Content uploaded by Tsukasa Mori
Author content
All content in this area was uploaded by Tsukasa Mori on Mar 12, 2014
Content may be subject to copyright.
GROWTH OF PACIFIC SAURY,
COLDLABIS SAZRA,
IN
THE NORTHEASTERN AND NORTHWESTERN PACIFIC OCEAN'
YOSHIRO
WATANABE,~
JOHN
L.
BUILER;
AND
TSUWSA
Mom4
ABSTRACT
Growth
of the Pacific
saury,
Cololabis
saira,
from the northeastern and northwestern Pacific Ocean was
studied using otolith
growth
increments. We found
that
growth
of Pacific sauries
from
the western Pacific
was higher than that from the eastern Pacific. Assuming that otolith
growth
increments are deposited
daily, average
growth
rates
from
hatching
up
to
1
year old were 0.62 mndd in the eastern and 0.85 mndd
in the western
Pacific.
Because
the
growth
rate
changes at around
100
mm.
two curves were
used
to
model the
growth
of
Pacific
saury
in the western Pscific: one for fish
up
to
100
mm
and the other for
fish larger than
100
mm.
Based
on counts of daily increments, Pacific sauries may
be
short lived. The
oldest specimen examined was only
14
months old.
The Pacific
saury,
Coblabis
saira
(Brevoort),
is
distributed throughout the North Pacific Ocean and
is one of the most important commercial fishes in
the northwestern Pacific. The average annual catch
of Pacific
saury
in Japan has been approximately
200,000
t
(metric tons) in the last 20 years (Statistics
and Information Department, Japan 1985). The
catch has varied by an order of magnitude in the
last
20
years from
a
minimum of 63,000
t
in 1969
to
a
maximum of 406,000 tin 1973. Fluctuation in
stock size is
a
major factor in catch variability
although economic factors such
as
fish price may
also affect total landings. However, the causes of
stock fluctuation in the western Pacific remain
unknown. In the eastern Pacific, the Pacific
saury
has not been exploited but is recognized
as
a
poten-
tial
fishery resource (Ahlstrom 1968; Smith
et
al.
1970).
Investigations of the Pacific
saury
have mainly
been devoted to such subjects
as
systematics, abun-
dance, distribution, migration, and formation of
fishing ground in relation
to
oceanographic condi-
tions (e.g., Hubbs and Wisner 1980; Smith et al.
1970; Odate 1977; Fukushima 1979; Sablin and
Pavlychev 1982; Gong 1984). Age determination and
growth, however, remain controversial (Hatanaka
1955;
Hotta
1960; Novikov 1960; Sunada 1974;
Kim
'Contribution
No.
429 from Tohoku
Regional
Fisheries
Research
Laboratory.
ZTohoku Regional Fisheries Research Laboratory, Fisheries
Agency, Shiogama, Miyagi
985,
Japan.
'Southwest Fisheries Center La Jolla Laboratory. National
Marine Fisheries Service,
NOAA,
P.O.
Box
271, La Jolla, CA
92038.
'Faculty of Fisheries, Hokkaido University, Hakodate. Hokkaido
041.
Japan.
Manuscnpt accepted March 1988.
FISHERY
BULLETIN:
VOL.
86.
NO.
3.
1988.
and Park 1981), notwithstanding their critical im-
portance for fish stock assessment.
The discovery
of
daily increments in the otoliths
of fishes (Pannella 1971) has made
it
possible to
estimate
age
and growth of larval and juvenile fishes
accurately. Daily increments have been used
to
age
many species of fishes (Jones 1986). Nishimura et al.
(1985) reported the presence of growth increments
in Pacific saury otoliths observed by scanning elec-
tron microscopy and suggested that it is possible to
estimate
age
and growth of Pacific
saury
by using
daily increments in the otolith. The purpose of this
paper is
to
determine the age of Pacific sauries from
the eastern and western North Pacific using daily
increments and
to
compare
the
growth
rates
in these
areas.
MATERIALS AND METHODS
We read otoliths of
75
Pacific sauries from the
northeastern and 172 from the northwestern Pacific
Ocean. Details of sampling and methods of reading
otoliths are summarized in Table
1
and Figure
1.
Additional samples from the western Pacific were
used to determine the relation between otolith size
and fish length. Fish from the eastern Pacific were
futed and preserved in 80% alcohol after capture,
and those from the western Pacific were stored
frozen and thawed when processed. Because speci-
mens frequently have damaged upper jaws, knob
length (the distance from the tip of the lower jaw
to
the posterior end of the muscular
knob
at
the base
of a caudal peduncle)
is
the standard measure of
body size in Pacific
saury.
All body lengths in this
paper are knob length.
489
FISHERY BULLETIN: VOL.
86.
NO.
3
TABLE
1
.-Collection records
of
saury
samples
from the eastern
(d
1-10)
and the western
(#
11-22)
North
Pacific.
N
-
Neuston
net; G
-
Gill
net;
D
=
Dip net.
Sample
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
Date
80
06
29
80
06
29
800629
81 1026
81 1027
81 1028
81 1029
81 1030
81 11 01
81 11 02
840526
84
06
02
84
07 16
84
10
06
850520
85
05
21
85
05
23
85
05
24
85 05 29
Location
Lat.
Long.
36O02'N 124O04'W
36O07" 123O55'W
36'07N 123O45'W
46O36'N 127O49.W
47O19" 126OO9'W
47O20" 124O30'W
46O21'N 127O38'W
46O38'N 125*54'W
45O21'N 127O38'W
45O38'N 124O51'W
37O01'N 164O02'E
37OW" 15S000'E
42O00" 172"WE
43OWN 153O14'E
MWO'N 150°00'E
38O15" 149O59E
38'30" 152'00'E
38'00" 152°00'E
38O45'N 156°00'E
No.
10
10
10
5
6
12
1
14
4
3
19
114
7
5
3
2
2
9
3
Size range
(KnL
mrn)
19.8-
66.0
15.3-
96.0
47.8-235.0
38.0-142.0
23.9-216.0
20.6- 85.0
27.0- 71.0
70.0-206.0
145.0-230.0
19.8-109.0
209.0
38.0-125.0
213.0-282.0
300.0-330.0
21.0- 69.5
8.3- 12.5
26.5- 45.5
29.0- 85.0
16.0- 33.5
Gear
N
N
N
N
N
N
N
N
N
N
G
N
G
D
N
N
N
N
N
Micro-
scope
LM
LM
LM
LM
LM
LM
LM
LM
LM
LM
SEM
SEM
SEM
SEM
LM
LM
LM
LM
LM
-
20
850529 38O30" 156"WE 2 27.0- 33.0 N
LM
21 850529 38O15" 156°00E 3 24.6- 65.0 N
LM
22 850530 38WO'N 156°00E 3 31.5- 68.5 N
LM
..........
7------
11
'
*I
.........
+
.........
0,
............
'3.
I
-
.........
+-
170'E
50'
40'
3
0'
FIGURE I.-Locations
of
Pacific saury collection in the
North
Pacific. Figures
by
490
WATANABE ET AL.:
GROWTH
OF
SAURY
Sagittae were dissected out from fish and left to
dry
after removing tissues and membranes. We used
a dissecting microscope with
a
polarizing filter to
dissect otoliths from small larvae and juveniles. The
otoliths were read either by light microscopy (LM)
or by scanning electron microscopy (SEM). Otoliths
that were to be read by LM were mounted in
EUKITTj after dissection. Otolith radius was
measured from the focus to posterior margin and
the increments were counted along the same tran-
sect using the otolith reading system, which was
developed by the Southwest Fisheries Center of
the National Marine Fisheries Service, NOAA, and
which consists of a light microscope, a video
monitor,
a
micro-computer, and
a
digitizer (Methot
1981).
For SEM, otoliths were mounted in epoxy or
methacrylate resin. The otolith radius was measured
from the focus to the posterior margin with an
6Reference to trade names does not imply endorsement by the
National Marine Fisheries Service. NOAA.
dots indicate sample numbers in Table
1
optical comparator. The otoliths were ground
oblique to the sagittal plane parallel to the long
axis of the otolith in order to have a flat plane
through the otolith nucleus. The polished surface
was washed in xylene, using an ultrasonic washer,
then dried and etched for 50 seconds with 0.2M
EDTA-2Na (disodium
ethylenediaminetetracetic
acid). The etched surface was coated with palladium
platinum and observed under an SEM (JSM-25)
at
15
kV.
The three authors of this paper read saury oto-
liths independently: the senior author read fish from
the western Pacific up to 85 mm by LM, the second
author read otoliths from the eastern Pacific by LM,
and the third read otoliths from the western Pacific
larger than
38
mm by SEM.
To
confirm that we
were all interpreting the same structure
as
growth
rings by SEM and LM, we compared 50 data points
read by SEM and 14 points read by LM for west-
ern Pacific sauries between
38
and 85 mm. The
distribution of increment number versus knob length
was the same. We
also
checked for possible biases
for the
two
readers using LM by having each read
the same set of otoliths independently.
RESULTS
The nucleus of
a
Pacific saury sagitta is approx-
imately 20 pm in diameter and is composed of four
to
six
small dense bodies which appear
to
be assem-
blages of calcareous spherules
(Fig.
2a). These dense
bodies are separated from one another and each
is
surrounded by a small concentric ring.
We observed the nucleus areas of otoliths from
Pacific sauries collected in the western Pacific in
1985 (sample
#
15-22) and found that most of
them had
a
distinct ring of about
27
pm in radius.
Between the nucleus and this distinct ring, four
(or five) indistinct growth rings were detected (Fig.
2b).
We measured knob lengths of 27 larval and juven-
ile Pacific sauries before freezing and after thaw-
ing, and found that the ratio of these two measure-
ments was 0.997. There was virtually no shrinkage
by freezing and thawing. Theilacker (1980) found
that preservation of larval northern anchovy,
Engraulis
mordax,
in 80% alcohol did not cause
additional
shrinkage of the body
after
net treatment.
Thus, knob lengths after both
80%
alcohol preser-
vation and freezing are comparable to each other,
and this measurement corresponds
to
the size after
net treatment. Since shrinkage factors by net
treat-
ment are not known for the
saury,
lengths
are
un-
corrected for net shrinkage.
491
FISHERY
BULLETIN:
VOL.
86.
NO.
3
FIGURE
2.-Light micrographs
of
Pacific saury sagittae.
a)
Otolith nucleus composed
of
5
or
6
separate dense bodies with surrounding
cores.
b)
Assumed
4
embryonic and
1
hatching (arrow) rings.
The daily periodicity of growth increment forma-
tion in the Pacific
saury
has not been verified. For
that reason we plotted the number of increments
versus knob length instead of age versus length. We
used the Laird-Gompertz equation
to
describe the
relations of increment number and length
as
growth
curves for both the eastern and western Pacific
saury. Hatching size of artificially fertilized and in-
cubated Pacific
saury
from the western Pacific was
reported to be 7.19 mm in average live total length
(Yusa 1960). From the drawing of
a
newly hatched
larva in
Yusa's
paper, we estimated live knob length
to be 6.60 mm. Shrinkage factors of northern an-
chovy in the size range from 6.00 to 7.99 mm were
0.90 for
a
5-min net treatment and 0.85 for
a
10-min
net treatment (Theilacker 1980). Using these values,
the capture size of a newly hatched larva of Pacific
saury after
a
5-min net treatment was estimated to
be 5.95 mm and a 10-min treatment to be 5.61 mm.
We fixed the hatching size from 5.85 to 5.95 mm
in the growth curve, because the Pacific saury lar-
vae at this size are in
a
more advanced develop-
492
mental stage and shrank less by net treatment than
northern anchovy.
The resulting growth equation for the eastern
Pacific saury was
KnL
=
5.85 exp((0.0427/0.115)(1
-
e'-O
011q'-5))))
and the equation
for
the western Pacific saury was
KnL
=
5.95 exp((0.0504/0.0128)(1
-
e(-'
0128('-5))))
where
KnL.
is a knob length in mm and
Z
is
the total
number of increments observed in an otolith. The
term,
I-
5, indicates that five increments were pre-
sumed to have been present
at
hatching. Data from
the western Pacific saury appear to consist of two
curves separated around
100
mm in
KnL.
Two
Laird-Gompertz curves fit much better than one
curve. The intersection of the
two
curves was at 114
increments and
100
mm. The growth equation for
fish smaller than
100
mm was
WATANABE
ET
AL.:
GROWTH
OF
SAURY
KnL
=
5.90 exp((0.0865/0.0293Xl
-
e(-o.029q1-6))))
and for fish
larger
than
100
mm
KnL
was
KnL
=
3.01 exp((0.0592/0.0126)(1
-
e(-0.0126(r-5))))
The estimated mean square error,
215.7,
of the two
curves was smaller than that for
a
single curve,
351.7.
The two-curve model fits much
better
for the
smaller size range up to
100
mm. The estimated
mean square
error
of the two-curve model for this
size range,
75.7,
was much smaller than that of the
one-curve model,
240.8.
The growth
rate
of Pacific
saury
in
the
eastern
Pacific was slower than that in the western Pacific
(Figs.
3,4).
The knob length of
saury
in the
eastern
Pacific would
be
about
75
mm
at
100
rings,
170
mm
at
200
rings, and
220
mm
at
300
rings, whereas in
the western Pacific knob length would be about
100
mm
at
100
rings,
230
mm
at
200
rings, and
300
mm
at
300
rings. Assuming that the rings
are
formed
daily, overall growth
rates
of the first one year
of their life were
0.62
mdd and
0.85
rndd for
2oo
t
n
E
E
v
the eastern and western Pacific
Ocean,
respectively.
The
largest specimen examined was
330
mm from
the western Pacific, which
had
328
increments, and
the oldest fish, also from the western Pacific, which
measured
320
mm, had only
418
increments. These
fish would be classified
as
very large or large by
Novikov's categories of fish size composition
(Novi-
kov
1960,1973).
The largest fish examined from the
eastern
Pacific was
235
mm and had
241
incre-
ments. Hughes
(1974),
however, reported larger fish
from the
eastern
Pacific.
In Pacific
saury
from the western Pacific, the rela
tion between otolith radius in pm
(OR)
and knob
length in mm
(KnL)
was linear on logarithm-
logarithm coordinates (Fig.
5).
The equation com-
puted by the geometric mean regression (Ricker
1973)
was
In (OR)
=
2.33
+
0.749
In
(KnL)
(r
=
0.979).
The otolith radius
at
hatching
(5.9
mm
KnL)
cal-
culated by this formula was
38.9
pm, which was
12
pm larger than the radius of the presumed hatch-
ing ring.
A
100
m
e
Y
50
I
--I
,-
KnL:
KNOB LENGTH
I:
NUMBER
OF
INCREMENTS
01
I
I
I
I
0
50
100
200
300
NUMBER
OF
INCREMENTS
FIGURE
S.-Crowth
curve of the eastern Pacific
saury.
493
FISHERY
BULLETIN:
VOL.
86,
NO.
3
300
n
E
200
E
I
Y
6
5
rn
4
*
100
J
50
0
0.0504
(l-e(-O.Ol
28(1-5)))
---
KnL
=
5.95e
0.0128
0.0865
(1-e(-0.0293(1-5)))
KnL
=
5.90e
o.0293
I
)114
KnL
=3.0le
0.0126
0.0592
(l-e(-O.O
126(1-5)))
KnL KNOB LENGTH
I:
NUMBER
OF
INCREMENTS
50
100
200
300
400
NUMBER
OF
INCREMENTS
FIGURE
I.--Crowth
curve
of
the
western
Pacific
saury.
DISCUSSION
The microstructure of otolith growth increments
of the Pacific
saury
is similar to that of daily incre-
ments in some other fishes (Nishimura
et
al. 1985).
Thus, the following discussion is based on the
assumption that
the
increments
are
daily growth
rings. A rearing experiment of larval
sauries
is
under way
in
the senior author's laboratory
to
verify
daily periodicity of the increment formation.
Formation of
a
few embryonic growth rings or
a
lamellar structure has been reported in California
grunion,
Leuresthes tauis,
(Brothers
et
al. 1976);
mummichog,
Fundulus heteroclitus,
(Radtke and
Dean 1982); and walleye pollock,
Therugra
chalco-
gmmmu,
(Nishimura and Yamada 1984). Radtke
and Dean (1982) mentioned
that
deposition of
growth rings in the embryonic stage might
be
494
related
to
a
long incubation period. Pacific
saury
has
a
long incubation period-about 17 days under
13.5"-15.7"C
(Yusa
1960). At this temperature, eye
pigmentation begins 7 or
8
days before hatching,
and pectoral fins show constant movement from
5
or
6
days before hatching
(Yusa
1960). Notochord
flexion occurs about midway through embryonic
development
at
14"-22"C (Uchida et
al.
1958).
Thus
saury
is more advanced
at
hatching than killifish
based on the embryonic development of killifish
reported by Armstrong and Child (1965). Observ-
ing the central
area
of otoliths, we found four faint
rings and
a
dark ring immediately outside of those
rings. We assumed therefore that the four faint
rings
are
embryonic rings and the dark ring is the
hatching ring. This assumption may be confirmed
by examining otoliths of late embryos and newly
hatched larvae of
saury.
WATANABE
ET
AL.:
GROWTH
OF
SAURY
1000
r
500
I
I-
-
$
loo
0
In
OR
=
2.33
+
0.749
In KnL
r
=
0.979
-
OR:
OTOLITH
RADIUS
KnL:
KNOB
LENGTH
I
I
I
I
I
I
10
50
100
200
300
KNOB LENGTH
(mm)
FIGURE
5.-Knob
length
and
otolith
radius
relationahip
of
the
western
Pacific
saury.
Previous studies on age and growth of the Pacific
saury have
based
on annuli on scales andor otoliths.
Sunada
(1974)
found five age groups in Pacific
sauries off southern Oregon, California, and Baja
California. Mean fork lengths of
age
groups were
171, 220,246, 270,
and
268
mm for age
0,
1, 2, 3,
and
4,
respectively. Hughes
(1974)
examined age
composition of
5,248
sauries
collected in waters
off
California up
to
Vancouver Island. He found spring-
and autumn-born
fish
in
his
samples, but little differ-
ence was noted
in
growth
rates
between two groups.
Approkte
knob
lengths of
1.0-
to
5.0-year-old fish
were
180,230,255,290,310
mm. The growth
rates
given in these two papers
are
not very much dif-
ferent. The saury grows
at
0.5-0.6
mm/d up to
1
year old, which is almost equal to our growth rate
in the eastern Pacific,
0.62
mm/d. Hatanaka
(1955)
found five
age
groups in the western population of
the saury,
0-4
years old, and estimated mean body
length of age groups
to
be
80,160,230,265
mm for
1-
to 4-year-old fish, respectively. Novikov
(1960,
1973)
divided sauries captured in autumn into five
size groups, very small
(-
200
mm), small
(201-240),
medium
(241-290),
large
(291-320),
and very large
(321
+),
and assigned the small, medium, and large
to
1-, 2-,
and 3-year-old with maximum 5-year-old
fish.
A different model of Pacific
saury
growth in the
western Pacific was proposed by
Hotta
(1960)
based
upon
a
hypothesis of
two
subpopulations. He separ-
ated
the
saury into springspawning and autumn-
spawning populations based upon the observations
of
fish
size composition, scales, otoliths, and num-
bers
of vertebrae. He assigned
four
ages
of
half
year
intervals,
1.0, 1.5,2.0,2.5
years old,
to
fish
210-240,
260-280,290-300,
and
310-330
mm, respectively.
The growth
rate
up
to
1
year was
0.6-0.7
mxdd.
Kim
and Park
(1981)
examined Pacific sauries from
Korean waters and found four size groups of four
different
ages
of half-year intervals
as
well. They
presented two growth models for each of two sub-
populations, spring and autumn spawning, based
upon the von Bertalanffy equation. The
sizes
at
ages
were almost identical
to
those of Hotta
(1960).
How-
ever, the hypothesis of two saury subpopulations in
the western Pacific is not supported by electro-
phoretic analyses
of
genetic separation (Numachi
1971;
Hara et al.
1982).
The average growth rate of the western Pacific
saury in this paper was
1.1
mdd from
0
to
8
or
495
FISHERY BULLETIN:
VOL.
86.
NO.
3
Pacific may differ from year-to-year due
to
environ-
mental factors and may result in changes in size
composition of the fish. Between
1968
and
1972,
mean knob length of exploited sauries in the west-
ern Pacific was
170-250
mm, whereas in the
1980’s
the major mode in the size composition was
290-310
mm
(S.
Kosaka6). This increase could have been
due to an acceleration of growth rate or
a
shift of
spawning season
to
early months
or
both in recent
years. The high growth rate of western Pacific
saury
presented in this paper has come from specimens
collected in
1984
and
1985.
The growth rate in the
late
1960’s
and early
1970’s
may have been lower
than that presented in this paper. Investigation of
the interannual variation in growth
rates
using daily
increments would distinguish between these two
hypotheses.
We used three different gears to collect Pacific
saury samples in the western Pacific. Knob lengths
of sauries collected were from
8.3
to
125
mm by ring
net,
145
to
282
mm by
gill
net, and
300
to
330
mm
by stick-held dip net. Sauries of
125-145
mm might
not be available either to the ring net or to the gill
net. Further, the ring net may select small juveniles
of
a
cohort in the size range over
100
mm, and this
may have produced the two growth curves. This
problem needs to be examined further with data on
gear selectivity.
We do not know how long Pacific sauries survive
after becoming adult. The oldest specimen in our
sample was about
14
months old
after
hatching
(418
increments). The largest saury aged
(330
mm),
which had
328
growth rings, is close to the max-
imum size. Although the maximum known length
of the Pacific saury was reported to be about
400
mm (Hubbs and Wisner
1980),
the
largest
fish ex-
ploited in Japan is about
340
mm. Therefore, the
lifespan of the Pacific saury is about one year in the
western Pacific. Our results
are
more consistent
with those of Kosaka
(1979)
who found two age
groups
(0
and
1
year) than those (Sablin
1979)
who
found three age groups
(0,
1,
and
2
years).
In Japan, fishing efforts of the Pacific saury is
regulated by fishing season
as
well
as
by the num-
ber
of fishing boats. The fishing season
starts
in mid-
August. Pacific sauries hatched in the main spawn-
ing season
are
about
250
days old
at
this time of year
(approximately
270
mm) and are growing at the
rate
of
0.8
mm/d. Thus
a
2-wk
postponement
at
the
beginning of fishing season would result in an
11
mm
(10-15
g
body weight) increase
of
average fish
9
months old. It
was
still faster than Novikov’s
growth
rate
of the corresponding age period
(0.83
mdd), which
was
the highest
rate
of
all
the previous
reports. The fish would become
316
mm in one year
according to our model.
Support for the fast growth
rate
of Pacific saury
presented in this paper in the western Pacific can
be found in rearing experiments. Hotta
(1958)
reared young sauries caught by a
set
net. He reared
them in
a
crawl and
fed
them minced anchovy and
mackerel twice
a
day. Young sauries
116
mm in
mean length became
172
mm in the rearing period
after
72
days. The growth
rate
was
0.78
mm/d. The
sauries fed
three
times
a
day grew
130-143%
faster
than the group fed twice
a
day. Thus growth rates
of young sauries may
be
higher than
1.1
mm/d
(0.78
x
1.4
mm/d) when food is readily available. Our
growth
rate
of sauries in
this
size range was approx-
imately
1.5
mm/d in the western Pacific.
For
Atlan-
tic saury,
Scomberesox
saurus
scombroides,
reared
by Brownell
(1983),
the average growth rate of the
larvae was
0.62
mm/d from hatching
(7.5
mm
SL)
to 47-day old
(36.8
mm
SL).
The growth
rates
of
Coblabis
saira
in
a
corresponding period were
0.48
mm/d in the
eastern
Pacific and
1.0
mmld in the
western Pacific.
Our
results indicate that the growth
rate
of Pacific
saury in the western Pacific is much higher than in
the eastern Pacific. This could be due to
a
differ-
ence in food availability between the
two
areas.
However, mean zooplankton standing stock in
1951-66
was
34.8
g/mZ in the California Current
region in June (Smith and Eppley
1982),
whereas
that
of
Kuroshio water off southern Japan was
4.7
g/m2 and of Oyashio area
off
northern Japan was
25.7
g/m2 in May
to
July (Odate
1986).
Thus, differ-
ences in zooplankton standing stock do not explain
the difference in growth
rates.
On the other hand, there seems
to
be
a
reasonable
explanation for the reacceleration of growth
rate
at
around
100
mm in the western Pacific saury. The
western sauries hatch out mainly in offshore water
of the Kuroshio Current
Oat.
31-33’N)
off
Japan
in winter. They migrate north to the Oyashio area
(up to
46-50’N)
where copepods
are
highly avail-
able. Young and adult sauries feed actively and gain
fat. They are in the northward migration stage in
early summer when they
are
about
100
mm, and are
moving from poor Kuroshio water to rich Oyashio
water (Fukushima
1979).
High zooplankton stand-
ing stock in the Oyashio water and
its
derivatives
might be responsible for the reacceleration of
growth rate in fish older than
100
days.
The growth rates of Pacific saury in the western
496
‘S.
Kosaka,
Tohoku Regional Fisheries Research Laboratory,
Fisheries Agency, Shiogama,
Miyagi
985,
Japan,
pers.
commun.
WATANABE ET
AL.:
GROWTH
OF
SAURY
length. The biomass and yield of Pacific
saury
need
to
be
reestimated based on faster
growth
rates
presented here.
ACKNOWLEDGMENT
We thank A. Nishimura of Hokkaido Salmon
Hatchery;
J.
Yamada of Hokkaido University; R.
Lasker,
S.
Tsuji, and N. C.
H.
Lo of Southwest
Fisheries Center of National Marine Fisheries
Ser-
vice (NMFS), NOAA; and
S.
Kosaka and T. Wata-
nabe of the Tohoku Regional Fisheries Research
Laboratory for valuable suggestions and reading the
manuscript. We
also
thank A. W. Kendell
Jr.
of the
Northwest and Alaska Fisheries Center, NMFS,
NOAA, for providing
us
with some specimens from
the
eastern
Pacific, and
D.
Abramenkoff of the
Southwest Fisheries Center for help in reading Rus-
sian papers. Y. Watanabe would like to thank the
Science and Technology Agency of Japan for fund-
ing his stay
at
the Southwest Fisheries Center.
LITERATURE CITED
AHLSTROM,
E.
H.
1968.
An evaluation of the fishery resources available to
California
fishermen.
In
The future
of
the
fishing
industry
in
the
United
States,
p.
65-80.
Univ.
Wash.
Publ. Fish.,
New
Ser.
IV.
ARMSTRONG,
P.
R.
AND
J.
S.
CHILD.
clitus.
Biol. Bull.
128143-168.
1965.
Stages
in the normal development of
Fundulus
hew
BROTHERS,
E.
B.,
C.
P. MATHEWS,
AND
R.
LASKER.
1976.
Daily growth increments in otoliths from larval and
adult fishes. Fish. Bull.. US.
74:l-8.
BROWNELL, C. L.
1983.
Early growth rate and feeding of a
small
group of
laboratory-reared
saury,
Scomberes?
saurus
scombroides
(Pisces: Scomberesoeidae).
S.
Afr.
J.
Mar. Sci.
1:245-248.
1979.
Synoptic analysis of migration and fishing conditions
of saury in
the
Northwest Pacilic Ocean. (In Jpn.) Bull.
Tohoku Reg. Fish. Res. Lab.
41:l-70.
FUKUSHIMA.
S.
GONG, Y.
1984.
Distribution and movements
of
Pacific saury,
Cololubis
saira
(Brevoort). in relation to oceanographic conditions
in
watersoff Korea. Bull. Fish. Res. Dev. Agency
331-156.
1982.
Genetic study on population structure of Pacific saury.
(In Jpn.) Bull. Tohoku Reg. Fish. Res. Lab.
45:l-18.
HATANAKA.
M.
1955.
Biological studies on the population
of
the saury,
Cob
labia
saira
(Brevoort). Part
I.
Reproduction and growth.
Tohoku
J.
Agric. Res.
6:227-269.
HARA.
M.,
A.
KIJIMA,
AND
Y.
FUJINO.
HOTTA, H.
1958.
On the growth of the young saury,
Cololabis
saira.
in
the rearing experiment. (In Jpn.) Bull. Tohoku Reg. Fish.
Res. Lab.
11:47-64
1960.
On the analysis of the population of the saury
(Colo-
labis
saira)
based on the scale and the otolith characters,
and their
growth.
(In Jpn.) Bull. Tohoku, Reg. Fish. Res.
Lab.
1641-64.
Hvses,
C.
L.,
AND
R. L. WISNER.
1980.
Revision of the sauries
(Pisces,
Scomberesoeidae) with
description of two new genera and one new
species.
Fish.
Bull..
US.
77:521-566.
HUGHES,
S.
E.
1974.
Stock composition, growth, mortality, and availability
of Pacific saury,
Cololubis
saira,
of
the northeastern Pacific
Ocean. Fish. Bull., U.S.
72:121-131.
JONES,
C.
1986.
Determining age of larval fish with the otolith incre-
ment technique. Fish. Bull., US.
84:91-103.
KIM, Y.
M..
AND
Y.
J.
PARK.
1981.
A study on the growth
of
saury,
Cololabis
saira
(Bre
voort), based on the length composition in the Korean
waters. (In Korean) Bull. Fish. Res. Dev. Agency
27:
59-70.
KOSAKA,
S.
1979.
Sanma no nenrei seicho ni
tsuite
(Age and growth of
saury). (In Jpn.) Rep. Japan-Soviet
Coop.
Res. Meet.
Saury Scomber
11
:118-122.
METHOT. R. D.
1981.
Growth
rates
and age distributions of larval and juven-
ile northern anchovy,
Engraulis
mor&,
with inferences on
larval survival. Ph.D. Thesis, Univ. California, San Diego,
209
p.
NISHIMURA, A,,
Y.
WATANABE,
AND
J.
YAMADA.
1985.
Daily growth increment-like microstructure in otoliths
of the Pacific
saury
Colohbia
saim.
(In Jpn.)
Buli.
Tohoku
Reg. Fish. Res. Lab.
47:33-35.
NISHIMURA, A,,
AND
J.
YAMADA.
1984.
Age and growth of larval and juvenile walleye pollock
Theragra
chnlcogrammu (Pallas).
as
determined by otolith
daily growth increments.
J.
Exp. Mar. Biol. Ecol.
82191-
205.
NOVIKOV, Yu.V.
1960.
Oprjedjenije vamta
po
chjeshchuje i varastnoi sostav
sairy
(Cololabia
saira
(Brevoort)) v raionje ziuzhnykh
kuril'skikh
ostrovov (Determination of age by scales and age
composition of saury
(Cololubia
saira
(Brevoort)) in region
of southern Kuril Islands). (In Russ.)
IN.
TINRO
46:233-
241.
1973.
Izuchjenije populiatsionnoi struktury
i
vozrastncgo
sostava sairy sjejero-zapadnoi chasti tikhogo okjeana (The
study
of
population structure and
age
composition of saury
of north-western
prt
of the Pacific
Ocean).
(In Russ.) Izv.
TINRO
87:149-154.
NUMACHI. K.
1971.
Kouso no takei ni yoru sanma keitougun ni kansuru
kenkyu (Study on the population of the Pacific saury based
upon analyses
of
enzymatic polymorphism). (In Jpn.)
Ocean Res. Inst., Univ. Tokyo, Tokyo,
57
p.
ODATE,
S.
1977.
On the distribution of Pacific saury in the North Pacific
Ocean. (In Jpn.) Res. Inst. North Pac. Fish. Sp.
Vol.,
p.
353-382.
Fac. Fish., Hokkaido Univ.. Hakodate, Ja-
pan.
ODATE.
K.
1986.
Automatic data processing for estimation of abundance
of zooplankton in the waters off northeastern Honshu,
Japan,
1951-1976.
(In Jpn.) Bull. Tohoku Reg. Fish. Res.
Lab.
4831-47.
PANNELLA. G.
1971.
Fish otoliths: daily growth layers and periodical pat-
terns. Sci.
173:1124-1127.
497
FISHERY BULLETIN: VOL.
86.
NO.
3
Southern California Bight: Comparison of time series.
Limnol. Oceanogr. 27:l-17.
STATISTICS
AND
INFORMATION
DEPARTMENT,
JAPAN.
1985. Fisheries
Statistics
of
Japan 1984.
Minist.
Agric.. For.
Fish., Jpn., 288 p.
SUNADA.
J.
S.
1974. Age and growth of the Pacific saury,
colohbis
sai7a.
Calif. Fish Game 6064-73.
THEILACKER.
G.
H.
1980. Changes in
body
measurements
of
larval northern
anchovy,
Engraulis
mmdaz,
and
other
fishes
due
to
handling
and preservation. Fish. Bull., U.S. 78685-692.
UCHIDA,
K.,
S.
IUI,
S.
Mim,
S.
FUJITA,
M.
UENO.
Y.
SHOJIMA,
1958. Studies on the eggs, larvae and juvenile of Japanese
YUSA,
T.
1960. Embryonic development of the saury
Coblabis
saim
T.
SENTA.
M.
TAHUKU,
AND
Y.
~sv.
fishes. (In Jpn.)
Kyushu
Univ., Fukuoka. Japan, 89 p.
(Brevoort). Bull. Tohoku
Reg.
Fish.
Res.
Lab. 17:l-14.
RADTKE, R. L.,
AND
J.
M.
DEAN.
1982. Increment formation in the
otolith8
of
embryos,
larvae,
and juveniles of the mummichog,
Fundulus
hetcroclatus.
Fish. Bull., US. 80:201-215.
RICKER.
W.
E.
1973. Linear regressions in fishery research.
J.
Fish. Res.
Board Can. 30409-434.
SABLIN.
V. V.
1979. Sanma no nenrei
ni
kansuru
mondai
ni
tsuite
(Problems
on
age
composition of saury). (In Jpn.) Rep. JawSoviet
Coop.
Res.
Meet. Saury Scomber 11:114-117.
SABLIN,
V.
V..
AND
V.
P. PAVLYCHEV.
1982. Dependence of migration and
catch
of
Pacific saury
upon thermal
conditions.
Bull. Tohoku
Reg.
Fish.
Res.
Lab.
44~109-117.
SMITH, P.
E., E.
H.
AHLSTROM,
AND
H.
D.
CASEY.
1970. The saury
as
a
latent resources of the California Cur-
rent. Calif. Coop. Oceanic Invest. Rep. 14%-130.
SMITH. P.
E.,
AND
R.
W.
EPPLEY.
1982. Primary production and the anchovy population in the
498
