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DOI 10.1007/s12562-014-0807-x
ORIGINAL ARTICLE
Fisheries
Received: 6 February 2014 / Accepted: 2 August 2014 / Published online: 19 September 2014
The Author(s) 2014. This article is published with open access at Springerlink.com
Introduction
Sustainable stock of Japanese eel Anguilla japonica is
important to freshwater fisheries in Japan where the eel
stock is primarily exploited for two objectives: human
consumption and seeds for aquaculture. Traditional fisheries, such as pod, long-line and set net, exploit fishery
stock consisting of yellow-stage and silver-stage eels [1] of
commercial size (referred to as exploitable stock in the
following text) for human consumption. Juveniles (eels
smaller than the commercial size) can be harvested by
some fisheries either incidentally on a case-by-case basis or
under special permit for seeds. Japanese eel fisheries have a
long history, going back many hundreds of years. Smallscale fisheries, such as hand-net and bag net, are only
allowed to exploit glass eel stock for aquaculture seed
under prefectural special permit, in association with
aquaculture.
Japanese eel aquaculture began in 1879 [2], at which
time all seeds were natural juveniles caught by the traditional fisheries. Commercial utilization of glass eels for
seed first succeeded in 1923, followed by the rapid development of aquaculture systems using glass eels in the
1930s [3, 4], as well as takes of glass eels. The takes
dropped in World War II, but by 1957 had recovered to the
level before World War II [5]. Eel aquaculture utilizing
glass eels rapidly developed in the 1970s1980s in China,
Korea and Taiwan [6, 7]. Currently, glass eels for seed are
taken from the coastal zones of all countries and regions
and are traded internationally.
According to official Japanese statistics (http://www.
maff.go.jp/j/tokei/kouhyou/naisui_gyosei/index.html), the
average catch of exploitable stock [300 tons (rounded off)]
during the period 20062010 was greater than that of glass
eels (12 tons) and much smaller than that from aquaculture
123
1130
123
500
3,000
400
300
2,000
200
1,000
100
0
1960
1970
1980
1990
2000
0
2010
600
4,000
Year
600
500
Fishing effort
(1,000 day-boat)
(21,000 tons). Therefore, aquaculture production is economically relatively more important than fisheries in terms
of exploitable stock. According the statistics of the Food
and Agriculture Organization of the United Nations (FAO;
http://www.fao.org/fishery/statistics/software/fishstat/en),
other countries and regions mainly exploit glass eels for
seeds in aquaculture while Japanese fisheries share the
majority of exploitable stock catch because the consumption of eels as food in countries other than Japan is
local.
The catch of exploitable stock and glass eel in Japan has
been steadily decreasing in recent years, with the latest
estimates of the average catch of exploitable stock at
approximately 8 % of the maximum post-World War II.
Depletion of stock is a matter of grave concern, and three
reasons have been suggested to account for the reduced
catch [8]: overexploitation of stock, (commercial/industrial) development of rivers and changes in the ocean
environment. Although the reduction in stock size can be
accurately assessed, there is much uncertainty about the
level of depletion since it is estimated based on the catch
amount.
Another reason for the decreasing catch amount is
associated with the drastic decline in freshwater fisheries in
Japan [9]. Both the fishery census in 1968 [10] and successive censuses have revealed that the total amount of
fishing units in major Japanese lakes (except for gillnet,
throw net, shellfish and seaweed fisheries) have decreased
from 4,082 units in 1968 to 977 in 2008 and that the units
targeting exploitable stock of eels have decreased from 551
units in 1968 to 91 in 2008 (Fig. 1). In Lakes Kasumigaura
and Kitaura, where the eel catch in 1963 was 11 % of the
total catch in Japan, the fishing effort has decreased from
588 (1,000 day-boats) in 1963 to 44.1 in 2001 (Fig. 2; see
annual statistics in 1954 [11] and successive volumes for
sources). Thus, the decline in fisheries had a serious impact
on productivity. Therefore, conducting some stock assessment is necessary to reliably estimate the level of depletion.
Stock assessments have been carried out for the European eel Anguilla anguilla and the American eel Anguilla
rostrata [12, 13] using current statistical methods, such as
stock synthesis [14]. Age- and sex- (or length-) structured
models were used in the latter assessment as dynamical
models of stock size [15, 16], and the effect of natural
mortality after spawning was incorporated into the models
to express the more realistic dynamics of eel stock size.
Stock assessment of the Japanese eel needs data such as
catch statistics, abundance index, CPUE (catch per unit
effort) and biological and operational parameters (including age and growth, natural mortality, sexual maturity and
gear selectivity).
Studies on the biological aspects essential for a stock
assessment of Japanese eel have produced an abundance of
400
300
200
100
0
1950
1960
1970
1980
1990
2000
2010
Year
1131
Stock
component
Countries and
region
Data
Exploitable
stock
East Asia
Catch
Glass eels
Japan
Japan Aquaculture
Newspaper, unpublished, 2013
Sources
19582010
FAO statisticsa,
19031911
19121943
19501953
19542010
China
19731992
Taiwan
Aquaculture
19581980
China
19902010
Newspaperd
Korea
19722010
Newspaperd
Japan
19722010
Newspaperd
Takes
Aquaculture production
19572010
19291963
Malaysia
19911996
Newspaperd
Taiwan
19802010
Newspaperd
Catch
Period
123
1132
Table 2 Japanese data sources
for series of index of stock
abundance
Stock
component
Areas
Data
Period
Sources
Exploitable
stock
19542006
Annual statisticsa
[11]
19541969
Annual statisticsa,
[26]
Catch effort
19682008
Annual statisticsa
[11]
19682008
Censusa,
19771997
Lake Hamana
Abundance catch
1972, 19742006
Research reporta
[29]
Research report [5,
2732]
19542010
Tokushima Prefecture
Seasonal
abundance
5-year interval
123
19731974,
19761977
5,000
4,000
Catch (ton)
3,000
2,000
1,000
0
1900
1920
1940
1960
1980
2000
1980
2000
Year
250
200
Catch (ton)
[10]
Nine prefectures
150
100
50
0
1900
1920
1940
1960
Year
1133
123
1134
collected over 30 years were sufficient for effectively estimating long-term changes in abundance.
Along the coast of Tokushima Prefecture, surveys using
hand-nets were simultaneously carried out in 19731974 and
19761977 [33] at various sites and during different months.
CPUE records according to day and region (north or middle)
were reported, and data from December to April in the
subsequent year were used for estimating the parameters.
Let ci;t;m;k;j be CPUE (eel/operation) of the
j 1; 2; . . .; JMAX;z;i th operation of survey i 2; 3 in
month m 1; 2; . . .; mMAX of year t 1972; . . .; 2002;
or 1973,. . .; 1977 for site k 1; 2; . . .; KMAX;z;i . In the
abundance survey of glass eels, the frequency of zero
CPUE was so high that zero-inflated models, such as those
used in the analysis of data from tuna longline fishery [37,
38], were applied. The CPUE (fish/operation) is expressed
by the following zero-inflated negative binomial
distribution:
8
> 1 di;t
<
vi
ci;t;m;k 1
C vi ci;t;m;k:j 1
vi
mi;t;m 1
P ci;t;m;k;j
>
d
: i;t Cv c
mi;t;m vi 1
i;t;m;k;j 1 ! mi;t;m vi 1
i
1 exp b2;t 12;k
1 exp b3;t n3;m 13;k
di;t exp j0;i j1;i t ;
j0;i ; j1;i [ 0 :
6b
Here bi;t , ni;m and 1i;k are year, month and site effects,
respectively, and j0;i and j1;i are coefficients.
Parameters were estimated by the MLE method, and the
best model was statistically selected using the Akaike
information criterion from several models, such as 1i;k 0
and/or j1;i 0. The selected model was expressed by
m2;t;m 1 exp b^2;t and d2;t exp j^0;2 j^1;2 t for
Lake Hamana, and by m3;t;m 1 exp b^3;t n^3;m and
di;t 1 for Lake Tokushima.
For each survey, expectation of CPUE using only the
year effect was used for stock assessment and is expressed
by
n
o
E u0;i;t; d^i;t 1 exp b^i;t
i 2; 3:
8
123
ci;t;m;k;j 0
5a
ci;t;m;k;j 1; 2; . . . :
5b
Pz;t
8
<1R
t
2
:
Ns;a1;t1 exp Ma1;t1 Fs;a1;t1 1 ms;a1
8
w0 Rt
>
>
>
aMAX
>
<P P
Ss;a ws;a Ns;a;t
s a1
>
MAX
>
P aP
>
>
ws;a Ns;a;t
:
s
a 0
z 0
10a
z 1
10b
z 2;
10c
a1
11
12
Fs;a;t
f0;t
ln 1 Ss;a 1 expf1;t
Ma;t
9a
a 1; . . .; aMAX ; 9b
M0 dM t
Ma
a 0
1 a;
a 0
a 1; . . .; aMAX ;
14a
14b
13a
13b
1135
Table 3 continued
Variable
Description
Ys;a;t
Yz;t
Variable
Description
a0;s
yt
am;s
aS;s
bw
BMAX
Bt
ci;t;m;k
az;i;k
bi;t
vi
c0 ,c1
Ez;t
di;t
dM t, dR t,
du t
ez;i;t;k
fz;t
eSR;t
Fs;a;t
gt
gBt
h
Recruitment function of Bt
Steepness (0:2\h\1, shape parameter in Beverton
and Holt model of stock and recruitment)
j0;i , j1;i
mi;t
q
r2z;i
r2SR
ni;m
Ks
1i;k
ln
n 1; 2; . . .th likelihood
Month
ms;a
Maturity rate
Ma
Ns;a;t
pM , pR , pu
Pz;t
qz;i;t
rm;s , rS;s
RMAX
Rt
s
Ss;a
tM;1 , tM;2
tR;1 , tR;2
ts
tu;1 , tu;2
uz;i;t;k
CPUE (kg/effort)
vi
Conversion parameter
w0
ws;a
W1;s
xf
Rt gBt
8
>
>
<
4hRMAX dR t
MAX
5h 1 1hB
Bt
8
>
>
<
16a)
t tM;1 1; . . .; tM;2
(16b)
t tM;2 1; . . .; 2010;
(16c)
t 1903; . . .; tR;1
17a)
t tR;1 1; . . .; tR;2
(17b)
t tR;2 1; . . .; 2010;
(17c)
t tR;1
dR t 1 pR 1
tR;2 tR;1
>
>
:
pR
Bt
t 1903; . . .; tM;1
t tM;1
dM t 1 pM 1
t
tM;1
>
M;2
>
:
pM
15
wfemale;a mfemale;a1 Nfemale;a1;t1 exp Ma1 Ffemale;a1;t1 ;
a1
18a
BMAX
a1
X
1X
wfemale;a mfemale;a1 RMAX exp
Mi
2 a1
i1
a1
Y
1 mfemale;k1 ;
k1
18b
ms;a
8
<0
:
1 exp rm;s a am;s
a 0; 1
19a
a 2; . . .; aMAX ;
19b
123
1136
Ss;a
ws;a
8
<1
:
1 exp rS;s a aS;s
8
< w0
a 0
20a
a 1; . . .; aMAX ;
20b
oib
a 0
21a
a 1; . . .; aMAX :
21b
22a
gt eSR;t qgt1
eSR;t N 0; r2SR
22b
Growth parameters
Maturity
Selectivity
Natural mortality coefficient
ln :
23
n1
123
w0 (g)
0.15
0.15
W?, s (g)
372.4
1,365.1
Ks (1/year)
a0, s (year)
0.18
-1.07
0.13
-1.14
bw,
3.42
3.42
rm s (1/year)
1.47
1.47
am, s (year)
5.00
9.00
rS s (1/year)
2.94
2.94
aS, s (year)
2.50
2.50
M0 (1/half of year)
3.40
3.40
M1 (1/year)
0.20
0.20
M2 (1/year)
0.10
0.10
Ma (a [ 2, 1/year)
0.05
0.05
2 Xh n
o
1X
ln 2p V ln u1;i;t; r^21;i
2 i1 t
2 #
ln E u1;i;t; =du t ln q1;i;t Pt
;
V ln u1;i;t; r^21;i
l1
3 Xh
h
i
1X
ln 2pV ln u0;i;t; r^20;i
2 i1 t
2 #
ln E u0;i;t; =du t ln q^0;i vi Rt
;
V ln u0;i;t; r^20;i
q^1;1 t ts ; . . .; tc 1
q1;1;t ^0
q 1;1 t tc ; . . .; 2006;
24
l2
vi
w0
1
8
>
>
<
Log-likelihoods
Female
Biological parameters
Table 4 shows the values of the biological and operational
parameters used for estimations under the base case scenario (S1) that uses a standard data set. Parameter values,
with the exception of natural mortality coefficients and
parameters in the selectivity function, are estimates for
eels in the Tone River, Chiba Prefecture [21]. Values of
natural mortality coefficients in Table 4 are assumed from
values used to indicate values of European eels [3841],
and the values of parameters in the selectivity function are
assumed from the shape of the length frequency in the
catch [21].
Value
Male
22c
Notation
i 1
i 2; 3;
25
26a
26b)
27a
27b)
t tR;1
du t 1 pu 1
t
tR;1
>
R;2
>
:
pu
t 1903; . . .; tu;1
28a)
t tu;1 1; . . .; tu;2
(28b)
t tu;2 1; . . .; 2010:
(28c)
ln
gB
q^lnRt1 lngBt2 g2
t
t1
2
ln 2pr^SR
:
2 tts 1
r^2SR
29
1137
f0;t
8^
f0;t
>
>
>
<
t ts 1 ^
f0;ts 1 f0;ts 1
f0;ts 1
3
>
>
>
t ts 3m 1 ^
: f^
f0;ts 3m2 f^0;ts 3m1
0;ts 3m1
3
Ns;a;1903
8
1
>
>
R1
>
>
2
>
>
(
)
>
>
a1
X
<1
R
exp
M
F
1
n
s;n1;1
2
>
n0
>
>
>
>
a1
>
Q
>
>
1 ms;n1
:
a 0
30a
a 1; . . .; aMAX :
30b
n1
^
Given the data on Y1;1903 and the values of R^MAX and h,
the value of f1;1903 is uniquely determined by solving the
following simultaneous equations:
x0
R^MAX
^
^
4h 1 h
R1903
;
31a
xf1;1903
5h 1
31b
Y1;1903 R1903 xf0;1903 1 exp f1;1903 ;
(
)
a1
a1
Y
X
1X
xf
wfemale;a exp
Mn
2 a1
31c
n0
n1
1 mfemale;n1 1 Sfemale;n1 f1 expf g :
After 1904, the value of fz;t is deterministically computed by
Yz;t
fz;t ln 1
Pz;t
t 1904; . . .; ts 1 ;
32
Y0;t
:
w0 1 expf0;t
33
t ts 3m 1; m 1; 2;
34a
t ts ; ts 1
34b
elsewhere:
34c
^ f^0;t 3m1 ; q^1;1 ; q^0 1;1 ; q^0;j ;
Unknown parameters R^MAX ; h;
s
^ maximizing Eq. 23 are numerically searched
r^2z;i ; r^2SR ; q
for under the following constraints on variances:
ol
ol
2 0:
2
or^z;i or^SR
35
123
1138
Code
number
Scenario
Description
Parameter
Point
estimates
Lower 95 %
CL
Upper 95 %
CL
S1
Base case
q 0, pM pR pu 1
3,435.8
2,259.9
5,223.6
S2
Small M0
0.662
0.214
0.894
S3
Large M0
0.285
0.178
0.456
S4
0.5 Ma and
M0 =2.9
50 % of Ma a 1 and M0 2:9
0.452
0.285
0.717
0.561
0.366
0.861
S5
1.5 Ma and
M0 =3.9
0.308
0.183
0.517
S6
M0 t
0.284
0.654
0.174
0.445
0.466
0.961
0.322
0.215
0.482
0.729
0.481
1.104
0.516
0.386
0.690
0.691
0.529
0.901
0.514
0.365
0.724
0.594
0.395
0.892
0.503
0.333
0.760
0.847
0.565
1.268
0.433
0.211
0.890
0.747
0.396
1.409
0.480
0.270
0.852
S7
0.7 W1;s
S8
Sa shift
S9
ma shift
S10
AR (1)
q is estimated
S11
h 0:3
h 0:3
S12
h 0:8
h 0:8
S13
RMAX decrease
S14
CPUE
corrected
Underreported
catch A
S15
S16
S17
S18
Underreported
catch B
0.487
0.258
0.920
Underreported
catch C
Excluding one
CPUE
0.346
0.178
0.671
0.016
0.011
0.021
0.012
0.007
0.022
0.021
0.014
0.032
0.012
0.007
0.022
0.015
0.009
0.026
0.035
0.008
0.153
r1,1
r1,2
1.405
1.376
1.128
0.834
1.750
2.269
r0,1
3.991
2.669
5.970
r0,2
1.221
1.133
1.316
rSR
1.376
0.834
2.269
Results
The results of the estimation for the base case scenario are
summarized in Table 6 and Figs. 48. Table 6 shows the
values of the parameter estimate and indicates that the
value of h^ was 0.66 (95 % CL 0.210.89), that of P^2;2010
(stock size of individuals aged C1 year) was 18.7 (95 %
CL 9.228.4) thousand tons and that of P^2;2010 =K was 24 %
of the initial size (544 %).
Figures 4 and 5 compare data and MLE of CPUE in the
base case scenario. In CPUE for exploitable stock in Lake
Kasumigaura and Kitaura, there was a discrepancy in the
second half of the 1980s between the data and the estimates, suggesting the existence of locality of stock abundance in the data. Comparison of three series of CPUE for
glass eels revealed that the goodness of fit of estimates to
data of commercial fisheries were better than those of the
123
1139
3.0
Ln CPUE(kg/fisher)
Ln CPUE (ton/day-boat)
-6.0
-7.0
-8.0
-9.0
-10.0
-11.0
1950
2.0
1.0
0.0
-1.0
1950
1960
1970
1980
1990
2000
1960
2010
1970
1980
1990
2000
2010
Year
Year
1.0
Ln CPUE(fish/operation)
0.0
-1.0
-2.0
-3.0
1950
1960
1970
1980
1990
2000
8.0
6.0
4.0
2.0
0.0
-2.0
-4.0
1970
1990
2000
2010
Year
2010
Year
1980
Discussion
Under various scenarios, the results of the estimation firmly
supported the recent recovery of P^2;t . It was hypothesized
that the recovery would be caused by the reduction of scale
of eel fisheries, as discussed earlier. Since the main fishing
gears for catching eel stock are traditional and conservative, a decrease in the number of fishing units would
therefore contribute significantly to the reduction of excess
effort.
To understand the unified effect of two fishing mortalities (f0;t and f1;t ) to the exploitable stock, the value of f0;t is
converted to that of f1;t , giving the same spawning per
recruit (xf1;t ). In the biological and operational parameters in the base case scenario (S1), the unified value is
approximately expressed by f1;t 0:123f0;t . The averages
123
80
70
1140
4
3
2
1
60
50
40
30
20
10
0
0
10
15
20
0
1950
0.6
0.5
0.4
1970
1980
1990
2000
2010
Abundance
(billion fish)
0.7
1960
Year
4
3
2
1
0.3
0
1950
0.2
1960
1970
1980
1990
2000
2010
Year
0.1
0.0
1950
1960
1970
1980
1990
2000
2010
Year
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
1950
1960
1970
1980
1990
2000
2010
Year
123
In the sensitivity tests, S15S17 cover the entire plausible range and, consequently, the underreported catch does
not affect the present results. It is a possibility that Japanese statistics do not include part of the catch in the coastal
zone [44]; however, the effect of the catch is small because
the scale of fisheries exploiting the stock in the sea is relatively tiny compared to that in freshwater fisheries.
Using ad hoc data to calculate the rate of the annual
aquaculture production to the annual input of utilization of
glass eels as seeds (for example, 690 times for Chinese
aquaculture), the recent Chinese input of glass eels was
estimated to be sevenfold the total amount adopted in our
study. The estimate of total takes of glass eels in East Asia
has increased since the 1970s; however, the increase has
been quite inconsistent with respect to the decreased input
of glass eels in each of the countries and regions. Thus,
data on eel stock should be carefully examined.
A potential for overreported takes of glass eels in Japanese statistics was presented when the annual demand of
glass eels was estimated by converting aquaculture production of commercial size eels to constant survival and
growth rates [45]. However, in general, demand does not
1141
l1
l2
l3
pa
fb0,*
fb1,*
P1980/Kt
P1980 (ton)
P2010 (ton)
S1
-145.8
-38.3
-77.3
-30.2
32
3,436
0.662
0.430
0.022
0.241
16,453
18,783
S2
-145.4
-48.2
-76.0
-21.3
32
2,931
0.547
0.509
0.013
0.268
26,028
29,257
S3
-152.7
-43.9
-77.2
-31.6
32
8,782
0.383
0.247
0.015
0.221
24,117
26,175
S4
-145.2
-44.7
-75.6
-24.9
32
2,826
0.506
0.660
0.013
0.213
33,618
28,496
S5
-153.6
-42.3
-77.5
-33.9
32
9,538
0.424
0.213
0.016
0.233
20,916
24,287
S6
-136.6
-36.8
-75.3
-24.5
32
4,771
0.902
0.197
0.042
0.087
23,580
9,417
S7
-152.8
-39.4
-77.6
-35.8
32
4,344
0.640
0.348
0.024
0.247
16,075
17,093
S8
-145.0
-42.1
-77.7
-25.2
32
3,092
0.665
0.466
0.025
0.249
15,963
17,454
S9
-141.8
-40.1
-77.0
-24.7
32
3,147
0.650
0.483
0.020
0.231
19,553
20,886
S10
-140.2
-38.3
-77.3
-24.7
33
3,398
0.665
0.432
0.022
0.242
16,321
18,685
S11
S12
-154.9
-145.8
-40.9
-37.0
-77.5
-78.2
-36.5
-30.7
31
31
8,456
2,874
0.300
0.800
0.375
0.433
0.012
0.023
0.155
0.281
35,635
14,101
29,654
18,311
S13
-135.7
-37.0
-75.8
-23.0
32
4,313
0.770
0.453
0.021
0.661
20,724
19,394
S14
-152.8
-45.9
-75.7
-31.3
32
4,484
0.497
0.426
0.018
0.222
21,644
22,561
S15
-141.5
-39.9
-76.8
-24.9
32
4,274
0.654
0.443
0.019
0.217
19,507
21,030
S16
-151.0
-39.8
-77.9
-33.3
32
4,579
0.657
0.311
0.022
0.272
22,931
28,215
S17
-145.6
-37.8
-77.0
-30.8
32
5,708
0.641
0.321
0.020
0.241
26,674
31,155
S18
-100.0
-3.9
-76.2
-19.8
32
4,599
0.448
0.347
0.013
0.289
19,385
30,191
123
1142
300
S1
4.0
Sustainable yield
(thousand ton)
S2
250
S3
S4
200
S5
150
S6
100
50
0
1950
1960
1970
1980
1990
2000
3.0
2.0
1.0
0.0
2010
Year
40
60
80
100
0.3
S1
S7
100
S8
S9
80
Sustainable yield
(thousand ton)
20
S10
S11
60
S12
S13
S14
40
0.2
0.1
20
0
1950
1960
1970
1980
1990
2000
2010
Year
0.0
0
20
40
60
80
100
120
100
S1
S15
80
S16
S17
S18
60
40
20
0
1950
1960
1970
1980
1990
2000
2010
Year
123
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