Fisheries Research 225 (2020) 105475

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The Sea of Amakusa playing the role of a distributor of swordtip squid T

(Uroteuthis edulis) migrating from the East China Sea to the east and west
sides of Japan
Tadanori Yamaguchia,*, Katsumi Takayamab, Naoki Hiroseb, Michiya Matsuyamac
a
Saga Prefectural Genkai Fisheries R & D Center, Karatsu, 847-0122, Japan
b
Research Institute for Applied Mechanics, Kyushu University, Fukuoka, 816-8580, Japan
c
Faculty of Agriculture, Kyushu University, Fukuoka, 819-0395, Japan

A R T I C LE I N FO A B S T R A C T

Handled by: S Xavier Cadrin We investigated the monthly numbers of tracers moving from the southern East China Sea to the east side of
Keywords: Japan via the Sea of Amakusa, on the west side of Japan, by tracer experiments. We estimated the migratory
Kuroshio route of large Swordtip squid (Uroteuthis edulis) caught in Sagami Bay, on the east side of Japan, by the micro-
Migratory route increment counts and variation of strontium:calcium ratios of the statoliths to test our hypothesis that the Sea of
Statolith Amakusa plays the role of a distributor of this species to the east and west sides of Japan. Results of the monthly
Sr:Ca ratio numbers of the tracers were generally consistent with the season of peak catches on the east side. The older
Swordtip squid tracers reaching the east side of Japan were found to have passed through the Sea of Amakusa at larger rates.
Tracer experiment
Moreover, the estimated empirical water temperature history of the large squid caught in Sagami Bay showed
two peaks, in winter and early summer, implying that the squid experienced the Kuroshio current twice and that
during spring, they stayed in waters whose temperature was a few degrees lower than that of the Kuroshio; that
is, in the Sea of Amakusa. Frequent and extensive marine observations will be needed to fully understand the
distribution of this species around Japan.

1. Introduction
The swordtip squid (Uroteuthis edulis) is a neritic species widely
distributed in the western Pacific, from the southern Japan Sea and the
East China Sea in the north through to the Java Sea and coastal waters
of Indonesia, Malaysia, and Thailand in the tropics, and as far south as
the waters off northern Australia (Jereb and Roper, 2010; Arkhipkin
et al., 2015). U. edulis has become an important commercial species and
source of revenue for many fishers, particularly in the southern Japan
Sea and eastern Tsushima Strait (Fig. 1; Yoda and Takahashi, 2018) on
the west side of Japan. However, this species is also caught on the east
side of Japan, for example in Sagami Bay, which is open to the north-
western Pacific. Here, the peak of catches of U. edulis appear between
January and March when they tend to be small in size and immature;
large mature squid are caught from June to August (Kuboshima, 1992;
Sukramongkol et al., 2006). Ando and Kato (2008) reported that the
catches of this species by set nets around the Izu Islands, near Sagami
Bay, were largest in May, although the sizes of the squid were not
mentioned.
Yamaguchi et al. (2015) estimated the hatching site and migratory
route of U. edulis caught in the eastern Tsushima Strait by using the
variation of strontium:calcium ratios in statoliths (Fig. 2). However,
dynamics of elemental uptake into statoliths can be regulated by many
intrinsic and extrinsic factors, such as physiology, ontogenetic stage,
reproduction, diet, stress, temperature, salinity, and water chemistry.
For example, strontium incorporation is considered to have a negative
relationship with temperature, but temperature effects can be different
at different temperatures (Townsend et al., 1992), and water chemistry
and diet are also thought to be drivers of incorporation into other
calcified structures (Elsdon and Gillanders, 2005; Zumholz et al., 2005).
Then, by using statolith analyses (micro-increment count and variation
of Sr:Ca ratios) and particle-tracking experiments, Yamaguchi et al.
(2015, 2018) have shown that small individuals of U. edulis caught in
Sagami Bay and the waters off Owase, both of which are located on the
Pacific side of Japan, were revealed to have hatched in the southern
East China Sea and moved with the Kuroshio, the western boundary

Abbreviations: BW, body weight; ML, mantle length; TWC, Tsushima Warm Current
⁎
Corresponding author.
E-mail address: yamaguchi-tadanori@pref.saga.lg.jp (T. Yamaguchi).

https://doi.org/10.1016/j.fishres.2019.105475
Received 29 September 2019; Received in revised form 3 December 2019; Accepted 16 December 2019
Available online 21 January 2020
0165-7836/ © 2019 Elsevier B.V. All rights reserved.
T. Yamaguchi, et al.
Fig. 1. Map of the study area in the East China Sea and Northwestern Pacific.
current of the North Pacific subtropical gyre (Isobe and Imawaki,
2002), without contradictions. However, the hatching site and mi-
gratory route of the large mature squid caught on the Pacific side of
Japan are still unknown.
On the other hand, using both the methods above, Yamaguchi et al.
(2019) showed that there are differences in the migratory routes taken
by large mature and small immature U. edulis caught in the eastern
Tsushima Strait. Although squid of both sizes were assumed to have
hatched in the southern East China Sea, the large mature squid took
many more days to make the journey. It is believed that the large squid
move slowly from the northernmost Kuroshio region to the eastern
Tsushima Strait because of slow currents in the Sea of Amakusa (the
waters to the south of the Goto Islands on the west side of Kyushu Is-
land), whereas small immature squid move rapidly in fast currents in
the northern East China Sea. Through animations of the tracer experi-
ments (Yamaguchi et al., 2019), it was discovered that some particles
moved slowly northward to the eastern Tsushima Strait, but some
moved back down south and moved around the southern end of Kyushu
Island to the Pacific side of Japan after they had remained for quite a
few days in the Sea of Amakusa. From this observation, the hypothesis
was formulated that the large U. edulis caught in Sagami Bay could have
hatched in the southern East China Sea and then moved to the Sea of
Amakusa before leaving for the Pacific side of Japan. If so, the Sea of
Amakusa could be another distributor area for the squid (in addition to
the waters around 30 °N, 128 °E), where small squid moving from the
southern East China Sea are separated to the east or west side of Japan
by the Kuroshio and one of its branches (Fig. 2).
To test our hypothesis and understand the estimated migratory
routes of the small and large squid caught on the east and west sides of
Japan, we examined whether the monthly numbers of tracers moving
from the southern East China Sea to the Pacific side of Japan via the Sea
of Amakusa and the seasonal catches of the large squid caught on the
Pacific side of Japan would be consistent and whether the distribution
2
Fisheries Research 225 (2020) 105475
of the tracers would support the migratory route of the large squid
caught in Sagami Bay, which were estimated by the variation of Sr:Ca
ratios in the statoliths in this study. Moreover, we examined whether or
not the monthly number of the tracers would be consistent with the
seasonal catches of the small squid, whose migratory route had been
estimated by Yamaguchi et al. (2015, 2018).
2. Materials and methods
2.1. Distribution of seawater temperatures and currents and tracer
experiments
The ocean environment was inferred using a high-resolution data
assimilation output, and the distribution of the tracers was simulated
using the Japan Sea Data Assimilation Experiment ver.2 dataset, fol-
lowing Yamaguchi et al. (2018). The currents in which the squid moved
were assumed to be at a depth of 50 m (Yamaguchi et al., 2018).
On the first day of every month from January 2013 to December
2017, 10,000 Lagrangian tracers were released at 26°00′N, 122°30′E
(Fig. 3, “Χ”). The release point presumably belongs to the hatching sites
of the small and large U. edulis caught in the eastern Tsushima Strati
and the small ones caught on the Pacific side of Japan (Yamaguchi
et al., 2018, 2019), which led us to hypothesize that the release point
also belong to the hatching site of the large squid caught on the Pacific
side of Japan. The tracers’ advection and diffusion were calculated
using the method of Kato et al. (2014). The area of the sea, including
the Sea of Amakusa and the waters around the Goto Islands (Fig. 3, area
“A”: 31°16′–33°12′N, 128°25′–130°30′E), was thought to be very im-
portant for not only the large squid caught in the eastern Tsushima
Strait (Yamaguchi et al., 2019) but also the squid caught on the Pacific
side of Japan as well. Thereafter, the tracers entering area “A” were
identified and recorded daily for the first 365 days and then monthly
until they reached the Osumi Strait (30°36′–31°30′N, 131°00′E) or
T. Yamaguchi, et al. Fisheries Research 225 (2020) 105475

Fig. 2. Estimated migratory routes of Uroteuthis edulis from the southern East China Sea to the east and west sides of Japan. a and b arrows show the migratory routes
of small and large squid, respectively, estimated in Yamaguchi et al. (2019). c arrow shows that of small squid estimated in Yamaguchi et al. (2018). d arrow shows
those of small and large squid estimated in Yamaguchi et al. (2015) and this study, respectively. Broken-line circles show the waters presumably playing the role of a
distributor of squid migrating from the southern East China Sea.

Tokara Strait (27°00′–30°36′N, 131°00′E).
2.2. Sampled squid
To determine the migratory route of the large U. edulis that appear
on the Pacific side of Japan in summer, 11 mature specimens (four
females and seven males) were caught with a fixed net in Sagami Bay
(35°11′N, 139°17′E; Fig. 1) on August 7, 2014. Mean mantle length
(ML) and body weight (BW) ( ± standard deviation [SD]) of the females
and males were 208 ± 54 mm and 254 ± 123 g and 194 ± 61 mm
and 213 ± 187 g, respectively. The specimens were considered mature
based on there being spermatophores in the Needham’s sac in the males
or ovulated mature eggs in the oviducts in the females.
2.3. Micro-increment counts
The number of micro-increments in each of seven randomly selected
statoliths from the specimens was counted from the nucleus to the edge
of the lateral dome under a light microscope, as described by
Yamaguchi et al. (2015). The distances from the nucleus to the 50th,
100th, 150th, 200th, and last micro-increment in each statolith were also
measured to reveal the relationship between the micro-increments and
the measurement points of the wavelength-dispersive spectrometer that
was used to measure the concentrations of Sr and Ca (see Section 2.4).
2.4. Sr and Ca measurements
Differences in the Sr:Ca ratios from the nucleus to the lateral dome
edge in the statoliths of the specimens were assessed along a line of
continuous beam spots through electron probe microanalysis using a
wavelength-dispersive spectrometer (JXA-8900RLS; JEOL Ltd., Tokyo,
Japan), based on the methods of Yamaguchi et al. (2015). The following
measurement conditions were used: measurement interval, 5 μm; dia-
meter-focused beam, 5 μm; accelerating voltage, 10 kV; beam current,
100 nA; and counting time, 20 s. CaSiO3 and SrTiO3 were used as
standards for Ca and Sr, respectively, and 10-point running means were
calculated to smooth any small fluctuations of unknown origin. Dif-
ferences between the 10-point running means and the final

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T. Yamaguchi, et al. Fisheries Research 225 (2020) 105475

Fig. 3. Tracer release experiment showing the distribution of tracers on day 140 after the release day (September 1, 2017); image taken from Animation 1 of
Supplementary Material. Χ: Release point (26°00′N, 122°30′E). Area “A”: the area of the sea, including the Sea of Amakusa and the waters around the Goto Islands
(31°16′–33°12′N, 128°25′–130°30′E). Also shown are the locations of the Osumi Strait (30°36′–31°30′N, 131°00′E) and the Tokara Strait (27°00′–30°36′N, 131°00′E).

measurements were then presented as means ± SD. mmol), and C is a constant (Yamaguchi et al., 2018).

2.5. Water temperature history 3. Results

Yamaguchi et al. (2015) previously showed that a 2.8 °C increase in
water temperature corresponded with a 7.8 × 10−4 ± 2.1 × 10−4
(mean ± SD) mmol/mmol decrease in the Sr:Ca ratio. On this basis,
the Sr:Ca ratios in the statoliths were used to estimate the water tem-
perature history of the squid using the following equation derived from
this finding:
Y = –3.6 × 103 ( ± 9.7 × 102) X + C,
where Y is the water temperature (°C), X is the Sr:Ca ratio (mmol/
3.1. Tracer experiments
The mean number ( ± SD) of tracers remaining in area “A” on the
first day of each month decreased gradually from 1,274 ± 299 in
March to 675 ± 222 in October (Fig. 4). The number of tracers, which
reached the Osumi or Tokara Straits, aged four or five months, also
decreased from spring to winter (Fig. 5). On the other hand, the
(1)
monthly percentage of tracers reaching the Osumi or Tokara Straits via
area “A” among the total number reaching the straits indicated that

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T. Yamaguchi, et al. Fisheries Research 225 (2020) 105475

Fig. 4. Mean number ( ± SD) of tracers remaining in area “A” on the first day of
each month in the tracer experiments.
Fig. 7. Number of 6-, 7-, and 8-month-old tracers reaching the Osumi or Tokara
Straits via area “A”.
older tracers generally passed through area “A” at larger rates in all
months (Fig. 6). This implies that old tracers remained for a few more
months in and around the Sea of Amakusa before passing through the
Osumi or Tokara Straits. Moreover, relatively many old tracers (i.e. 6–8
months of age) reached the Osumi or Tokara Straits from January to
April and from August to September via area “A” (Fig. 7).

3.2. Age determination

Based on the micro-increment counts, the mean age ( ± SD) of the

squid caught in Sagami Bay was 260 ± 20 d (n = 7), which corre-
sponds with estimated hatching dates in the middle of November 2013.

3.3. Ontogenetic variation in statolith Sr:Ca ratios

The mean number ( ± SD) of 10-point running means of the Sr:Ca

Fig. 5. Number of 4- and 5-month-old tracers reaching the Osumi or Tokara
Straits by month.
ratios from the nucleus to the lateral dome edge in the statoliths of the
11 specimens were 128 ± 8 (Fig. 8a). Ontogenetic Sr:Ca ratios on the
50th, 100th, 150th, 200th, and 250th days from hatching corresponded to
the following 10-point running mean measurements from the nucleus:
31st ± 4, 58th ± 4, 83rd ± 5, 105th ± 6, and 123rd ± 5. The mean dif-
ference ( ± SD) between the first and last 10-point running mean
measurements of the Sr:Ca ratios was 3.4 × 10−4 ± 5.2 × 10−4
(mmol/mmol).

3.4. Water temperature history

Examination of the Sr:Ca ratios showed that the difference between

the mean empirical water temperature ( ± SD) on the estimated day of
hatching (in the middle of November 2013) and just before sampling (at
the beginning of August 2014) for the specimens was −1.2 °C ± 1.9 °C
(Fig. 8b). The mean water temperature presumably increased by
3.1 °C ± 2.2 °C for ca. 110 days from the estimated day of hatching to
the middle of March 2014, decreased by 2.5 °C ± 1.7 °C for ca. 70 days
to the middle of May, 60 days to the middle of July, and decreased
again by 1.0 °C ± 0.5 °C for ca. 20 days just before sampling.

4. Discussion

4.1. The Sea of Amakusa as a distributor

From spring to autumn, not only did the number of tracers re-
maining in area “A” decrease, but the numbers of 4–5-month-old tracers
reaching the Osumi or Tokara Straits also decreased (Figs. 4 and 5).
These results appear to show an inverse relationship with the extent of
Fig. 6. Monthly percentages of 4-, 5-, 6-, 7-, and 8-month-old tracers reaching
the Tsushima Warm Current (TWC) in the Japan Sea, which increased
the Osumi or Tokara Straits via area “A”.
during the same period (Fig. S1). The Taiwan–TWC System operates at

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T. Yamaguchi, et al. Fisheries Research 225 (2020) 105475

4.2. Migration of large squid caught off the Pacific coast of Japan

The water temperature at a depth of 50 m in Sagami Bay at the

beginning of August 2014 was ca. 18 °C (Fig. S2). The specimens caught
in the bay on August 7, 2014, would experience a water temperature of
ca. 20 °C in February and March (the squid were three to four months
old), when the only waters whose temperature was > 20 °C were in the
region of the Kuroshio (Fig. S3). Then, from March to May, the squid
had moved from the Kuroshio region to some waters whose tempera-
ture was 2–3 °C lower. Also, because the squid were finally transported
to Sagami Bay via the Kuroshio, the waters in which they resided must
have contained currents flowing into the Kuroshio region again. As the
tracer experiments in this study have shown, the Sea of Amakusa seems
to satisfy these requirements. In addition, the first period of the em-
pirical water temperature history of the squid (December to May) re-
sembles that of squid caught in the eastern Tsushima Strait in May and
June (Yamaguchi et al., 2015, 2019). Of course, there may be several
candidates for such waters along the Pacific side of Japan; but the
various bays and channels there are too small for the squid to stay for a
few months. Therefore, the large squid caught in Sagami Bay would
have hatched in the southern East China Sea and moved along the
Pacific coast of Japan with the Kuroshio via the Sea of Amakusa, ac-
counting for the two peaks in the empirical water temperature history
of the squid (Fig. 8b).

4.3. Migration of small squid caught off the Pacific coast of Japan

Many of the small squid (young squid) caught on the Pacific side of
Japan would pass through the Osumi or Tokara Straits directly, and not
via the Sea of Amakusa (Yamaguchi et al., 2018). This is reflected in the
tracer experiment results in this study (Fig. 6) and supported by the
Fig. 8. (a) Ten-point running means of the Sr:Ca ratios from the nucleus to the study by Yamaguchi et al. (2018) who estimated the migratory route of
lateral dome edge of the statoliths from 11 Uroteuthis edulis caught in Sagami small squid caught in the waters off Owase in the Kii Peninsula (Fig. 1)
Bay on August 7, 2014. Each value is shown relative to the first running mean
on the Pacific side of Japan on January 23, 2016 (ca. 104 mm ML and
Sr:Ca ratio. (b) Ontogenetic variations in the mean ( ± standard deviation [SD])
159 days of age). However, some small squid would come via the Sea of
water temperature to which the specimens were exposed relative to the water
temperature when they were caught. The historic water temperatures and time
Amakusa; empirical water temperature history of small squid caught in
of year were estimated based on Sr:Ca ratios and micro-increment counts in the Sagami Bay on May 23, 2014, was shown to exhibit two peaks (ca.
statoliths. 148 mm ML and 171 days of age; Yamaguchi et al., 2015). In fact, the
squid would have stayed for a relatively short time in the waters of the
Sea of Amakusa, and the duration of onward travel would have been
least from April to August (Isobe, 1999); this means that the TWC is
short, perhaps because of a difference in speed or other factors of the
probably strongest in autumn and weakest in winter. In winter then,
Kuroshio along the Pacific coast at that time, accounting for the young
when the TWC is weakest, more tracers would remain in area “A” and,
age (small ML).
rather than flow to the eastern Tsushima Strait, would flow to the Pa-
cific side of Japan through the Osumi or Tokara Straits with the Kur-
4.4. Conclusion and further study
oshio. This is probably the reason why many small U. edulis are caught
off the Pacific side of Japan in winter (Kuboshima, 1992; Sukramongkol
Some individuals of U. edulis hatching in the southern East China
et al., 2006). Moreover, the numbers of 6–8-month-old tracers reaching
Sea would enter the Kuroshio region and be moved northeastward
the Osumi or Tokara Straits in winter and summer were greater than in
along the shelf with the currents (Fig. 2). Approaching Japan, at around
the other seasons. In fact, few large squid are caught on the Pacific side
30 °N, 128 °E, some would be separated eastward to the Pacific side of
of Japan in winter because fewer U. edulis hatch in the southern East
Japan through the Tokara Strait along with the Kuroshio main stream,
China Sea between June and August (Wang et al., 2010) (the period
whereas some would be taken northwestward towards the Tsushima
during which these tracers were released there), whereas it is said
Strait with the fast currents. Then, these waters would play the role of a
(private communication from the Uchinoura Branch of Japan Fisheries
distributor of the squid migrating from the southern East China Sea to
Cooperative) that of the squid caught in the Osumi Strait from July to
the east and west sides of Japan. A little later, some of the remaining
September, the majority are ca. 100–300 g BW (ca. 140–270 mm ML)
squid would again be carried eastward to the Pacific side of Japan, this
and a minority are over 500 g BW (over 330 mm ML), and large mature
time through the Osumi Strait in the northernmost Kuroshio currents,
squid were caught by fixed nets in Sagami Bay. On the other hand,
whereas the remaining squid would gradually leave the northernmost
when the TWC increases in strength from spring onward, the many
Kuroshio region and enter the Sea of Amakusa, accumulating in the
tracers that stayed in area “A” over the winter move not only northward
complex of currents there. The Sea of Amakusa would then play an
but also southward, although the distribution factor is unknown. In this
important role as another distributor of the squid because its currents
case, the number of tracers could be accounted for not by the currents
would often move the squid northward to the eastern Tsushima Strait
but by the increase in tracers that passed over the Osumi Strait in
and sometimes southward to the Osumi Strait. The direction of the
summer, reflected in the catch of large squid on the Pacific side of
currents seems to be affected by the balance in strength between the
Japan in summer.
Kuroshio and the Taiwan–TWC system.
It should become increasingly possible to estimate the migratory

6
T. Yamaguchi, et al.
direction and speed of the large squid from the Sea of Amakusa to the
eastern Tsushima Strait and the Osumi Strait as observation data in the
Sea of Amakusa accumulates. Due to the lack of information about the
number of squid entering these waters, it would be difficult to forecast
when good catches would be likely, but it would be possible to forecast
when poor catches are likely; this would be when the currents are
arising not from the Sea of Amakusa but other waters. Such predictions
could not only help improve operating efficiency for local fishers but
could also support sustainable stock management of this species.
Potentially, similar predictions could be made for the Japanese horse
mackerel (Trachurus japonicus) because large numbers of eggs and
larvae of this species arising in the southern East China Sea would also
be transported northeastward by the Kuroshio and its branches into the
shallow waters off the west coast of Kyushu Island and the Pacific coast
of southern Japan (Sassa et al., 2006). Then, some of the fish might
migrate from the southern East China Sea to the Pacific coast via the
Sea of Amakusa as U. edulis would. On the other hand, some of the
Japanese common squid (Todarodesu pacificus) are thought to migrate
from the East China Sea along the Pacific coast of Japan by the Kuroshio
before traveling around the Japanese archipelago in the counter-
clockwise direction to return the hatching site and spawn eggs (Rosa
et al., 2011). The migratory route of the species might be almost the
same as that of the small U. edlis because of the fishing grounds and
sizes of the squid. Additionally, some of the T. pacificus are thought to
migrate from the East China Sea to the Japan Sea through the Tsushima
Strait with the currents. Therefore, frequent and extensive marine ob-
servation on the east and west sides of Japan will be needed to fully
understand the distribution of not only U. edulis but also some other
species around Japan.
Funding
This study was supported by funding from the Saga prefectural local
government and the Fisheries Agency of Japan.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgements
We express our sincere thanks to R. Shimomura for the specimens of
Uroteuthis edulis. We are grateful to the Industrial Technology Center of
Saga for their permission to use the wavelength dispersive spectro-
meter. Sincere thanks are also expressed to N. Kasaoka (Nakayama Steel
Works, Ltd.) and D. Hamawaki (the Uchinoura Branch of Japan
Fisheries Cooperative), who kindly provided us with the information
Fisheries Research 225 (2020) 105475
about catches of the squid in the Osumi Strait. We express our gratitude
to M. Maeda and C. Nakatsuka (Saga Prefectural Genkai Fisheries R & D
Center) for careful statolith preparation.
Appendix A. Supplementary data
Supplementary material related to this article can be found, in the
online version, at doi:https://doi.org/10.1016/j.fishres.2019.105475.
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