marmoratus),
which
despite
being
quite
abundant
and
popular
in
historic
recreational
fisheries,
has
not
been
very
thoroughly
studied,
especially
in
California
(
O'Connell
1953;
Lauth
1987,
1988).
We
propose
to
concentrate
on
age
and
growth
studies
that
utilize
more
modern
techniques
that
involve
sectioning,
and
we
will
conduct
marginal
increment
analysis
to
validate
the
growth,
age
at
maturity,
and
longevity
of
this
important
nearshore
species.
To
determine
length
at
maturity,
the
size
and
degree
of
maturity
of
the
gonads
will
be
examined
throughout
the
year.
Cabezon
are
being
collected
from
northern
and
southern
California
to
determine
if
differences
exist
between
fish
from
these
two
geographic
areas.
These
data
will
provide
valuable
information
for
fishery
management.

Other
species
(
see
list
below
under
genetics
and
demography)
will
be
studied
in
next
year's
PSMFC
project,
and
will
concentrate
on
age,
growth,
reproduction
and
age­
at­
maturity
studies.
Inclusion
of
additional
species
in
this
list
will
depend
upon
the
samples
available
and
the
graduate
student's
interest
in
them
as
thesis
material.

Dr.
Ralph
Larson
(
SFSU)
and
his
graduate
students
analyzed
existing
and
newly
collected
samples
of
otoliths
and
morphometric
data
from
kelp
rockfish
(
Sebastes
atrovirens)
and
black
and
yellow
rockfish
(
S.
chrysomelas)
as
part
of
their
on­
going
studies
of
age
and
growth
of
these
two
species.

2)
Genetics
and
Demography
A
comparative
life
history
study
of
blue
rockfish
(
Sebastes
mystinus)
subpopulations
will
be
performed
at
five
sites
along
California:
1)
Santa
Barbara
mainland,
2)
San
Miguel
Island,
3)
central
California
(
Big
Creek
or
Monterey
Bay),
4)
the
Farallon
Islands,
and
5)
northern
California
(
Mendocino
or
Bodega
Bay).
Genetic
analyses
of
tissue
samples,
using
the
control
region
(
D­
loop)
of
mitochondrial
DNA,
will
be
used
to
measure
genetic
variance
and
population
substructure
among
geographic
sites.
In
addition,
a
stage­
based
(
larvae,
pelagic
juvenile,
settled
juvenile,
and
adult)
stochastic
demographic
model
will
be
produced
using
age
and
growth
information,
adult
mortality
(
estimated
from
VBGF
parameters
and
gonadal
indices),
age
at
first
maturity,
and
longevity
estimates
from
each
geographic
area.
This
model
will
provide
vital
statistics
 
such
as
intrinsic
rates
of
increase
and
rebound
potentials
 
for
each
subpopulation
(
Caswell
1989,
Au
and
Smith
1997).
FINAL
REPORT
31
AUGUST
2000
7
The
combination
of
both
genetic
and
demographic
measures
of
each
subpopulation
will
allow
a
better
understanding
of
population
"
health,"
adaptability,
and
longevity
(
Lacy
1987,
Lande
1988,
Nunney
1992,
Soltis
et
al.
1999).
This
unique
combination
will
also
provide
insight
into
nearshore
rockfish
genetic
relatedness
in
the
California
Current
and
the
effectiveness
of
marine
reserves
to
protect
residential
nearshore
species
with
long
larval
durations.
This
approach
will
be
a
further
attempt
to
formulate
an
index,
which
may
be
comparable
across
taxa
and
populations.

The
blue
rockfish
was
chosen
as
the
model
species
to
represent
the
nearshore
rockfish
complex,
because
sufficient
samples
are
obtainable;
whereas,
local
population
depletions
and
the
solitary
nature
of
other
species
(
S.
atrovirens,
S.
auriculatus,
S.
carnatus,
S.
caurinus.
S.
chrysomelas,
and
S.
rastrelliger)
makes
obtaining
sufficient
numbers
of
samples
more
difficult.
Although
this
study
will
focus
on
California
blue
rockfish
populations,
its
results
and
the
general
approach
will
be
applicable
to
fishes
with
similar
life
histories.
The
proposed
index
will
also
provide
a
means
of
comparison
for
fisheries
managers
to
evaluate
sub­
stock
structure
and
interactions.

3)
Seasonal
Reproductive
Timing
of
Nearshore
Fish
Species
Simply
setting
size
limits
does
not
insure
that
reproductively
capable
and/
or
active
females
are
protected
sufficiently.
For
example,
there
is
some
disagreement
in
the
literature
on
the
size
and
age
at
maturity
of
the
leopard
shark,
Triakis
semifasciata
(
Smith
and
Abramson
1990,
Cailliet
1992,
Kusher
et
al.
1992,
Au
and
Smith
1997,
and
Smith
et
al.
1998).
Thus,
the
California
size
limit
may
not
protect
female
leopard
sharks
so
that
they
can
successfully
reproduce
before
entering
the
fishery.
Likewise,
without
seasonal
closure
for
some
species
of
nearshore
rockfishes
during
their
spawning
season
(
Echeverria
1987),
it
is
possible
that
fishing
pressure
could
deleteriously
affect
reproductively
capable
females,
unless
there
were
closed
seasons
during
that
period
of
time.

In
response
to
this
need,
we
will
study
the
reproductive
cycle
and
stage
of
maturity
of
the
leopard
shark
by
monitoring
steroid
concentrations
in
their
blood.
The
purpose
of
this
study
will
be
to
examine
the
serum
steroid
hormones
in
female
leopard
sharks.
Previous
studies
have
correlated
17ß­
estradiol,
progesterone,
testosterone,
and
dihydrotestosterone
with
reproductive
events
(
Rasmussen
and
Gruber
1993,
Rasmussen
and
Gruber
1990,
Heupel
et
al.
1999,
Koob
and
Callard
1999,
Rasmussen
et
al.
1999).
Blood
samples
will
be
taken
seasonally
in
several
estuaries
along
the
central
California
coastline
and
analyzed
using
radioimmunoassays.
Monitoring
of
reproductive
hormones
can
provide
information
regarding
reproductive
mode,
gestation,
frequency
of
pregnancy
and
location
of
breeding
areas.
These
details
are
critical
in
formulating
effective
management
of
these
fisheries
(
Pratt
and
Otake
1990).
Future
extension
of
this
kind
of
research
will
prove
useful
to
many
other
groups
of
nearshore
fishes,
especially
the
viviparous
rockfishes,
for
which
seasonal
closure
might
be
an
effective
management
tool.
FINAL
REPORT
31
AUGUST
2000
8
LITERATURE
CITED
Au,
D.
W.
and
S.
E.
Smith.
1997.
A
demographic
method
with
population
density
compensation
for
estimating
productivity
and
yield
per
recruit
of
the
leopard
shark
(
Triakis
semifasciata).
Canadian
Journal
of
Fisheries
and
Aquatic
Sciences
54:
415­
420.

Cailliet,
G.
M.
1992.
Demography
of
the
central
California
population
of
leopard
shark
(
Triakis
semifasciata).
Australian
Journal
of
Marine
and
Freshwater
Research
43(
1):
183­
193.

California
Department
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Fish
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Game
(
CDFG).
1998.
Live
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In:
Review
of
some
California
Fisheries
for
1997.
California
Cooperative
Oceanic
Fisheries
Investigations
Reports
39:
17­
18.

Caswell,
H.
1989.
Matrix
Population
Models:
Construction,
Analysis,
and
Interpretation.
Sinauer
Associates,
Inc.,
Sunderland,
Massachusetts.
328
p.

Echeverria,
T.
W.
1987.
Thirty­
four
species
of
California
rockfishes:
maturity
and
seasonality
of
reproduction.
Fishery
Bulletin
85(
2):
229­
250.

Heupel,
M.
R.,
J.
M.
Whittier
and
M.
B.
Bennet.
1999.
Plasma
steroid
hormone
profiles
and
reproductive
biology
of
the
epaulette
shark,
Hemiscyllium
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Journal
of
Experimental
Zoology
284(
5):
586­
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Karpov,
K.
A.,
D.
P.
Albin,
W.
H.
Van
Buskirk.
1995.
The
marine
recreational
fishery
in
northern
and
central
California:
A
historical
comparison
(
1958­
86),
status
of
stocks
(
1980­
86),
and
effects
of
changes
in
the
California
Current.
California
Department
of
Fish
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Game,
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Bulletin
176,
192
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Koob,
T.
J.
and
I.
P.
Callard.
1999.
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endocrinology
of
female
elasmobranchs:
lessons
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the
little
skate
(
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dogfish
(
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Journal
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284(
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574.

Kusher,
D.
I.,
S.
E.
Smith
and
G.
M.
Cailliet.
1992.
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age
and
growth
of
the
leopard
shark,
Triakis
semifasciata,
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Lacy,
R.
C.
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managed
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immigration,
selection,
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R.
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R.
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Lauth,
R.
R.
1988.
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spawning
cycle,
spawning
frequency,
and
batch
fecundity
of
the
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marmoratus,
in
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Sound,
Washington.
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Lea,
R.
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R.
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A.
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rockfishes
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the
genus
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central
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with
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Game,
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177,
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REPORT
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2000
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W.
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Dewees,
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of
California,
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the
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of
grass
rockfish,
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rastrelliger,
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brown
rockfish,
S.
auriculatus,
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McKee­
Lewis,
K.
K.
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C.
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H.
L.
and
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work
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to
increase
our
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relative
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fishery
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S.
H.
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T.
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in
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S.
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mortality
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yield,
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replenishment
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on
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in
San
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88:
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