CULTURE METHODS AND RESPONSE OF LABORATORY-REARED PINK SHRIMP
IPUtDALUS JORDANI) LARVAE TO SELECTED SIMULATED ENVIRONMENTAL FACTORS
INFORi4ATION REPORT
.
Michael J Hos ie
Fish C m i ssion o f Oregon
Groundfish and Shrimp Investigations
A p r i l 1975
CULTURE METHODS AND RESPONSE OF LABORATORY-REARED PINK SHRIMP
(PANDALUS JORMIYI) LARVAE TO SELECTED SIf51ULATED ENVIRONMENTAL FACTORS
L i t t l e i s known o f the l a r v a l l i f e of P . jordmri.
Published studies o f
P . jordmri 1arvae have been 1argely r e s t r i c t e d t o descri ptions o f 1aboratory-reared
l a r v a l stages (Hodin and Cox 1967; Lee 1969).
Pearcy (1972) attempted t o c o l l e c t
P . jordani o f f the Oregon coast, w i t h 1it t l e success.
The major objective o f t h i s study was t o r e f i n e c u l t u r e methods f o r shrimp
larvae.
Further objectives were t o ascertain influence o f water current and l i g h t
i n t e n s i t y on survival and behavior of i n i t i a l zoeal stages.
The time period of
l a r v a l emergence was also studied.
METHODS AND WTERIALS
Collecting and Holding Adult Shrimp
I n Late i.tarci.1 1971, ovigerous P. jord7mr.i were caught by shrimp trawl off
Coos Bay, Oregon.
They were held i n 136.5 l i t e r (30 gallon) p l a s t i c containers
and transported t o the Fish Commission o f Oregon marine laboratory a t Newport,
Oregon.
These female shrimp were he1d i n a r e c i r c u l a t i ng seawater aquarium designed
by Lukas (1973).
Each shrimp used t o obtain larvae was removed from the aquarium and placed
i n a 2000-ml beaker containing 1500 m l o f sea water.
Adult shrimp were kept near
the bottom o f the beakers by a large-mesh p l a s t i c screen t o prevent cannibalization
of larvae.
tlli nced fresh cock1e clams (~Zinomdiwnnuttat Z i i ) o r gaper clams
(Treaus capex) were fed t o these shrimp.
Larvae Culture Techniques
Shrinip larvae c u l t u r e methods were patterned a f t e r those of Reed (1969) and
Gaumer (1969, 1970) f o r Dungeness crab (Cmcer magiater) larvae.
Only larvae less than one day o l d and a c t i v e l y swimning near the water surface
were used.
)hen larvae appeared i n beakers, tiley were transferred by large-mouth
pipette t o eittier 250-ml Erlenmeyer flasks containing 200 ml of water or 500-ml
Erlenmeyer flasks f i 1 led w i t h 400 nil of water.
Plastic caps were placed 1oosely
over flasks t o eliminate air-borne contamination.
an a i r 1 ine tllro~qhtops of the caps.
Flasks were aerated by inserting
The aniount of a i r introduced was regulated
so that a cortit~cous,b u t slow, series of b c 2 . 1 5
discharged.
In one experi-
ment, to det.enline ir~lpnrtanceof water aeration of survival , flasks were not
aerated.
A71 rearing containers were placed in a constant temperature water bath initia7ly
maintained a t 10 c 5 0.5 c.
T h i s temperature closely approximated ocean conditions
(9.3 t o 10.9 c surface and 9.5 to 10.0 c near bottom) when the majority of larvae
hatched naturally on the northern Oregon shrimp grounds in idarch 1968 (Robinson and
iljilburn i n press).
llhen sktriap reached zoeal stage 9, the water bath temperature
- 0.5 c) to simulate known ocean floor temperwas lowered and maintained a t 7.2 c (+
atures of 6.1 t o 7.8 c (Lukas and Iiosie 1973) in spring and early sumner.
I t was
assumed t h i s was the time of inetamorphosis.
Sea water was obtained from Yaquina Bay and f i l t e r e d through a Plicroflocpolyvinyl chloride f i l t e r to el ininate particles larger than 5
was suppressed by ultraviolet treatment.
p.
Bacterial growth
Sal inity was maintained a t 30 O/oo,
2
0
1 /oo, by dilutfon w i t h d i s t i l l e d water.
Shrinlp were exposed to 12 hours of flourescent 7ight of 69 t o 73 f t . c from
0600 t o 1800 hours, w i t h complete darkness d u r i n g the r e s t of a 24 hour day.
Light
measurements were made w i ti) a rrlodel 756 Weston il7umi nation meter.
Water in the containers was changed, shrimp fed, development noted, and the
presence of mortalities and exuviae recorded three times per week.
Larvae were
considered dead i f they failed t o respond t o agitation w i t h a sma71 pipette.
were cleaned periodical1.y to prevent algae growth.
Flasks
Larvae were fed newly hatched
n a u p l i i of San Francisco v a r i e t y b r i n e shrimp (Artenria satins) a t a concentration of
15 naupl i i / m l of r e a r i n g water v i a a c a l i b r a t e d automatic p i p e t t e .
E f f e c t s of L i g h t on Survival
The effects of l i g h t on s u r v i v a l was tested by p l a c i n g 59 newly-hatched zoeae
(about l 6 / f l a s k ) i n t o four black painted 250-ml f l a s k s containing 200 m l of water.
Four transparent 250-1111 f 1asks containing 52 newly-hatched zoeae (about 1 3 l f l a s k )
were used as a c o n t r o l .
Effects of Aeration and Lack o f Food on Survival
Experiments were conducted t o a s c e r t a i n e f f e c t s o f a e r a t i o n on s u r v i v a l o f :
(a) unfed newly-i~atched larvae and (b) feeding larvae reared through the zoeal
stages.
I n the f i r s t experiment, s u r v i v a l comparisons were made between two groups
o f 60 unfed recently-hatched zoeae.
They were reared u n t i l death i n e i t h e r aerated
o r non-aerated 500-ml flasks a t 30 zoeae/fl ask.
I n the second experiment, the e f f e c t o f a e r a t i o n on feeding larvae was tested
by p l a c i n g 60 newly-hatched zoeae i n t o two non-aerated 500-ml flasks (30 zoeae/flask)
containing food.
four a w a t e d 500-ml f 1asks containing 120 newly-hatched deeding
zceae (3CISlask) were used as % control.
The maximum number of days newly-hatched zoeae could l i v e without food was
determined i n the unfed 1arvae a e r a t i o n experiment.
Larvae Emergence
To determine number o f Jays necessary f o r a l l eggs t o hatch and t h e s p e c i f i c
time o f larvae emergence during a day, an experiment was conducted using three
ovigerous shrimp.
Each shrimp was placed i n an i n d i v i d u a l 1000-ml beaker.
vations f o r emergent larvae were made each day f o r 24 days.
Each morn-
hours newly-emerged 1arvae ?!ere r m o v d and thei r ntrmher.: rncurded
.
O W -
a t 0800
TO a s c e r t a i n
specific emergence time, beakers were checked during four continuous days a t eight
times per day (0400, 0600, 0800, 1200, 1700, 2000, 2200, and 2400 hours).
Effects of L i g h t Intensity and \dater Current on Behavior
Observations were made on response of f i r s t and t h i r d stage P. jordani zoeae
t o l i g h t intensity and water current u s i n g a l i g h t chamber and a current cylinder
(Figure 1) developed by Gaumer (1970). Only early stage shrimp zoeae were utilized
because Gaumer ( F i s h Comission of Oregon, pers. comm.) found l a t e stages of
1aboratory-reared Oungeness crab 1arvae d i splayed di fferent reactions i n these
containers than ocean-col 1ected 1arvae.
The l i g h t chamber and current cylinder were i n a darkened controlled temperature
mom.
A n ~ e t r i cscale was attached t o the side of each container t o measure 1arval
movement.
Botb containers were f i l l e d with f i l t e r e d and ultraviolet-treated 30
(+ 1 O/oo) sea water maintained a t 12c
O/OO
(2lc).
The l i g h t chamber was v e r t i c a l , plexiglass, 150 cni in height by 15 cm i n
diameter and closed a t both ends.
An adjustable incandescent microscope 1i g h t was
placed a t the top of the chamber. Water and zoeae were introduced through a 2.5 cm
hole a t the top of the chamber.
Zoeae were exposed t o one of s i x l i g h t intensities
(0, 100, 500, 1000, 2000, and 8000 f t . c. ) . The 8000 f t . c . (576 1argleys/day) i s
about equivalent t o the maximum recorded s o l a r energy (622 largleys/day) a t the
ocean surface a t Manairno, British Col umbia (Takahashi and Parsons 1973). L i y h t
intensqties *re
w s u r e d w i t h 4 photqpleter place4 d i r e e t l y above t h water
~
surface.
In darkness, the location of larvae was detemiined by u s i n g a flashlight covered
with red cell ophane.
Newly hatched (1ess-than-one-day-old)
f i rst stage zoeae, three-day-01 d f i r s t
zoeae, and seventeen-day-01 d t k i rd zoeae were used.
Newly hatched zoeae, because
of small s i z e and transparent appearance, were introduced one per t r i a l into the
F i g u r e 1.
L i g h t Chamber (background) and Current Cyl i n d e r ( f o r e g r o u n d )
Used t o Measure Swimming Behavior o f P. jordani Zoeae
l i g h t chamber.
Four t r i a l s were conducted a t each 1ight intensity with a total
sample s i z e of 24.
Older f i r s t stage zoeae an3 third stage zoeae, because of
greater v i s i b i l i t y , were placed into the chamber a t f i v e per t r i 3 l .
One t r i a l was
conducted a t each l i g h t intensity, w i t h a sample s i z e of 30 for each o f these l a t t e r
two zoeal types.
12 minutes.
Location of larvae i n the chamber was recorded a t 5, 7, 10, and
Water was changed a f t e r each t e s t t o avoid a temperature increase.
The cylinder used t o ascertain response of zoeae t o a water current was vertical,
plexiglass, and 110 cm i n height by 10 cm i n diameter, with a base-mounted pump
connected to a sea%ater source.
A vertical upward ~ o v i n gcurrent was used because
Gaumer (1969) reported Oungeness crab placed in
P
horizontal current container d i d
not stay i n the current, b u t instead s e t t l e d t o the bottom and remained there,
possibly because of gravity.
Flow was regulated by a rheostat.
F i r s t and third
stage zoeae were introduced (one p r t r i a l , w i t h f i v e t r i a l s a t each zoeal stage)
througii a 1.5 crn hole into the middle of the chamber, half f i l l e d with water.
The
opening was then closed, the cylinder slowly f i l l e d w i t h water and velocity adjusted
t o a flow of 33.0 mm/sec (0.7 knots).
About 15 seconds were a1 lowed f o r acclimation
of zoeae t o the upward current i n the cylinder.
Position of the larvae was then
recorded a t 'the end of eitller one and two rilinutes or when they l e f t the chamber.
RESULTS
Larvae Culture Techniques
E f f e c t s o f L i g h t on Survival.
Zoeae reared t o 13 days i n l i g h t versus dark
= 0.19<3.84n.s.).
f l a s k s showed no s i g n i f i c a n t difference i n i i ~ o r t ai lt y (x:~. f.
S u r v i v a l was 42.3% and 49.2% r e s p e c t i v e l y f o r l i g h t reared and dark reared zoeae
(Figure 2).
A l l s u r v i v i n g l a r v a e had reached o r passed through t h e second zoeal
stage when t h e experiment was terminated.
I t was more d i f f i c u l t t o determine t i l e a e r a t i o n r a t e and c i l l t u r e zoeae i n t h e
dark f l a s k s .
Effects of Aeration and Lack o f Food on Survival.
Some aerated unfed zoeae
l i v e d t o 16 days w h i l e non-aerated zoeae were dead by 12 days (Figure 3).
However,
r e s u l t s were too s u i l a r and number o f r e p l i c a t e s too few t o j u s t i f y conclusions
about e f f e c t s o f a e r a t i o n on unfed l a r v a e s u r v i v a l .
another.
Unfed zoeae cannibalized one
Hence, some unfed l a r v a e reached t h e second zoeal stage before dying.
Survival o f feeding larvae a t zoeal stage 6 (maximum age 47 days) was 7.0%
and 42.7% r e s p e c t i v e l y i n non-aerated and aerated r e a r i n g conditions (Figure 4).
The experiment on non-aerated feeding zoeae was terminated a t stage 6, as o n l y
4 i n d i v i d u a l s were s t i l l a l i v e .
Larvae Emergencc
Seven, nine, and ten days were required f o r complete hatching o f eggs from
three females (Table 1). Most zoeae emerged w i t h i n a f o u r day p e r i o d from each
female.
Total numbers of zoeae hatched p e r fen~alewere 717, 1083, and 1463
respectively
.
Hatching occurred o n l y between 1700 and 0600 hours, w i t h 77.4% of t h i s emergence
between 2200 and 2400 (Figure 5).
Hatching was usual 1y accompanied by vigorous beating o f t h e female's pleopods,
V)
3
vr
-C rULL
a, cTI
-3
C,
(3 Y
T
c-.l
Is-
!al
C3
0
t
0
I
4
1
2
1
t
I
I
1
&
t
I
3 4
5
6 7
8
9 1 0
(11) (lb) (25) (33) (39) (47)(52) (60)(80) (83)
ZOEAL STAGE
Figure 4.
I
I
I
I1
(92) (93)
MEGALOPAL STAGE
Survival o f Feeding P. jordani Larvae Cultured from Eniergence t o a
Given Stage i n Aerated Versus Non-aerated Flasks, Numbers i n parentheses
a r e maximum ages, i n days, o f each stage.
LARVAE
causing l a r g e numbers of larvae t o be l i b e r a t e d .
\/hen each larvae emerged, i t swam
upward and assurned a v e r t i c a l . p o s i t i o n near t h e t ~ a t e rsurface w i t h i t s carapace
down and t e l s o n up.
Table 1.
ilo pre-zoeae were seen.
tjumber of Days Necessary f o r Zoeae t o Emerge From Eggs o f
Each of Three Ovigerous Female P.iordani.
Day
Female I
T o t a l zoeae p e r fernale
Number o f Emerqent Zoeae
Female I 1
Female III
1,083
Female carapace ?ength(mn1)21.5
E f f e c t s o f t i q l 7 t I n t e n s i t y and Hater Current on Behavior
There were no major d i f f e r e n t i a l 1 i g h t resporises w i t h i n each o f the zoeae
types tested, even though responses were s t a t i s t i c a l l y d i f f e r e n t between t h e t h r e e
ayes o f zoeae used (Table 2 and Figure 6).
No noticeable v a r i a t i o n occurred i n
t h e mean r e a c t i o n t o l i g h t between t h e 5 t o 12 minute observation period.
Individual
shrimp however, showed a wide v a r i a t i o n i n t h e i r response t o l i g h t .
Table 2.
Analysis o f Variance on E f f e c t s o f D i f f e r e n t L i g h t I n t e n s i t i e s on
Behavior o f Newly Hatched and Thi r d Stage P. jordmi Zoeae.
Source o f
Variation
Sun1 of
Squares
df
Treatments
Light Intensity
Zero i t . c. Versus o t h e r ft. c.
Zoeal Stages
Stages x L i g h t s
Residual
0.3229
1.3923
0.2418
5.0116
11
5
1
1
4
42
Total
7.5220
53
-1/
2.5104
0.13758
S i g n i f i c a n t d i f f e r e n c e between zoeal stages a t 1%l e v e l .
Mean
Square
[I!:{
0.1193
F
2.70 n.s.
11.681~
<1 n.s.
(Top)
0
10
@
n
20
a&'
30
w
40
.-
T
50
-
.-
60
-
9
K
5
70
W
u
II
a
W
A
n
80 .-
LL
0
n
z
90
-
100
-
110
-
0
Y
F
P.
W
0
2
!+
-'-
120
,-
130
-
140
-
= ADVAiJCED FIRST STAGE
u
= THIRD STAGE
Q
A
(Bottom) 150
A
L
A
A
A
I
t
t
0
100
590
I
I
1000
2000
5
8000
LIGHT Ii4'IENSITY ( f t.c. )
Figure 6.
i-iean Depth iabintained i n a 150. cm V e r t i c a l L i g h t chamber by Three Types
of P. jordmri Larvae Exposed t o S i x D i f f e r e n t L i g h t I n t e n s i t i e s . L i g h t
Source was atToop o f t h e Chamber.
Newly hatched f i r s t stage zoeae primarily stayed i n tile upper one-fourth of
the ci~arnber near the light source a t mean dcptlis o f 6.1 cnl (1000 f t . c . ) to 45.5 cm
(500 f t . c . ) .
Three-day-old f i r s t stage zoeae swan) t o the bottom of the chamber and
remained either on the bottom or near there.
Seventeen-day-old third stage zoeae
Here more scattered i n the chamber, swinrminy a t a mea:-r depth o f 17.8 cm ( 0 ft.c. )
to 106.8 cm (2000 f t . ~ . ) .
The f i r s t and third stage zoeae subjected to the upward moving 33.8 mm/sec.
current were swept out of the chanber within approximately one minute of release.
First stage zoeae had a mean positive movement of 4 . 2 mm/sec. in the water current
used.
Third stage zoeae were swis~ti~ing
against the current a t a mean velocity of
15.1 nnn/sec.
A1 1 zoeae oriented t)~eri~seives
!with carapace down and telson up and
actively swain against the current.
DISCUSS ION
Results of t h i s study provided no conclusive evidence survival of cultured
newly-hatched larvae differed in dark versus clear flasks.
This suggests t h a t
rearing newly-iiatciied P. jordani in dark containers would not reduce mortal i t y
of f i r s t stage zoeae cultured i n transparent vessels.
However, the 1i ght-survival
experlment was terminated a f t e r only 13 days, due t o other commitments.
Reeve (1969)
and Gaunler (1973) found f o r otiier decapod 1arvae a significantly higher mortal it y
i n dark containers, attributed t o probable increased d i f f i c u l t i e s of capturing food.
Tile higher r a t e of survival of feeding P. jordani zoeae in aerated conditions
showed future 1arvae reari ny in 500-ml Erlenmeyer flasks si~ouldinclude mi ld aeration.
This agrees with Modin and Cox (1967), who briefly mention no success i n culturing
P. jordani zoeae in non-ci rculatiny water.
Results of the aeration experiment on non-fed zoeae suggest laboratory-reared
larvae have a rnaximum of 16 days a f t e r ?mergence t o find food before starving t o
death.
The 7-to-10 day laboratory hatching period of P. jordani larvae i s identical
t o what Price and Chew (1972) reported f o r laboratory held P. pktyceros.
Also,
the 4 day peak of larval emergence from each female i s the same Kurata (1955) found
f o r P. kessZeri.
A large tiurllber of eggs on female I I I (Table 1) hatched on the f i r s t day of
the study, suggesting some of i t s eggs had hatched previously t o collection.
Hatching of P. jordcmi in t h i s study primarily occurred from 2000 t o 2400 hours,
w i tti 77.4% of t h i s fronl 2200 t o 2400.
Peter Rothlisberg (Oregon State University,
Department of Zoology, pers. comw. ) in uncontrolled 1ighting conditions, observed
P. jordani zoeae primarily hatching between 2000 and 0200 hours.
Night time pref-
erence for emergence has also been reported for other Pacific ocean panalid shrimp
(Berkeley 1930, Kurata 1955, Price and Chew 1972).
Fiesul t s o f t h i s l a r v a l emergence study suggests t h a t i n nature P. jordani
zoeae are hatched a t n i g h t .
I t i s probable t h a t n i g h t emergence o f P. jordani
zoeae i s advantageous t o t h e s u r v i v a l o f t h e species.
Newly hatched l a r v a e might
have a greater chance o f avoiding predation i f emergence occurred i n darkness.
Lack o f l a r v a e r e a c t i a n t o varying 1i y h t i n t e n s i t i e s i n the l i g h t chamber
suygest t h a t behaviour patterns were caused by some other s t i m u l i .
Gaumer (1973)
usins Dungeness crab zoeae i n the s a w l iy h t chamber, found s i m i l a r differences
between zoeal stages.
However, u n l i k e t n i s study, he and Cook (1968) found
d i f f e r e n t responses based on l i g h t condition, i.e.,
a p o s i t i v e p h o t o t a c t i c response
t o low l i g h t and a negative response t o high l i g h t .
The mean srqimn~ingv e l o c i t i e s found f o r f i r s t stage zoeae (4.2 mn~/sec) and
t h i r d stage zoeae (15.1 mm/sec), though measured i n the abnormal c o n d i t i o n o f an
upward moving current, are probably about what would be t h e i r maximum v e l o c i t i e s
i n nature, as the larvae a c t i v e l y swan1 i n t o t h e current.
Gaumer (1973), using t h e
same chamber and c u r r e n t v e l o c i t y , reported f i r s t stags Dungeness crab zoeae swam
a t 5.8 mm/sec.
Steur (1910) found t h a t swimming v e l o c i t i e s o f f i v e species o f
European decapod l a r v a e ranged frob1 110.9 mm/scc. f o r PorceZZana s p . t o 22.2 mm/sec
f o r GaZathea strigosa.
He reported i n d i v i u u a l s o f a PandaZus s p . , which were l a t e
larvae, swam a t speeds from 14.9 t o 20.6 mm/sec.
The l a b o r a t o r y swiriin~ing reactions o f f i r s t stage and t h i r d stage P, jordani
zoeae show them t o be weak swimmers, unable t o withstand a c u r r e n t o f 33.8 mm/sec.
This v e l o c i t y i s below o r close t o t h e minimum h o r i z o n t a l c u r r e n t reported over the
continental she1 f i n Oregon waters by previous workers, i n c l u d i n g bJyatt, Burt, and
P a t t u l l o (1972) f o r surface currents; P i 11sbury, Smi ti], and P a t t u l 1o (1970) f o r
midwater currents; and, Gross, Worse, and Barnes (1969) as ell as H a r l e t t (1972)
f o r bottom currents.
She actions o f these currents are not f u l l y understood, b u t
i t i s probable t h a t the i n i t i a l m e a l stages o f P. jordani,
being feeble swimmers,
a r e a t the mercy of whatever water mass they occupy.
I t is t~ypothesizedfronl the r e s u l t s of t h i s and other studies t h a t ~ c e a n i c
transport can be impor-Lant i n a f f e c t ir:g P. jordani 1 arval d i s t r i b u t i o n and survival
I wish t o thank Jack Robinson and J i ~ i9ieehai-r
i
f o r i n i t i a l l y offering encourage-
ment and support.
TIw-oughout t h i s interim study, I was he1 ped by Tom Gaumer who
gave valuable advice on rearing techniques.
Thanks a r e due a l s o t o Lou Fredd nho
did the s t a t i s t i c a l analysis of the data and commented on i t s significance.
I am
especially grateful t o J e r r y 8 u t l e r and Peter Rothlisberg who spent many patient
hours reviewing t h e r~~anuscript.
LTiEKATUilE CITED
Berkeley, A.A.
1930. The post-er!~bryonic development of the conmon panda1 ids of B r i t i s h
Colur,~bia. C o i i t r i ~ . Can. 3iol ., N.S. G(6): 79-163.
Cook, H.L.
1968. Taxononly and c u l t u r e of shritnp larvae. I n Report of t h e Bureau of
Commercial Fisheries Biological Laboratory, ~ a l z s t o n ,Texas Fiscal year
1957. U.S. Fish Wildl. Serv., Circ. 295, p.7-8.
Gaumer, T.F.
1969. Conirollcd rearing of Dungeness crab larvae and t h e influence o f
environmental conditiotls of t h e i r survival. Cormn. Fish. Res. and Dev. Act,
Contract 14-17-0001-1907. Fish Co~nin. Ore. Progress Rep., July 1, 1963 t o
June 30. 1969. 17 p.(Eiimeoyr.)
1970. Controlled raaring of Dungeness crab larvae and t h e influence of
environmental conditions on t h e i r survival. Comnl. Fish. Res. and Dev. Act,
Contract 14-17-0001-2131. Fish Comm. Ore. Progress Rep., July 1, 1969 t o
June 30, 1970. 13 p. (Mirneogr. )
1973. Controlled rearing of Dungeness crab larvae and the influence of
environmental conditions on t h e i r survival. Coml. Fish. Res. and Dev. Act,
Contract 14-17-0001-2325. Fish Comm. Ore., Final Rep., Wov. 16, 1965 t o
June 30, 1971. 43 p. (:.linleogr. )
.
Gross, M.G., 8. Morse and C.A. Barnes
1969. i;iovernent of near-bottom waters on the continental shelf off the
northwestern United S t a t e s . J . Geophysical Sci . 74(28):7044-7047.
Harlett, J.C.
1972. Sediment transport on tile northern Oregon continental s h e l f .
Tiiesis, Oregon S t a t e University. 120 p .
Kurata, 11.
1955.
Bull.
PhD.
The post-embryoni c cievel op111ent of the prawn, PunduZus kessleuri.
Hokkaido reg. Fish. Res. Lab. 12:l-15.
Lee, Y.J.
1969. Larval development of pink shrimp, PandaZus jordani Ratl~bun reared
i n laboratory. i.1. Sc. Thesis, Univ. Eashington, 62p.
Lukas, G.
1973. An evaluation of methods f o r determining movement of shrimp. Phase 11:
Shrimp holding studies. Comm. F i s h . Res. and Dev. Act, Contract 14-17-0001-2383.
Fish Conlm. Ore., Final Rep., July 1, 1970 t o June 30, 1971. 11 p. (Mimeogr.)
Lukas, G. and i4.J. Hosie
1973. Investigation of t h e abundance and benthic d i s t r i b u t i o n of pink shrimp
o f f the northern Oregon coast. Comrn. Fish. Res. and Dev. Act, Contract
14-17-0001-2323.
F i s h Comm. Ore., Final Rep., July 1, 1969 t o June 30, 1970.
45 p. (i4imeogr. )
Modin, J.C. and K.W. Cox.
1967. Post-embryoni c devel opment of 1aboratory reared ocean shrimp,
PandaZus jordani Rathbun
Crustaceana 13(2) : 197-219.
.
Pearcy , bi.
G.
1972. Distribution and die1 changes in the behavior of pink shrimp,
PandaZus jorduni, off Oregon. Proc. ilat. S h e l l f i s h e r i e s Assoc. 62: 15-20.
Pillsbury, R . D . , R.L. Smith and 3 . 6 . P a t t u l l o .
1970. k compilation of observations front moored current meters and thermographs,
Vol . 111. Oregon continental s h e l f , May - June 1967, Apri 1 - September 1968.
Dept. of Oceanography, Oregon S t a t e University, Corvall is, (Data Rep. 40,
Ref. 70-3).
Price, V.A. and K.W. Chew.
)
1972. Laboratory r e a r i ng of spot s hrinip 1arvae (~andazusp ~ u t ~ c e r o sand
descriptions of stages. J. Fish. Res. Board Can. 29(4): 413-422.
Heed, P.t!.
1969. Culture n~ethodsand e f f e c t s of temperature and s a l i n i t y on survival
and growth of llungeness crab (cancer magister) larvae i n the laboratory,
J. Fish. Res. Board Can. 26(2) : 389-397.
Reeve, M.R.
1969. The laboratory culture of the prawn Palaemon serratus. Fishery Invest.
Lond. Ser. 2, 26(1): 38p.
Robinson, J.G.
1971. The d i s t r i b u t i o n and abundance o f p i n k shrimp (~andaZusjordcmi) o f f
Oregon. Fish Coml. Ore., Invest. Rep. 8 , 48 p.
Robinson, J.G. and Gary Milburn.
I n press. The v e r t i c a l d i s t r i b u t i o n and die1 m i g r a t i o n o f p i n k shrimp
(PandaZus jordmni) o f f Oregon. F i s h Comm. Ore. Research Reports.
Steur, A d o l f
1910. Planktonkude.
L e i p z i g und R e r l i n , B.G.
Takahaski, N. and T. Parsons
1973. B i o l o g i c a l Oceanographic Processes.
I;Jyatt, B.,
Teubner.
723 p.
Pergamon Press, l-le\br York.
186 p.
i.J.V. B u r t and J.G. P a t t u l l o
1972. Surface currents o f f Oregon as determined from d r i f t b o t t l e returns.
J. Phys Oceanography 2 (3) : 286-293.
.