QL
155
.
S63
Biological
Report
82(
1
l­
125)
TR
EL­
82­
4
December
1989
Species
Profiles:
Life
Histories
and
Environmental
Requirements
of
Coastal
Fishes
and
Invertebrates
(
Pacific
Southwest)

CRANGONID
SHRIMP
no.
82­
11.125
Fish
and
Wildlife
Service
U.
S.
Department
of
the
Interior
Coastal
Ecology
Group
Waterways
Experiment
Station
U.
S.
Army
Corps
of
Engineers
.
Biological
Report
82(
11.125)
TR
EL­
82­
4
December
1989
Species
Profiles:
Life
Histories
and
Environmental
Requirements
of
Coastal
Fishes
and
Invertebrates
(
Pacific
Southwest)

CRANGONID
SHRIMP
bY
Clifford
A
Siegfried
Biological
Survey
New
York
State
Museum
and
Science
Service
Albany,
NY
12230
Project
Officer
David
Moran
U.
S.
Fish
and
Wildlife
Service
National
Wetlands
Research
Center
1010
Gause
Boulevard
Slidell,
LA
70458
Performed
for
U.
S.
Army
Corps
of
Engineers
Coastal
Ecology
Group
Waterways
Experiment
Station
Vicksburg,
MS
39180
and
U.
S.
Department
of
the
Interior
Fish
and
Wildlife
Service
Research
and
Development
National
Wetlands
Research
Center
Washington,
DC
20240
DISCLAIMER
The
mention
of
trade
names
does
not
constitute
endorsement
or
recommendation
for
use
by
the
Federal
Government.

This
series
may
be
referenced
as
follows:

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

This
profile
may
be
cited
as
follows:

Siegfried,
CA
1989.
Species
profiles:
life
histories
and
environmental
requirements
of
coastal
fishes
and
invertebrates
(
Pacific
Southwest)­­
crangonid
shrimp.
U.
S.
Fish
Wildl.
Serv.
Biol.
Rep.
82(
11.125).
U.
S.
Army
Corps
of
Engineers,
TR
EL­
82­
4.
18
pp.
PREFACE
This
species
profile
is
one
of
a
series
on
coastal
aquatic
organisms,
principally
fish,
of
sport,
commercial,
or
ecological
importance.
The
profiles
are
designed
to
provide
coastal
managers,
engineers,
and
biologists
with
a
brief
comprehensive
sketch
of
the
biological
characteristics
and
environmental
requirements
of
the
species
and
to
describe
how
populations
of
the
species
may
be
expected
to
react
to
environmental
changes
caused
by
coastal
development.
Each
profile
has
sections
on
taxonomy,
life
history,
ecological
role,
environmental
requirements,
and
economic
importance,
if
applicable.
A
three­
ring
binder
is
used
for
this
series
so
that
new
profiles
can
be
added
as
they
are
prepared.
This
project
is
jointly
planned
and
Gnanccd
by
the
U.
S.
Army
Corps
of
Engineers
and
the
U.
S.
Fish
and
Wildlife
Service.

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

Information
Transfer
Specialist
U.
S.
Fish
and
Wildlife
Service
National
Wetlands
Research
Center
NASA­
Slide11
Computer
Complex
1010
Gause
Boulevard
Slidell,
LA
70458
or
U.
S.
Army
Engineer
Waterways
Experiment
Station
Attention:
WESER­
C
Post
Office
Box
63
1
Vicksburg,
MS
39180
.
.
.
111
CONVERSION
TABLE
Multiply
BY
To
Obtain
millimeters
(
mm)
centimeters
(
cm)
meters
(
m)
meters
kilometers
(
km)
kilometers
square
meters
(
m')
square
kilometers
(
km2)
hectares
(
ha)
0.03937
inches
0.3937
inches
3.281
feet
0.5468
fathoms
0.6214
statute
miles
0.5396
nautical
miles
10.76
square
feet
0.3861
square
miles
2.471
acres
liters
(
1)
0.2642
cubic
meters
(
m3)
gallons
35.31
cubic
feet
cubic
meters
0.0008110
acre­
feet
milligrams
(
mg)
0.00003527
ounces
grams
(
g)
0.03527
ounces
kilograms
(
kg)
2.205
pounds
metric
tons
(
t)
2205.0
pounds
metric
tons
1.102
short
tons
kilocalories
(
kcal)
Celsius
degrees
("
C)

inches
inches
feet
(
ft)
fathoms
statute
miles
(
mi)
nautical
miles
(
nmi)

square
feet
(
ft2)
square
miles
(
mi2)
acres
3.968
1.8("
C)+
32
U.
S.
Customary
to
Metric
25.40
2.54
0.3048
1.829
1.609
1.852
British
thermal
units
Fahrenheit
degrees
millimeters
centimeters
meters
meters
kilometers
kilometers
0.0929
square
meters
2.590
square
kilometers
0.4047
hectares
gallons
(
gal)
3.785
liters
cubic
feet
(
ft3)
0.0283
1
cubic
meters
acre­
feet
1233.0
cubic
meters
ounces
(
oz)
28350.0
milligrams
ounces
28.35
grams
pounds
(
lb)
0.4536
kilograms
pounds
0.00045
metric
tons
short
tons
(
ton)
0.9072
metric
tons
British
thermal
units
(
Btu)
0.2520
kilocalories
Fahrenheit
degrees
("
F)
0.5556
("
F
­
32)
Celsius
degrees
Metric
to
U.
S.
Customary
iv
CONTENTS
Page
PREFACE
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111
CONVERSION
TABLE
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iv
ACKNOWLEDGMENTS
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vi
NOMENCLATURE/
TAXONOMY/
RANGE
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1
MORPHOLOGY/
IDENTIFICATION
AIDS
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1
REASON
FOR
INCLUSION
IN
SERIES
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4
LIFE
HISTORY
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4
Spawning
and
Larvae
..............................................
4
Postlarvae
and
Juveniles
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5
Migrations
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6
Adults
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6
GROWTH
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7
Mortality
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7
Disease
and
Parasites
..............................................
8
THE
FISHERY
ECOLOGICAL
R&
b
.
:
:
:
:
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:
:
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:
...........
8
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9
ENVIRONMENTAL
REQUIREMENTS
.................................
10
Temperature
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10
Salinity
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11
Temperature­
Salinity
Interactions
......................................
11
Substrate
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11
Freshwater
Flow
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12
Other
Environmental
Requirements
....................................
13
LITERATURE
CITED
..............................................
15
V
ACKNOWLEDGMENTS
.
I
am
grateful
for
the
reviews
by
Richard
Wahle
of
the
University
of
Maine
and
Kathy
Hieb
of
the
California
Department
of
Fish
and
Game,
Stockton,
California,
and
helpful
editorial
comments
by
an
unidentified
reviewer,
which
greatly
improved
the
readability
of
this
work.

vi
Figure
1.
Representative
crangonid
shrimp
(
from
Smith
and
Carlton
1975).

CRANGONID
SHRIMP
NOMENCLATURE/
TAXONOMY/
RANGE
Three
species
of
crangonid
shrimp,
commonly
called
bay
shrimp
(
Figure
l),
are
important
to
epifaunal
decapod
shrimp
communities
of
the
Pacific
Southwest.

Scientific
name
.
.
.
.
.
.
.
Crangon
franciscotum
(
Stimpson)
Preferred
common
name
.
.
.
.
.
.
.
sand
shrimp
Other
common
names
.
.
.
grass
shrimp,
common
shrimp.
Scientific
name
.
.
.
.
Crangon
nigricauda
(
Stimpson)
Preferred
common
name
.
.
.
.
.
.
.
black
shrimp
Other
common
names
.
.
.
.
deep­
water
shrimp,
black­
tailed
shrimp
Scientific
name
.
.
.
.
.
Crangon
nigromaculata
(
Lockington)
Common
name
.
.
.
.
.
.
.
.
.
blue­
spotted
shrimp
Class
.
.
.
.
.
.
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.
,
.
.
.
Crustacea
Order
.
.
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.
.
.
.
.
.
Decapoda
Suborder
Natantia
Tribe
.
.
.
:
:
:
:
:
:
.
.
.
.
:
:
.
.
.*
.
.
.*
.*
.
.
.*
.
.
Caridea
Family
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Crangonidae
Geographic
range:
sand
shrimp
and
black
shrimp
are
widely
distributed
along
the
Pacific
coast
of
North
America­­
sand
shrimp
from
southeastern
Alaska
to
San
Diego,
California,
and
black
shrimp
from
Alaska
to
Baja
California
(
Figure
2)
(
Rathbun
1904)
and
possibly
as
far
west
as
Japan
(
Schmitt
1921).
Both
shrimp
are
abundant
in
bays
on
mud
and
sand
bottoms
and
offshore
in
deeper
waters
(
Carlton
and
Kuris
1975).
The
range
of
blue­
spotted
shrimp
is
more
restricted;
it
occurs
from
the
Gulf
of
Farallons
to
Baja
California
(
Rathbun
1904)
(
Figure
2).
It
is
rarely
abundant
in
embayments,
living
more
commonly
offshore
on
mud
and
sand
bottoms.

MORPHOLOGY/
IDENTIFICATION
AIDS
The
crangonid
shrimp
of
the
Pacific
Southwest
are
easily
distinguished
from
other
members
of
the
tribe
Caridea
by
four
features
(
Figure
3);
(
1)­
the
rostrum
is
very
short,
generally
not
extending
beyond
the
eyestalks,
(
2)­
the
body
is
dorsally
flattened,
(
3)­
the
chelipeds
are
not
strongly
developed,
i.
e.,
they
1
CAPE
MENDOCINO
I
/

NEVADA
CALIFORNIA
`/
:
.
.:
:
:
.
:
.
.:
.
:
.
:
:
:
.
.:
.
t.
b
:
:
0'
.
.
,*
:
.
.

~~
:
:
.
.
.
.
.

"
aQy::
I
"
q.
.
cj
:
:
.
a
.
..
.
:
:
R,
.*
:
.
:
:
:
:
.
:=
*
:
:
:

1
.
.
,
m
%
LOS
ANGELES
SAN
DIEGO
­
­
­
­
­
­
_
PACIFIC
OCEAN
Crangon
franciscorum
z::.*
and
C.
nigricauda
t.
.
.
.

C.
nigromaculata
ezd
Figure
2.
Distribution
of
crangonid
shrimp
in
the
Pacific
Southwest.

2
b
­
rostrum
Q
Figure
3.
Distinguishing
characteristics
of
crangonid
shrimp
(
from
Smith
and
Carlton
1975).

are
subchelate
in
form,
and
(
4)­
the
eyes
are
not
covered
by
the
carapace
(
Carlton
and
Kuris
1975).
Shrimp
of
the
genus
Crangon
are
further
distinguished
by
a
single
median
spine
in
the
gastric
region
of
the
carapace.

The
three
species
of
Crangon
are
easily
distinguished
by
the
structure
of
the
cheliped
or
the
presence
of
distinctive
markings.
The
"
hand"
of
the
cheliped
of
sand
shrimp
is
slender
and
elongate
(
Figure
3)
and
the
"
finger,"
when
closed,
turns
back
almost
longitudinally.
The
hand
of
the
cheliped
of
black
shrimp
is
more
robust:
the
closed
finger
of
the
hand
is
directed
almost
transversely.
The
hand
of
the
3
cheliped
of
blue­
spotted
shrimp
is
intermediate
in
form­­
not
as
elongate
as
that
of
sand
shrimp
and
not
as
robust
as
that
of
black
shrimp.
The
differences
in
cheliped
shape
are
not
always
distinctive
in
small
shrimp
but
sand
shrimp
have
a
pair
of
spines
on
the
fifth
abdominal
segment
that
can
be
used
to
separate
small
sand
shrimp
from
small
black
shrimp.
Blue­
spotted
shrimp
are
easily
recognized
(
in
life)
by
a
prominent
dark
circular
spot
with
a
blue
center
surrounded
by
a
black
and
then
a
yellow
ring
on
each
side
of
the
6th
abdominal
segment
(
Carlton
and
Kuris
1975).

The
sexes
of
mature
crangonid
shrimp
are
easily
distinguished.
Sand
shrimp
sexes
can
be
distinguished
at
about
26­
30
mm
while
bluespotted
and
black
shrimp
can
be
separated
at
22­
24
mm
(
Israel
1936;
Krygier
and
Horton
1975;
Siegfried
1980).
The
most
distinguishing
characteristic
separating
the
sexes
is
the
structure
of
the
endopodite
of
the
second
pleopod
(
Lloyd
and
Yonge
1947;
Meredith
1952).
Males
have
an
appendix
masculina
(
Figure
4)
on
the
endopodite
of
the
second
pleopod,
whereas
females
do
not,
i.
e.,
the
first,
second,
and
third
pleopods
look
alike.
The
structure
of
the
endopodite
of
the
first
pleopod
is
short
and
curved
inward
in
males
and
long
and
straight
in
females.
The
location
of
the
gonopore
is
still
another
distinguishing
characteristic,
but
it
is
often
difficult
to
recognize
in
preserved
shrimp
(
Siegfried
1980).
The
gonopore
is
at
the
base
of
the
fifth
pair
of
walking
legs
in
males
and
at
the
base
of
the
third
pair
of
walking
legs
in
females.

exopod
appendix
masculina
\

kl
lcm
endopod
cf
Figure
4.
Sexual
dimorphism
in
Crangon
ji­
anckorzun
(
from
Smith
and
Carlton
1975).
REASONS
FOR
INCLUSION
IN
SERIES
The
crangonid
shrimp
of
the
California
coast
have
been
fished
commercially
since
the
1800'
s.
This
commercial
fishery
was
centered
in
San
Francisco
Bay.
Before
the
1960'
s,
most
of
the
catch
was
dried
and
shipped
to
the
Orient,
but
part
of
it
went
to
the
fresh
fish
markets
(
Israel
1936).
After
the
1960'
s,
the
fishery
became
primarily
a
bait
fishery,
and
annual
harvests
were
less
than
200,000
pounds.

The
importance
of
crangonid
shrimp
in
the
food
web
of
coastal
estuaries
is
well
documented.
These
shrimp
are
often
the
predominant
food
of
the
principal
sport
and
commercial
fishes
of
the
Pacific
coast
estuaries
(
Johnson
and
Calhoun
1952;
Ganssle
1966;
Haertel
and
Osterberg
1966;
Boothe
1967).
Crangonids
are
opportunistic
predators
that
feed
on
the
most
abundant
small
epibenthic
and
benthic
fauna
and
thus
serve
as
an
important
step
in
the
transfer
of
energy
from
the
primary
consumers
and
detritivores
to
the
top
predators
of
estuarine
food
webs.

The
agitation
of
bottom
sediments
by
crangonids
in
search
of
food
and
protection
may
be
important
in
the
cycling
of
nutrients
in
coastal
systems
(
Lloyd
and
Yonge
1947).
Fecal
pellets
of
crangonids
are
important
substrates
for
bacterial
colonization
and
may
be
important
in
nutrient
regeneration
in
shallow
water
habitats
(
Knight
1978).
Nitrogen
excretion
by
large
populations
of
crangonids
in
shallow
water
may
have
important
influences
on
the
nitrogen
budget
of
coastal
systems
(
Nelson
et
al.
1979).
Crangonid
shrimp
have
recently
become
the
subject
of
research
on
aquaculture
potential
of
native
decapods
(
Knight
1978)
and
as
a
standard
bioassay
test
organism
for
west
coast
estuaries
(
Dr.
P.
Sheehan,
Aqua
Terra
Technical;
pers.
comm.).

LIFE
HISTORY
Spawning
and
Larvae
Crangonid
shrimp
carry
their
eggs
under
the
abdomen
attached
to
and
between
the
basal
joints
and
inner
rami
of
the
pleopods.
The
distribution
and
abundance
of
ovigerous
females
is
a
useful
index
of
reproductive
activity.
Several
investigators
have
reported
that
the
spawning
season
of
crangonid
shrimp
is
long.
Ovigerous
females
have
been
reported
to
occur
during
9
to
12
months
of
the
year
in
various
populations
(
Israel
1936;
Lloyd
and
Yonge
1947;
Meredith
1952;
Price
1962;
Krygier
and
Horton
1975).

Ovigerous
females
of
the
three
species
reviewed
in
this
report
can
be
found
year­
round
along
the
California
coast.
Ovigerous
sand
shrimp
are
usually
most
abundant
in
the
spring
and
summer
in
coastal
embayments
but
are
abundant
offshore
in
winter
(
California
Department
of
Fish
and
Game
1987).
The
abundance
of
ovigerous
black
shrimp
is
generally
bimodal,
peaking
in
winter­
spring
and
summer­
fall.
They
are
usually
found
in
embayments;
few
are
collected
in
nearshore
areas.
Ovigerous
blue­
spotted
shrimp
are
usually
most
abundant
in
nearshore
coastal
areas,
peaking
in
winter
(
California
Department
of
Fish
and
Game
1987).

Israel
(
1936)
reported
that
both
male
and
female
sand
shrimp
live
for
about
one
year.
More
recent
investigations
suggest
that
females
may
live
1.5
to
2.5
years
and
males
1.5
years
(
Hatfield
1985;
Kinnetic
Laboratories
1987).
Repeated
spawning
has
been
demonstrated
by
the
presence
of
females
bearing
ovarian
stages
5­
7,
early
in
the
spawning
season,
as
well
as
eggs
on
their
egg
pad
(
Israel
1936;
Krygier
and
Horton
1975).

Ovigerous
black
shrimp
females
usually
remain
partly
buried
in
the
sediment
during
the
day.
The
eggs
usually
hatch
at
night.
Females
emerge
from
the
sediment
and
beat
their
pleopods,
generating
currents
that
release
the
newly
hatched
larvae.
The
females
are
usually
free
of
eggs
by
dawn
(
Villamar
and
Brusca
1988).

The
brood
sizes
of
crangonid
shrimp
are
related
to
shrimp
size
and
species.
Siegfried
(
1980)
determined
the
best
fit
relationship
between
brood
size
of
sand
shrimp
from
San
Francisco
Bay
and
total
length
to
be:

Log
N
=
­
3.66
+
4.09
Log
TL,
d
4
Juvenile
(
and
adult)
black
shrimp
are
rare
in
waters
of
<
lo
ppt
salinity
(
California
Department
of
Fish
and
Game
1987).

Mgmtions
Both
sand
shrimp
and
black
shrimp
migrate
to
deeper,
more
saline
water
as
they
mature
(
Krygier
and
Horton
1975;
Siegfried
1980,
Hatfield
1985).
This
out­
migration
from
lowsalinity
water
appears
to
be
related
to
reproduction,
as
it
coincides
with
the
development
of
sexual
characteristics.
The
migration
is
particularly
pronounced
in
sand
shrimp.
Juveniles
are
often
found
in
the
upper
reaches
of
estuaries,
in
nearly
fresh
water.
As
the
shrimp
mature,
they
move
to
water
of
higher
salinity,
which
may
result
in
size
gradients
in
sand
shrimp
populations.
The
mean
length
of
sand
shrimp
collected
in
midsummer
in
the
San
Francisco
Bay
Estuary
ranged
from
31
mm
near
the
upstream
limit
of
their
distribution
(
1
ppt)
to
>
50
mm
in
the
central
bay
(
Siegfried
et
al.
1978;
Siegfried
1980).

Further
evidence
for
an
outward
migration
related
to
reproductive
state
is
provided
by
information
on
mean
salinity
of
occurrence
of
females
bearing
eggs
of
various
stages
(
California
Department
of
Fish
and
Game
1987).
The
range
of
salinity
for
the
three
crangonid
species
is
more
restricted
for
ovigerous
females
than
for
other
life
stages.
Females
bearing
stage­
l
eggs
are
found
at
salinities
of
0.1
to
33.8
ppt
(
mean
20
ppt).
Gvigerous
sand
shrimp
are
generally
found
only
at
salinities
greater
than
14.6
ppt
(
Hatfield
1985;
Krygier
and
Horton
1975).
The
average
salinity
appears
to
increase
with
egg
stage
(
up
to
24.6
ppt
for
those
with
stage­
4
eggs;
California
Department
of
Fish
and
Game
1987).
Females
bearing
stage­
4
eggs
were
not
collected
from
waters
of
salinity
less
than
3.7
ppt.

Ovigerous
black
shrimp
apparently
prefer
somewhat
higher
salinities
of
about
25
ppt
(
California
Department
of
Fish
and
Game
1987);
their
outward
migration
is
less
extensive
because
they
do
not
penetrate
as
Ear
inland
as
sand
shrimp.
Blue
spotted
shrimp
prefer
even
higher
salinities;
ovigerous
females
are
scarce
in
embayments
but
abundant
offshore
(>
30
ppt;
California
Department
of
Fish
and
Game
1987).

The
outward
migration
of
crangonid
shrimp
is
believed
to
be
related
to
temperature­
salinity
interactions.
Ovigerous
females
are
found
in
coastal
embayments
in
summer
but
are
uncommon
in
them
in
winter;
they
seemingly
migrate
offshore
in
winter,
possibly
in
response
to
water
temperature
fluctuation
(
Hatfield
1985).
This
offshore
population
then
contributes
larvae
and
postlarvae
for
the
spring
abundance
peaks.

Sand
shrimp
also
undergo
die1
vertical
migrations.
Siegfried
et
al.
(
1978)
first
reported
a
die1
pattern
in
which
the
shrimp
enters
the
water
column
and
disperses
through
the
water
column
at
night
but
remains
on
or
near
the
bottom
during
daylight.
The
ecological
significance
of
this
behavior
remains
unknown,
but
the
habit
may
serve
to
allow
feeding
near
the
surface
while
protected
by
darkness
from
visual
feeding
predators
(
fish).
Migration
into
the
water
column
may
also
be
a
response
to
the
movement
of
their
primary
food,
Neomysis
mercedis,
into
the
water
column
(
Welch
1970;
Siegfried
et
al.
1979).
Additional
studies
substantiated
the
die1
activity
patterns
of
sand
shrimp
and
N.
mercedis
(
Sitts
1978).
There
is
no
comparable
information
on
the
die1
activity
patterns
of
the
other
crangonids
of
the
California
coast.

A&
h
Male
black
shrimp
in
Yaquina
Bay,
Oregon,
have
been
reported
to
be
mature­­
i.
e.,
to
contain
ripe
sperm­­
at
lengths
of
26­
28
mm;
male
sand
shrimp
matured
at
34
mm
(
Krygier
and
Horton
1975).
Gvigerous
females
as
short
as
36.2
mm
for
black
shrimp
and
43.6
mm
for
sand
shrimp
were
reported
by
Krygier
and
Horton
(
1975)
in
Oregon
waters.
These
lengths
at
maturity
agree
well
with
findings
in
California
(
Israel
1936;
Siegfried
1980).

Israel
(
1936)
reported
a
seasonal
variation
in
the
sex
ratio
of
sand
shrimp
from
San
Francisco
Bay:
males
predominated
before
the
breeding
season
and
females
predominated
during
the
peak
of
the
breeding
season.
This
variation
can
Y
6
be
attributed
to
the
short
life
span
of
males,
which
are
believed
to
die
soon
after
copulation,
and
the
longer
life
span
of
at
least
some
of
the
breeding
females.
In
general
the
sex
ratios
of
crangonid
populations
of
the
Pacific
Southwest
appear
to
be
about
1:
l.
This
ratio
is
expected
in
nonsynchronously
spawning
populations
in
which
a
portion
of
the
population
has
more
than
one
brood
(
Krygier
and
Horton
1975).

GROWTH
In
all
crangonids,
males
and
females
grow
at
different
rates
(
Lloyd
and
Yonge
1947;
Meredith
1952;
Allen
1960;
Price
1%
2;
Krygier
and
Horton
1975;
Siegfried
1980).
The
length
of
male
sand
shrimp
from
San
Francisco
Bay
rarely
exceeded
50
mm
although
some
individuals
as
long
as
71
mm
long
have
been
collected
(
California
Department
of
Fish
and
Game
1987).
Female
sand
shrimp
longer
than
70
mm
were
commonly
collected
(
Israel
1936;
Siegfried
1980).
In
the
somewhat
smaller
black
shrimp,
males
generally
reached
40
mm
and
females
60
mm
(
Israel
1936).
The
longest
black
shrimp
reported
from
San
Francisco
Bay
were
59
mm
(
male)
and
64
mm
(
female)
long.
The
maximum
size
of
blue­
spotted
shrimp
is
believed
to
be
similar
to
that
of
black
shrimp.
The
length
of
crangonids
in
San
Francisco
Bay
are
somewhat
greater
than
in
Oregon,
where
Krygier
and
Horton
(
1975)
reported
maximum
lengths
of
~
40
mm
for
male
and
~
55
mm
for
female
black
shrimp
and
50
mm
(
males)
and
~
62
mm
(
females)
for
sand
shrimp.
The
shrimp
may
grow
larger
in
San
Francisco
Bay
because
water
temperatures
are
higher
there
than
in
Oregon,
presumably
leading
to
faster
growth
or
longer
growing
seasons.

Offshore
populations
of
crangonids
may
reach
much
larger
lengths.
Collections
of
sand
shrimp
off
the
mouth
of
the
Columbia
River
indicate
a
population
with
a
mean
length
>
80
mm
and
maximum
lengths
of
110
mm
(
Durkin
and
Lipovsky
1977).
Estuarine
populations
of
the
European
species,
C.
crangon,
attained
smaller
body
sizes
than
nearby
populations
of
the
same
species
in
marine
habitats
(
Maucher
l%
l).
Remane
and
Schlieper
(
1971)
suggested
that
reduction
in
size
of
marine
animals,
although
generally
slight
in
higher
Crustacea
living
in
brackish
water,
is
comparable
to
Bergmann's
Law:
size
is
related
to
features
of
the
physical
environment.
The
reduction
may
be
attributable
to
the
physiological
effects
of
salinity,
reduced
food
availability,
or
a
combination
of
these
and
other
factors.
Studies
of
osmotic
regulation
indicated
that
smaller
sand
shrimp
are
capable
of
better
hyper­
regulation
but
larger
ones
are
capable
of
better
hypo­
regulation
(
Shaner
1978).
Thus,
the
migration
of
larger
shrimp
to
high
salinity
waters
would
be
energetically
advantageous
and
may
lead
to
faster
growth.

Growth
rates
are
extremely
difficult
to
estimate
from
size­
frequency
histograms
derived
from
field
collections
of
crangonid
shrimp.
Immigration,
emigration,
temperature
and
salinity
effects,
and
differential
mortality
combine
to
obscure
growth
patterns.
Krygier
and
Horton
(
1975)
estimated
that
the
growth
of
juveniles
ranged
from
0.76
to
1.37
mm
per
week
in
Oregon.
Growth
rates
of
crangonids
in
California
are
somewhat
higher.
Kinnetics
Laboratories
(
1984)
estimated
male
and
female
sand
shrimp
>
30
mm
long
to
grow
1.7
to
2.4
mm
per
month.

Length­
weight
relationships
for
juvenile,
male,
and
female
sand
shrimp
were
given
by
Siegfried
(
1980).
The
regression
equations
describing
these
relationships
follow:

juveniles:
Log
W
=
­
5.41
+
2.58
LogTL
males:
Log
W
=
­
6.12
+
3.27
LogTL
females:
Log
W
=
­
6.62
+
3.57
LogTL
where
W
=
dry
weight
in
grams
and
TL
=
length
in
mm.
Analysis
of
covariance
revealed
significant
differences
in
slopes
between
the
length­
weight
regressions
of
juvenile
and
mature
shrimp.
The
difference
is
at
least
partly
attributable
to
gonadal
development.

Mortality
Annual
abundance
of
crangonid
shrimp
varies
widely
(
Siegfried
1980;
California
Department
of
Fish
and
Game
1987).
Annual
abundance
indices
for
sand
shrimp
in
San
Francisco
Bay
were
several
orders
of
magnitude
higher
in
some
years
than
in
others
from
1980
to
1985,

c
7
and
that
of
blue­
spotted
and
black
shrimp
varied
by
more
than
tenfold
(
California
Department
of
Fish
and
Game
1987).
Annual
abundance
of
crangonid
shrimp
appears
to
be
determined
mostly
by
mortality
of
larvae
and
postlarvae.
Mortality
due
to
predation
is
undoubtedly
high
and
may
explain
geographic
patterns
of
abundance
within
embayments
(
Kinnetic
Laboratories
1983;
Kuipers
and
Dapper
1984).
Recruitment
to
bay
populations
in
any
one
year,
however,
appears
to
depend
on
environmental
conditions.

Recruitment
of
crangonid
shrimp
to
San
Francisco
Bay
is
independent
of
the
abundance
of
ovigerous
females,
i.
e.,
the
parent
stock.
Correlations
between
annual
abundance
of
crangonid
larvae
and
postlarvae
and
of
ovigerous
females
are
non­
significant
(
California
Department
of
Fish
and
Game
1987),
suggesting
that
environmental
conditions
play
a
major
role
in
determining
annual
abundance.
Thus,
management
to
maintain
crangonid
populations
should
be
aimed
at
maximizing
recruitment
(
Christmas
and
Etzold
1977).

Annual
abundance
of
crangonid
shrimp
has
been
linked
to
the
volume
of
freshwater
flow
to
San
Francisco
Bay
(
California
Department
of
Fish
and
Game
1987).
The
volume
of
freshwater
inflow
determines
the
magnitude
of
seaward
and
landward
currents,
the
salinity
regime,
temperature,
and
the
distribution
and
abundance
of
other
organisms
including
crangonid
predators
and
prey
(
Siegfried
et
al.
1979;
Armor
and
Herrgesell
1985).
All
of
these
factors
play
major
roles
in
determining
crangonid
recruitment
and
mortality.

&
ease
and
Parasites
Crustaceans
are
subject
to
infection
by
bacteria,
fungi,
protozoans,
platyhelminths,
and
nematodes
which
can
cause
disease
(
Green
1968;
Couch
1978;
Overstreet
1978).
Although
infestation
of
crangonids
by
these
groups
has
been
observed
(
C.
A.
Siegfried,
pers.
obs.),
there
is
little
information
on
the
incidence
of
infection
or
the
effects
on
crangonid
populations.
In
crangonids
of
San
Francisco
Bay,
the
incidence
of
infection
by
microsporidian
protozoans
is
often
high
(
C.
A.
Siegfried,
pets.
obs.).
The
bopyroidean
branchial
isopod,
Argeia
­
­
pugettensis,
an
ectoparasite
in
the
branchial
chamber,
often
infects
crangonids
in
San
&
W
Francisco
Bay
(
Nelson
and
Simmons
1978)
and
in
Yaquina
Bay,
Oregon
(
Krygier
and
Horton
1975).
It
attacks
shrimp
in
San
Francisco
Bay
only
in
higher­
salinity
waters.
Krygier
and
Horton
(
1975)
reported
only
female
parasitized
sand
shrimp
in
Yaquina
Bay,
Oregon;
however,
no
parasitized
ovigerous
females
were
found.
In
San
Francisco
Bay
almost
all
parasitized
shrimp
appeared
to
be
females
(
Nelson
and
Simmons
1978).
Since
it
is
unlikely
that
the
isopod
would
attack
only
females,
and
since
castration
by
parasites
is
reported
for
other
crustacean
species,
it
is
likely
that
the
attachment
of
A.
pugettensis
results
in
castration
in
sand
shrimp
(
Nelson
and
Simmons
1978).
Castration
would
inhibit
gonadogenesis
and
castrated
male
shrimp
would
take
on
feminizing
characteristics,
including
larger
size.
A
larger
host
would
presumably
make
more
energy
available
to
the
parasitic
isopod.
Since
host
and
parasite
weights
are
positively
correlated,
early
attachment
of
the
parasite
and
growth
with
the
host
is
indicated
(
Nelson
and
Simmons
1978;
Kinnetics
Laboratories
1987;
Jay
1989).
i
'

Whether
female
or
castrated
male,
parasitized
crangonid
shrimp
are
still
significantly
smaller
than
nonparasitized
shrimp,
as
shown
in
a
field
study
conducted
in
Humboldt
Bay.
The
study
suggests
that
there
are
slower
growth
rates
in
the
parasitized
shrimp
(
Jay
1980).
Preliminary
laboratory
investigation
reveals
that
parasitism
by
A.
pugettensis
depresses
metabolic
rates
(
oxygen
consumption)
in
sand
shrimp
but
does
not
affect
excretion
rates
(
Nelson
and
Simmons
1978).

THE
FISHERY
The
earliest
commercial
fishing
for
crangonid
shrimp
along
the
California
coast
is
believed
to
have
been
done
in
San
Francisco
Bay
in
the
late
1860'
s
by
Italian
fishermen
(
Scofield
1919;
Bonnot
1932;
Israel
1936).
The
shrimp
were
taken
in
bag
seines
18.3
m
long
by
2.4
m
deep.
The
catch
of
crangonid
shrimp
greatly
increased
when
Chinese
fishing
camps
appeared
in
1871.
The
Chinese
introduced
the
use
of
the
Chinese
8
Ii
A\

t
shrimp
net,
a
funnel­
shaped
net
12.1
m
long
with
a
mouth
9.1
m
wide.
These
nets
were
held
stationary
by
a
system
of
lines
and
anchors,
and
shrimp
were
captured
as
they
were
carried
into
the
net
by
the
tide.
Chinese
nets
were
set
during
a
single
ebb
or
flood
tide
and
then
lifted
just
before
the
tide
turned.
By
1897,
26
Chinese
fishing
camps
operated
20
to
50
nets
each
in
San
Francisco
Bay,
landing
400
to
8,000
pounds
of
shrimp
per
camp
per
day.
In
the
early
1890'
s,
crangonid
shrimp
were
also
caught
in
Tomales
Bay,
north
of
San
Francisco,
but
the
fishery
was
abandoned
by
about
1895
(
Bonnot
1932).

The
local
market
for
crangonid
shrimp
was
saturated
soon
after
the
Chinese
began
shrimp
fishing.
However,
a
profitable
export
trade
soon
developed,
based
on
the
shipment
of
dried
shrimp
to
the
Orient.
The
use
of
Chinese
shrimp
nets
was
investigated
by
the
California
Fish
and
Game
Commission
in
1897
and
again
in
1910,
largely
to
assess
the
loss
of
young
fish
(
particularly
striped
bass,
Morone
saxatiZk)
in
the
Chinese
nets.
In
1901
the
California
State
Legislature
established
a
closed
season
to
shrimp
fishing
from
May
to
August.
By
1911
the
Chinese
shrimp
nets
were
prohibited,
but
in
1915
a
law
was
passed
to
allow
limited
use
of
the
nets
in
parts
of
San
Francisco
Bay
(
Scofield
1919).

Trawl
shrimp
fishermen
began
operating
in
San
Francisco
Bay
in
about
1910.
Trawl
fishermen
used
trawls
with
beams
of
5.5
to
6.1
m
and
an
18.3­
m
funnel­
shaped
net.
The
trawl
was
dragged
over
the
bottom
in
the
direction
of
the
tide.
A
single
haul
lasted
from
40
min
to
2
h,
and
a
day's
work
consisted
of
making
2­
4
hauls
and
catching
a
total
of
100
to
1,000
pounds
of
shrimp
(
Fry
1933).

The
annual
shrimp
catch
in
San
Francisco
Bay
exceeded
720
t
for
much
of
the
1920'
s
and
1930'
s;
the
peak
catch
of
more
than
1,591
t
was
made
in
1935
(
California
Department
of
Fish
and
Game
1987).
The
annual
catch
did
not
exceed
455
t
during
the
1940'
s
and
1950'
s,
and
had
declined
to
less
than
45
t
by
the
late
1950'
s,
it
did
not
exceed
114
t
except
in
1978
when
216
t
were
landed
(
California
Department
of
Fish
and
Game
1987).
The
catch
of
crangonid
shrimp
continued
to
be
used
for
fresh
or
dried
food
until
the
1960'
s.
However,
the
demand
declined
steadily
after
the
1930'
s
and
the
fishery
became
a
bait
fishery,
supplying
sport
fishermen.
Crangonid
shrimp
are
too
small
to
shell
and
market
economically.
The
bait
fishery
relies
entirely
on
trawls
to
capture
shrimp.

The
sport
fishermen
of
the
region
will
probably
continue
to
support
a
bait
fishery
landing
68­
91
t
of
crangonid
shrimp
annually.
The
prospect
of
expansion
of
the
fishery
is
poor.

ECOLOGICAL
ROLE
Little
is
known
about
the
ecology
of
larval
and
postlarval
crangonids.
The
larvae
are
presumably
predators
on
small
zooplankters,
such
as
copepods.
Larvae
have
been
maintained
in
the
laboratory
on
a
diet
of
Arlemiu
nauplii
(
Shaner
1978).

Juvenile
and
adult
crangonids
are
predaceous,
their
dietary
differences
being
related
to
shrimp
size
and
prey
availability
(
Siegfried
1982;
Wahle
1985).
Seasonal
and
geographical
dietary
studies
have
indicated
that
crangonid
prey
in
the
diet
is
generally
proportional
to
their
occurrence
in
an
estuary
(
Siegfried
1982;
Wahle
1985).
Wahle
(
1985)
who
studied
the
feeding
ecology
of
sand
shrimp
and
black
shrimp
in
San
Francisco
Bay,
found
that
these
species
feed
on
a
similar
array
of
benthic
prey
made
up
of
crustaceans,
polychaetes,
mollusks,
foraminiferans,
and
plant
material.
Amphipods
were
the
most
frequently
ingested;
barnacle
exuvia,
fish
eggs,
bryozoans,
hydrozoans,
and
mites
were
occasionally
ingested.
Black
shrimp
ate
significantly
more
amphipods
than
did
sand
shrimp.
Larger
crangonids
ate
larger
prey.
Foraminiferans,
copepods,
and
ostracods
were
taken
by
small
shrimp,
while
shrimp,
polychaetes,
and
isopods
were
taken
by
large
shrimp.

In
the
less
saline
regions
of
the
San
Francisco
Bay
Estuary­­
the
delta
region­­
the
most
important
food
of
sand
shrimp
is
the
opossum
shrimp,
Neomy.
sh
mercedis,
which
occurred
in
9
62%&
l%
of
all
sand
shrimp
gastric
mills
containing
prey
(
Siegfried
1982).
Larger
crangonids
ate
larger
mysids.
Sitts
and
Knight
(
1979)
suggested
that
predation
by
sand
shrimp
affected
the
population
structure
and
abundance
of
mysids
in
the
delta.

The
distribution
of
N.
mercedis
does
affect
the
distribution
of
sand
shrimp
in
the
San
Francisco
Bay
Delta
(
Siegfried
1980).
Not
only
is
crangonid
density
much
greater
in
locations
where
mysids
are
abundant,
but
crangonids
in
areas
of
high
mysid
density
take
more
prey
than
those
in
areas
of
low
prey
density
(
Siegfried
1982).
The
delta
region
of
San
Francisco
Bay
has
impoverished
benthic
communities
(
Nichols
1979)
and
thus
the
region
has
few
potential
prey
organisms.
This
may
be
an
important
factor
linking
the
distributions
of
crangonids
and
mysids
in
the
delta
region
of
San
Francisco
Bay.

Crangonid
shrimp
are
important
food
for
many
estuarine
fish.
They
have
been
reported
to
be
important
in
the
diets
of
striped
bass
(
Johnson
and
Calhoun
1952;
Huebach
et
al.
1%
3;
Ganssle
1966;
Kinnetic
Laboratories
1983);
white
sturgeon,
Acipenser
trunsmonfunus
and
green
sturgeon,
A.
medirostris
(
Ganssle
1966;
McKechnie
and
Fenner
1971);
and
staghorn
sculpin,
Leptocottus
armatus,
(
Ganssle
1966;
Boothe
1967;
Kinnetic
Laboratories
1983).
They
are
also
an
important
food
of
American
shad,
Alosu
supidissimu;
brown
smoothhound,
Mustelus
henlei;
Pacific
tomcod,
Microgudus
proximus;
and
white
catfish,
Ictulurus
cutus
(
Ganssle
1966).

Crangonid
shrimp
recycle
nutrients
during
their
feeding
activities.
Agitation
of
bottom
sediments
by
crangonids
searching
for
food
and
shelter
has
been
suggested
as
an
important
mechanism
of
nutrient
recycling
in
estuaries
(
Lloyd
and
Yonge
1947).
Nitrogen
excretion
by
large
populations
of
crangonids
can
have
important
effects
on
the
nitrogen
budget
of
estuarine
systems
(
Nelson
et
al.
1979).

ENVIRONMENTAL
REQUIREMENTS
TempemW
Water
temperature
is
a
critical
factor
not
only
in
survival
but
in
the
regulation
of
most
life
functions
of
cold­
blooded
organisms
such
as
crangonid
shrimp.
Water
temperature
affects
+
.
metabolic,
growth,
and
feeding
rates,
osmoregulation,
movement,
habitat
selection,
and
survival
(
Prosser
1950).
The
discharge
of
heated
effluents
may
restrict
the
distribution
of
crangonids
or
other
cold­
blooded
organisms
in
estuarine
systems,
and
sudden
temperature
changes
may
be
lethal.

The
seasonal
migrations
of
crangonids
have
been
linked
to
changing
water
temperatures.
The
spring
onshore
migration
of
juveniles
may
be
a
migration
to
warmer
waters
and
the
fallwinter
offshore
movement
of
mature
shrimp
may
be
a
migration
to
cooler
waters
(
Israel
1936;
Ktygier
and
Horton
1975;
Siegfried
1980;
Hatfield
1985).
Decreasing
water
temperature
resulted
in
the
movement
of
C.
septzknspinosu
from
shallow
to
deeper
areas
of
Chesapeake
Bay
(
Haefner
1976).
Havinga
(
1930)
suggests
that
the
seasonal
migrations
of
C.
crungon
can
be
explained
as
a
search
for
the
warmest
water
mass.

Crangonids
of
the
Pacific
Southwest
have
been
collected
over
a
wide
range
of
temperatures;
sand
shrimp,
6.3
to
23.9
"
C,
black
shrimp,
6.7
to
22.1
"
C;
and
blue­
spotted
shrimp,
7.8
to
20.2
"
C
(
California
Department
of
Fish
and
Game
1987).
Sand
shrimp
are
abundant
at
>
15
"
C,
black
shrimp
at
<
18
"
C,
and
bluespotted
shrimp
at
14­
18
"
C.

Temperature
tolerance
in
adult
sand
shrimp
under
laboratory
conditions
was
reported
by
Khorram
and
Knight
(
1977a).
The
research
indicated
a
significant
interaction
between
temperature
and
salinity
on
survival:
water
temperature
affected
survival
at
different
salinities
and
salinity
affected
survival
at
different
temperatures.
In
general,
survival
of
adult
sand
shrimp
was
poor
at
water
temperatures
below
10
"
C
or
above
20
"
C,
and
decreased
with
decreasing
salinity.
The
optimum
ranges
of
temperature
and
salinity
for
adult
sand
shrimp,
as
determined
by
response
surface
analysis
(
a
statistical
technique
that
determines
optimum
response
to
more
than
one
variable),
was
14.5­
17.0
"
C
and
18­
20
ppt.
The
authors
concluded
that
temperature
was
slightly
more
important
than
salinity
in
determining
adult
survival
(
Khorram
and
Knight
1977a).

10
Salinity
Crangonids
are
euryhaline,
occurring
at
salinities
from
nearly
fresh
water
to
seawater
(
Siegfried
1980,
Hatfield
1985).
Sand
and
black
shrimp
have
been
collected
from
San
Francisco
Bay
at
salinities
of
O­
1­
34.3
ppt
and
blue­
spotted
shrimp
at
the
somewhat
narrower
range
of
4.5
34.3
ppt
(
California
Department
of
Fish
and
Game
1987).
Blue­
spotted
shrimp
are
generally
abundant
only
in
water
with
salinity
>
23
ppt.
Black
shrimp
are
generally
more
abundant
in
water
with
salinity
>
23
ppt.
Black
shrimp
are
generally
most
abundant
in
waters
of
salinity
>
lO
ppt
and
sand
shrimp
at
salinities
~
19
ppt
(
California
Department
of
Fish
and
Game
1987).
These
salinity
ranges
are
based
on
the
abundance
of
juvenile
and
adult
stages.
Adult
crangonids,
and
particularly
ovigerous
females,
prefer
the
higher
end
of
the
salinity
range.

The
seasonal
distribution
of
crangonids,
particularly
sand
shrimp,
along
the
California
coast
is
closely
related
to
salinity.
Although
the
sand
shrimp
inhabits
brackish
water
during
much
of
its
life
cycle,
it
requires
relatively
high
salinities
for
reproduction.
Ovigerous
females
are
rarely
collected
where
salinity
is
low.
Ovigerous
females
are
found
year­
round
in
San
Francisco
Bay,
but
almost
never
in
the
less
saline
portions
of
the
bay
(
Israel
1936;
Siegfried
1980;
Hatfield
1985).
Energetic
demands
of
osmoregulation
at
low
salinities
may
preclude
egg
development
and
thus
reproduction
in
low
salinity
waters.
Broekema
(
1941)
showed
that
low
salinities
retard
egg
development
in
crangonids.
Salinity
is
thus
important
in
larval
survival;
preliminary
investigations
suggested
that
survival
of
larval
sand
shrimp
declined
at
salinities
below
12
ppt
(
S.
W.
Shaner,
Univ.
Calif.,
Davis,
unpubl.)

Salinity
tolerance
was
significantly
affected
by
water
temperature.
At
5
ppt
salinity,
the
96­
h
survival
was
zero
at
5
"
C
and
25
"
C,
but
ranged
from
30%
to
42%
at
intermediate
temperatures;
in
waters
of
25
ppt
salinity,
survival
after
%
h
was
>
80%
at
10
"
C
and
15
"
C
but
60%
at
5
"
C
(
Khorram
and
Knight
1977a).
More
recent
salinity
tolerance
investigations
indicated
that
juvenile
sand
shrimp
were
more
tolerant
of
low
salinity;
survival
was
100%
in
water
of
2
ppt
salinity
(
S.
W.
Shaner,
unpubl.)
Sand
shrimp
from
high
salinity
waters
(
20­
30
ppt)
acclimated
to
low
salinity
(
2.2­
5.5
ppt)
in
the
laboratory
within
5­
6
weeks.
At
this
time,
they
physiologically
resembled
sand
shrimp
from
low
salinity
(
l­
7
ppt)
waters
(
Shaner
et
al.
1985).

The
upstream
distribution
of
sand
shrimp
in
the
San
Francisco
Bay
Estuary
is
limited
by
low
salinity.
In
extensive
collections
from
this
system
between
1976
and
1985,
almost
no
crangonids
were
taken
in
water
of
<
l
ppt
salinity
(
Siegfried
1980,
California
Department
of
Fish
and
Game
1987).

The
response
of
crangonids
to
changes
in
temperature
and
salinity
are
highly
interdependent
(
Khorram
and
Knight
1977a).
Researchers
disagree
about
the
relative
importance
of
each
of
these
factors
in
crangonid
distribution.
Havinga
(
1930)
and
Haefner
(
1976)
suggsted
that
temperature
is
the
most
important
factor.
Krygier
and
Horton
(
1975),
Siegfried
(
1980),
and
Hatfield
(
1985),
indicated
that
salinity
was
the
most
important
factor
determining
sand
shrimp
distribution.
Both
factors
are
important
in
determining
black
shrimp
distribution;
temperature
is
most
important
in
summer
and
salinity
in
the
winter
(
Krygier
and
Horton
1975).

Laboratory
investigations
of
the
interactive
effect
of
temperature
and
salinity
on
the
survival
of
adult
sand
shrimp
indicated
optimum
temperature
and
salinity
ranges
of
14.5­
17.0
"
C
and
18­
20
ppt
(
Khorram
and
Knight
1977a).
The
range
of
temperature
and
salinity
optima
decreased
with
increasing
exposure
times.
The
optimum
ranges
of
temperature
and
salinity
were
8.0­
22.5
"
C
and
13­
25
ppt
for
a
24
h
exposure;
9.5­
21.0
"
C
and
14.0­
24.5
ppt
after
48
h;
and
12­
19
"
C
and
15.5­
22.0
ppt
after
72
h.

Sub&
ate
Little
information
is
available
relating
crangonid
distribution
to
substrate
type.
Crangonids
are
found
on
substrates
ranging
from
mud
to
peat
to
sand
(
Israel
1936;
Carlton
and
Kuris
1975).
They
appear
to
be
particularly
11
suited
to
sand­
mud
substrates
by
being
able
to
nestle
and
bury
themselves
into
the
substratum
using
their
pleopods
and
walking
legs.
Some
crangonid
species
are
reported
from
the
rocky
intertidal
zone
(
Carlton
and
Km­
is
1975).

The
dorsally
flattened
crangonids
usually
spend
days
in
shallow
depressions
in
the
substrate
with
just
the
eyes
above
the
substrate.
In
laboratory
aquaria
this
behavior
occurs
on
sand
as
well
as
on
softer
substrates.
Substrate
selection
by
crangonids
is
undoubtedly
influenced
by
prey
availability
as
well
as
habitat
considerations.
Benthic
prey
is
generally
not
abundant
on
coarse
sediments
in
San
Francisco
Bay
but
is
abundant
and
diverse
on
softer,
more
organic
sediments
(
Nichols
1979).
These
richer
sediments
are
often
the
preferred
substrates
of
crangonids.

Freshwater
Flow
Freshwater
flow
to
estuarine
systems
affects
water
temperature,
salinity,
substrates,
seaward
and
landward
currents,
and
the
distribution
of
potential
prey
and
predators.
During
periods
of
high
freshwater
flows,
seaward
and
residual
landward
currents
are
high
and
salinity
intrusion
is
low.
During
periods
of
low
freshwater
flows,
currents
are
reduced,
high
salinity
waters
intrude
higher
into
the
estuary,
and
water
temperatures
tend
to
be
higher.
These
differences
in
current
patterns
and
temperature­
salinity
regimes
lead
to
important
differences
in
recruitment
and
habitat
availability
for
crangonids
and
their
prey
and
predators.

The
recruitment
of
crangonids
to
the
San
Francisco
Bay
Estuary
appears
to
be
linked
to
variations
in
current
patterns
related
to
freshwater
flow.
Ovigerous
females
occur
in
high
salinity
portions
of
the
estuary
and
in
the
offshore
region
(
California
Department
of
Fish
and
Game
1987).
Larvae
are
released
in
these
regions,
and
since
they
are
generally
in
the
water
column
they
can
bc
entrained
by
surface
currents
moving
seaward.
Crangonid
postlarvae
congregate
near
bottom
and
are
thus
more
likely
to
be
transported
upstream
or
landward
in
the
landward
flowing
lower
layer
of
water
(
Siegfried
et
al.
1978).
The
interaction
of
currents
flowing
seaward
and
landward
thus
determines
recruitment
of
crangonids
to
the
estuary.
&
.

Correlation
analysis
of
annual
abundance
of
sand
shrimp
in
relation
to
freshwater
flow
to
San
Francisco
Bay
indicates
some
highly
positive
relations
(
California
Department
of
Fish
and
Game
1987).
The
annual
abundances
of
postlarvae,
juveniles,
and
all
sand
shrimp
stages
combined
were
positively
correlated
with
freshwater
flows
during
the
period
from
1980
to
1985.
The
abundances
of
early
and
mid­
stage
crangonid
larvae
in
San
Francisco
Bay
were
negatively
correlated
with
freshwater
flow.
This
is
consistent
with
a
life
history
pattern
in
which
larvae
are
planktonic
in
the
upper
layers
of
the
water
column
and
are
carried
offshore
during
high
freshwater
flows.
In
years
of
high
freshwater
outflow,
a
larger
proportion
of
the
reproductive
population
moves
from
embayments
to
the
neat­
shore
coastal
area,
resulting
in
more
larvae
hatched
outside
the
embayments
(
K
Hieb,
California
Department
of
Fish
and
Game,
Stockton,
CA;
pets.
comm.).
The
annual
abundance
of
black
shrimp
is
also
correlated
with
freshwater
inflow,
but
the
relation
is
not
as
strong
as
that
of
sand
shrimp
c
+

(
California
Department
of
Fish
and
Game
1987).
The
annual
abundance
of
blue­
spotted
shrimp,
which
does
not
show
the
extensive
migration
associated
with
development
of
sexual
characteristics,
is
not
correlated
with
freshwater
outflows.

The
volume
of
freshwater
outflow,
and
thus
the
location
of
the
sediment
entrapment
zone
associated
with
the
salinity
gradient
in
the
San
Francisco
Bay
Estuary,
has
been
linked
to
variations
in
the
annual
abundance
and
distribution
of
N.
merce&,
the
principal
food
of
sand
shrimp
in
the
upper
estuary
(
Siegfried
et
al.
1979).
The
distribution
of
sand
shrimp
in
the
upper
estuary
has
in
turn
been
related
to
the
distribution
of
N.
mercedis
(
Siegfried
1980).
Thus,
the
distribution
of
crangonids
can
also
be
affected
indirectly
as
well
as
directly
by
freshwater
flow.
The
distributions
and
abundances
of
many
species
of
fshes
in
the
San
Francisco
Bay
system
are
also
influenced
by
freshwater
flows
(
Armor
and
Herrgesell
1985).
Abundance
of
fishes
appears
to
be
higher
during
years
of
high
freshwater
flows.
Potential
13
predators
as
well
as
the
prey
of
crangonid
shrimp
are
thus
affected
by
freshwater
flow
to
the
San
Francisco
Bay
system.

OtherEnvironmentalRequirements
Other
environmental
factors,
such
as
dissolved
oxygen
concentration,
metals
concentrations,
pesticides,
and
other
agricultural,
municipal,
and
industrial
pollutants
may
affect
the
distribution
and
abundance
of
crangonids.
Khorram
and
Knight
(
1977b)
reported
the
toxicity
to
sand
shrimp
of
Kelthane,
an
organochlorine
miticide
once
commonly
used
on
vegetable,
fruit,
and
grain
crops
in
regions
tributary
to
San
Francisco
Bay.
The
lethal
threshold
was
estimated
to
be
about
100
ppb.
Kelthane
and
its
breakdown
products
bioaccumulate
in
body
tissue
(
Khorram
and
Knight
1977b).
Shrimp
exposed
to
this
pesticide
showed
a
characteristic
sublethal
response,
including
increased
physical
activity,
and
decreased
feeding
and
molting
rates.

Low
dissolved
oxygen
concentrations,
in
combination
with
high
water
temperatures,
are
believed
to
limit
the
occurrence
of
crangonids
in
several
streams
tributary
to
San
Francisco
Bay
(
Israel
1936;
Nichols
1979;
Kinnetic
Laboratories
1983).
Sand
shrimp
were
abundant
upstream
from
San
Francisco
Bay,
into
the
Sacramento
River,
during
years
of
low
freshwater
flows
(
Siegfried
1980).
Crangonid
shrimp
were
not
collected
from
the
San
Joaquin
River
in
Siegfried's
study
(
1980)
even
though
the
temperature
and
salinity
regimes
in
each
river
are
similar.
The
San
Joaquin
River
receives
more
industrial
and
agricultural
effluent
than
the
Sacramento
River,
relative
to
its
discharge.
This
increased
effluent
may
create
water
quality
differences
between
the
two
rivers
that
limit
shrimp
distribution.

e
13
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parasite
Argeia
pauper&
u
(
Crustacea:

16
Isopoda)
on
the
grass
shrimp
Crangon
franciscorum
(
Crustacea:
Crangonidae).
Comp.
B&
hem.
Physiol.
A
Comp.
Physiol.
83:
121­
124.

Nichols,
F.
H.
1979.
Natural
and
anthropogenic
influences
on
benthic
community
structure
in
San
Francisco
Bay.
Pages
409­
426
in
T.
J.
Conomos,
ed.
San
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Bay:
the
urbanized
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American
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the
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of
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Pacific
Division,
San
Francisco.

Overstreet,
R.
M.
1978.
Marine
maladies?
Worms,
germs,
and
other
symbionts
from
the
northern
Gulf
of
Mexico.
Mississippi­
Alabama
Sea
Grant
Consortium
MASGP­
78­
021.
104
pp.

Price,
KS.
1%
2.
Biology
of
the
sand
shrimp
Crangon
septimspinosa
in
the
shore
zone
of
the
Delaware
Bay
region.
Chesapeake
Sci.
3~
244­
255.

Prosser,
C.
L.
1950.
Temperature:
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380
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Prosser,
ed.
Comparative
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W.
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Saunders
Co.,
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P
A
Rathbun,
M.
J.
1904.
Decapod
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northwest
coast
of
North
America.
Harriman
Alaska
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A,
and
C.
Schlieper.
1971.
Biology
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Die
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25:
327
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Scofield,
N.
B.
1919.
Shrimp
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Calif.
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5:
1­
12.

Schmitt,
W.
L.
1921.
The
marine
decapod
crustacea
of
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Univ.
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No.
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470
pp.

Shaner,
S.
W.
1978.
Osmotic
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ionic
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(
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(
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121
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AW.
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ed.
Studies
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bioenergetics,
osmoregulation,
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behavior,
and
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Water
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Paper
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4507.
University
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California,
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Shaner,
S.
W.,
J.
H.
Crowe,
and
A.
W.
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1985.
Long­
term
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to
low
salinities
the
euryhaline
~
ncisc­.
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Crangon
J.
Exp.
Zool.
235~
315­
324.

Siegfried,
CA.
1980.
Seasonal
abundance
and
distribution
of
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fi­
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macrodactyhs
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192.

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1982.
Trophic
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Rathbun;
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89:
129­
139.

Siegfried,
CA.,
AW.
Knight,
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M.
E.
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1978.
Ecological
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the
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San
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1979.
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1978.
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R.
M.,
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W.
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1979.
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17
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D.
F.,
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J.
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1988.
Variation
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8~
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Wahle,
R.
A.
1985.
The
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&

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B.
L.
1970.
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MS.
Thesis,
University
of
Maryland,
Baltimore.
69
pp.

18
REPORT
E;
MENTATION
j
l.
REPoR:
No.
1
Btologtcal
Report
82(
11.125)*
,.
Title
and
Subt*
tls
Species
Profiles:
Life
Histories
and
Environmental
Requirements
of
Coastal
Fishes
and
Invertebrates
(
Pacific
Southwest)­­
Crangonid
Shrimp
i
December
1989
i6.

7.
Author(
a)

Clifford
A.
Siegfried
I
6.
Performing
Or6rmzation
Rept.
Nt
9.
Per(
ormnng
Organtzatlon
Name
and
Address
Biological
Survey
NYS
Museum
and
Science
Service
Albany,
NY
12230
11.
Contract(
C)
or
Grant(
G)
No.

_
_
_
_
__.
12.
Sponsonng
Or6aniration
Name
and
Address
U.
S.
Department
of
the
Interior
U.
S.
Army
Corps
of
Engineers
Fish
and
Wildlife
Service
Coastal
Ecology
Group
Research
and
Development
Waterways
Experiment
Station
Washington,
DC
20240
Vicksburg,
MS
39180
13.
TYIX
Of
Report
6
Period
Coveret
*
U.
S.
Army
Corps
of
Engineers
Report
No.
TR
EL­
82­
4
16.
Abstract
(
Limht:
200
words)

Species
profiles
are
literature
summaries
of
the
taxonomy,
morphology,
range,
life
history,
and
environmental
requirements
of
coastal
aquatic
species.
They
are
prepared
to
assist
in
environmental
impact
assessments.
Crangonid
shrimp
once
were
important
in
an
export
fishery
but
are
now
the
basis
of
a
bait
fishery
in
San
Francisco
Bay.
The
shrimp
are
important
components
of
the
estuarine
system,
serving
as
an
important
food
of
almost
all
sport
and
commercial
fshes
of
west
coast
estuaries.
Spawning
occurs
in
waters
of
~
15
ppt
salinity.
Gvigerous
females
are
found
year­
round;
abundance
peaks
in
spring
and
summer
in
embayments
and
in
winter
offshore.
Eggs
hatch
directly
into
planktonic
zoea
which
require
30­
40
days
to
develop
into
postlarvae.
Larvae
prefer
surface
waters,
while
postlarvae
prefer
bottom
waters.
Larvae
are
exposed
to
predominantly
seaward
currents
and
postlarvae
to
landward
moving
bottom
currents.
Juvenile
crangonids
are
generally
found
in
brackish
to
nearly
fresh
water
but
move
to
more
saline
waters
as
they
mature.
Crangonids
are
opportunistic
predators
of
epibenthic
and
benthic
forms.
Annual
abundance
of
crangonids
in
San
Francisco
Bay
has
been
linked
to
the
volume
of
freshwater
flow
to
the
estuary.
Maintaining
adequate
freshwater
flows
to
the
estuary
to
ensure
successful
recruitment
is
vital
to
maintaining
populations
of
this
important
component
of
the
estuarine
system.

17.
Document
Analysis
a.
Descriptors
Estuaries
Shrimp
Fisheries
b.
Idsntlfisn/
Opcn.
Endcd
Terms
Crangonid
shrimp
Crangon
nigricauda
Crangon
franciscorum
Crangon
nigromaculata
c.
COSATI
Field/
Group
16.
Avrolability
Statement
Unlimited
Distribution
j
19.
Security
Class
(
This
Report)
21.
No.
of
Pales
18
__­
22.
Price
:
See
ANSI­
239.16)
OPTIONAL
FORM
272
(
4­
7
(
Formerly
NTIS­
35)
Departm*
nt
Of
commcrc.
As
the
Nation's
principal
conservation
agency,
the
Department
of
the
Interior
has
responsibility
for
most
of
our
nationally
owned
public
lands
and
natural
resources.
This
includes
fostering
the
wisest
use
of
our
land
and
water
resources,
protecting
our
fish
and
wildlife,
preserving
the
environmental
and
cultural
values
of
our
national
parks
and
historical
places,
and
providing
for
the
enjoyment
of
life
through
outdoor
recreation.
The
Department
assesses
our
energy
and
mineral
resources
and
works
lo
assure
that
their
development
is
in
the
best
interests
of
all
our
people.
The
Department
also
has
a
major
responsibility
for
American
Indian
reservation
communities
and
for
people
who
live
in
island
territories
under
U.
S.
administration.

U.
S.
DEPARTMENT
OF
THE
INTERIOR
FISH
AND
WILDLIFE
SERVICE
TAKE
PRIDE
in
America
UN
ITED
STATES
DEPARTMENT
OF
THE
INTERIOR
FISH
AND
WILDLIFE
SERVICE
National
Wetlands
Research
Center
NASA­
Slide11
Computer
Complex
1010
Gause
Boulevard
Slidell,
LA
70458
