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Environ Biol Fish (2014) 97:921928

DOI 10.1007/s10641-013-0193-8

Do Japanese eels recruit into the Japan Sea coast?: A case

study in the Hayase River system, Fukui Japan
Kenzo Kaifu & Hideaki Maeda & Kazuki Yokouchi & Ryusuke Sudo &
Michael J. Miller & Jun Aoyama & Takehito Yoshida &
Katsumi Tsukamoto & Izumi Washitani

Received: 28 February 2013 / Accepted: 10 October 2013 / Published online: 18 October 2013
# Springer Science+Business Media Dordrecht 2013

Abstract The Japanese coastlines along the Sea of the present-day natural geographic distribution range of
Japan (Japan Sea) have been thought to be one of the this species, we searched for naturally recruited wild
margins of the distribution range of the Japanese eel eels in the Hayase River system, Fukui Prefecture,
Anguilla japonica, and there is evidence that eels had which flows into the Japan Sea. Multiple approaches
naturally recruited into these areas several hundred including investigation of glass eel recruitment, compar-
years ago. However, recruitment there is uncertain re- ison of body size, and estimation of habitat use types
cently, because there seems to be no study that reported was employed. During the observation period (from
glass eel or elver recruitment into the coasts along the January to July 2010), no glass eels were found at the
Japan Sea for a couple of decades, and the eels river mouth of the Hayase River in monthly sampling.
inhabiting these areas were probably stocked by fisher- Of eels collected in this study (n=127), no eels smaller
ies cooperatives. In order to improve understanding of than the initial body size of eels for stocking were found
in this water system and none were identified as being of
wild origin based on the habitat use type classifications
K. Kaifu (*) : I. Washitani from otolith microchemistry (n=48). This lack of evi-
Graduate School of Agricultural and Life Sciences, dence of Japanese eels recruiting into Japan Sea coast
The University of Tokyo, 1-1-1 Yayoi, Bunkyo-ku, waters suggests most eels present there may be stocked
Tokyo 113-8657, Japan
eels. Japanese eels could have been distributed naturally
along the Japan Sea coast more than in recent years,
H. Maeda indicating a possible decrease of the natural distribution
Fukui Marine Park Center, Fukui Prefecture, 18-2 Sekumi, range of this species.
Mikatakaminaka-gun, Fukui 919-1464, Japan

K. Yokouchi Keywords Distribution range . Fish stocking .

Institute for East China Sea Research, Nagasaki University, Japanese eel . Japan Sea . Sr:Ca
1551-7, Taira-machi, Nagasaki 851-2213, Japan

R. Sudo : M. J. Miller : J. Aoyama : K. Tsukamoto

Atmosphere and Ocean Research Institute, The University
of Tokyo, 5-1-5, Kashiwanoha, Kashiwa, Chiba 277-8563, Introduction
The Japanese eel, Anguilla japonica, is a catadromous
T. Yoshida
Department of General Systems Studies, Graduate School of
fish species that spawns in waters west of the Mariana
Arts and Sciences, University of Tokyo, 3-8-1 Komaba, Islands, and their leaf-like leptocephalus larvae drift to
Meguro-ku, Tokyo 158-8902, Japan East Asia from their offshore spawning area (Tsukamoto
922 Environ Biol Fish (2014) 97:921928

1992; Tsukamoto et al. 2011). After metamorphosis into stocking had not been conducted. It is questionable,
glass eels at the end of their larval migration, they enter however, how many Japanese eels are now actually
freshwater and estuarine growth habitats in areas adja- recruiting into this area at the margin of the species
cent to the Kuroshio Current in East Asia. Most of their range, because the Japanese eel population has de-
juvenile stage is spent as yellow eels in these continental creased dramatically. Although, glass eels have been
habitats, where they stay until the onset of sexual mat- studied in a number of locations in East Asia, to our
uration. At the beginning of their sexual maturation, eels knowledge, there seems to be no study that reported
metamorphose into silver eels and start their long mi- glass eel or elver recruitment into the coasts along the
grations to the spawning area where they were born Japan Sea in a couple of decades (Fig. 1). Fisheries
(Matsui 1972; Aoyama and Miller 2003; Tesch 2003; catch data in Japan confirm that many Japanese eels
Tsukamoto 2009). are living in rivers and lakes along the Japan Sea
The Japanese eel is an important commercial fish coast (Anonymous 19582008), but it is possible
species and recruiting glass eels are intensively cap- that these are all stocked eels caught and transported
tured for use in aquaculture. Fisheries catches of from other places.
recruiting glass eels and growth-phase juvenile The aim of this study is to examine whether the
Japanese eels, however, have continued to decrease Japanese eel is recruiting into the Hayase River system,
since the 1970s and appear to have reached a histor- which flows into the Japan Sea (Fig. 1), in order to help
ically critical situation in recent years possibly be- understand their natural recruitment into the Japan Sea
cause of changes of oceanic conditions, overfishing coast. This river system has a series of lakes and is
and loss and degradation of growth habitats relatively close to the mouth of the Japan Sea, so it may
(Tsukamoto et al. 2009). Finally, this commercially represent a good case study for detecting recruitment to
important species was listed as endangered (EN) in Japan Sea coastal areas.
the Red List revised in February 2013 by the Ministry Because the Japanese eel is a panmictic fish species
of the Environment of Japan. The serious declines of (Aoyama 2009; Han et al. 2010), stocked and wild
the Japanese eel stock require accurate ecological individuals cannot be discriminated genetically. We
information to manage this species for conservation therefore employed multiple approaches including
and sustainable use (EASEC 2012). sampling for recruiting glass eels, comparison of body
In order to develop an effective management plan size of yellow eels, and classification of habitat use
for a wild species, knowledge about the natural geo- types of yellow eels. If Japanese eels are naturally
graphic distribution range of the target species is nec- recruiting into the coast along the Japan Sea, the fol-
essary. Fish stocking however, sometimes disturbs the lowing things should be observed: (1) glass eel recruit-
natural distribution by transferring fish outside the ment, (2) eels smaller than those used for stocking, and
original distribution range. The Japanese eel is one of (3) eels that have not entered freshwater in their early
the fish species that are intensively stocked by fisheries growth period. For the third possibility, eels that have
cooperatives to enhance fisheries catches in various spent their early growth period in saline water should
areas. The primary method of this is to release small be wild, because stocked eels have been growing in
yellow eels from eel farms into rivers and lakes in freshwater ponds just after being captured in coastal
Japan. Evidence of this practice has been clearly shown waters as glass eels.
by detection of European eels, A. anguilla, migrating
downstream in the Uono River, Niigata, on the Japan
Sea coast (Miyai et al. 2004). Materials and methods
The Japan Sea coast of the Honsyu Island of Japan
has been thought to be the north-eastern margin of the Study area
natural distribution range of A. japonica (Tsukamoto
et al. 2009). In fact, eels had naturally recruited into This study was conducted during 20082012 in the
this water system several hundred years ago, because Hayase River system and its tributary rivers (Fig. 1).
Kojima et al. (2012) reported 18 literature sources The Hayase River system is a part of the Mikata-Goko
written in the Edo Period (16031868) that mentioned Lake system, and consists of Mikata, Suigetsu, Suga,
about eel fisheries along the Japan Sea coast, when eel and Kugushi lakes and their tributary rivers.
Environ Biol Fish (2014) 97:921928 923

Fig. 1 Maps of the study area (star) and places where glass 2012). Gray arrow indicates the Kuroshio Current, which
eels and elvers were collected (open circles) in previous stud- transports the eel larvae from the spawning area to their
ies (Tsukamoto 1990; Tzeng 1990; Tzeng and Tsai 1994; growth habitats. The channel between Suigetsu Lake and
Kawakami et al. 1999; Nishi and Kawamura 2005; Kim et al. Hiruga Lake has been closed, so brackish water enters Lake
2006; Fukuda et al. 2009; Han et al. 2010; Aoyama et al. Suigetsu and Mikata through Lake Kugushi

Observations for glass eel recruitment Yellow eel sampling

From January to July 2010, observations to detect glass Yellow eels were sampled in Lake Mikata and its
eel recruitment were conducted at the river mouth of tributary rivers. This lake is located at the upper-
the Hayase River, which is a single channel connecting most area of the Mikata-Goko Lake system and has
this water system and the Japan Sea. The standardized an area of 3.4 km 2 with a maximum depth of
procedure of Aoyama et al. (2012) was employed for approximately 2.5 m. The salinity of this lake var-
glass eel recruitment observations in this study. The ied from about 0 to 6, depending on the depth, area,
monthly observations consisted of two or more persons season and tide. Eels were captured in the lake by
using an underwater light and scoop net to detect and commercial fishermen using long-lines and refuge
catch recruiting glass eels during a 2-h period. Each of traps from November 2008 to August 2011. The
these standard observation periods was during night long-lines used shrimp captured in the lake as bait.
flood tides, because glass eels usually migrate up- The refuge traps were comprised of two or three
stream at night using selective tidal stream transport cylinders that were open at both ends. The cylin-
(STST) (McCleave and Robert 1982). Using the same ders were made of black PVC pipes of about 1 m
procedure, glass eels were easily caught in the Sagami length and 40 mm caliber. Sampling was conducted
River (Aoyama et al. 2012), Asahi River (Kaifu 2011) in the brackish water area near Lake Suigetsu in
and 6 other rivers located along the eastern or southern order to find eels smaller than the initial body size
coasts of Japan (N. Mochioka pers. comm.). of the eels for stocking, because Japanese eels
924 Environ Biol Fish (2014) 97:921928

spend their early continental life around the upper- Results

most brackish water areas after recruitment (Kaifu
et al. 2010). In the rivers, eels were sampled using Glass eel recruitment
an LR24 electroshocker (Smith-Root Inc., Washington,
USA) during April 2010 and June 2011. A total distance There were seven standard observation periods (14 h in
of 4,940 m of five rivers was electro-fished by two or total) that were made at the mouth of the Hayase River
more persons. during the study in 2010 according to the methods of
In order to evaluate the initial body size of stocked Aoyama et al. (2012). However, no glass eels were
eels, a sample of the eels bought by a fisheries coop- seen or captured.
erative of Lake Mikata for the purpose of stocking was
obtained in 2010 and 2011 immediately before stock- Size comparison
ing. All eels were measured to the nearest 1 mm total
length (TL). A total of 125 yellow eels (7 in 2008 by refuge trap, 43
in 2010 by long-line, and 75 in 2011 by long-line and
refuge trap) were captured in Lake Mikata. A total of
Classification of habitat use types 239 cultured eels for stocking (123 fish in 2010 and
116 in 2011) were obtained from fisheries coopera-
The Sr:Ca ratios of the otoliths of 82 eels (34 eels for tives. The total length (mean SD) was 483.6
stocking and 48 eels captured in Lake Mikata) were 110.6 mm for eels captured in Lake Mikata and
analyzed to estimate the habitat use type of each eel 291.063.8 mm for cultured eels used for stocking
using a wavelength-dispersive X-ray electron micro- (Fig. 2). The total length of the smallest fish was 268
probe (JEOL JXA-8900R) (See Kaifu et al. 2010 for and 144 mm for eels captured in Lake Mikata and
details). The electron beam was focused on a point cultured eels used for stocking, respectively.
10 m in diameter, with measurements spaced at Only two eels (one in 2010 and 2011 each) were
10 m intervals. electro-fished in tributary rivers flowing into Lake
The habitat use types of eels were categorized Mikata despite substantial sampling effort. The total
according to the mean Sr:Ca ratio of their otoliths length of these eels was 652 and 739 mm. Both of the
using the criteria of Kaifu et al. (2010), which had smallest eels captured in Lake Mikata and its tributary
been modified from Tsukamoto and Arai (2001): rivers were larger than the smallest cultured eels used
Sr:Ca ratios of 2.5 103 indicating residing in for stocking (144 mm TL).
brackish water and Sr:Ca <2.5103 indicating re-
siding in freshwater. In this study, it is important to Habitat use types
determine in which habitat eels had spent their
early continental life, freshwater or brackish water, For the eels for stocking, 34 eels were classified into
because eels that have not entered freshwater in the habitat use types using the patterns of Sr:Ca ratios
their early growth period should be naturally re- in their otoliths. For the eels captured in Lake Mikata,
cruited wild ones. Consequently, eels were catego- 48 eels (six fish from 2008 and 42 from 2010) were
rized into two groups: freshwater settlement group classified into the habitat use types. For all these eels of
and brackish water settlement group. Because the both groups, the mean Sr:Ca ratio 150440 m from
elver mark at about 150 m from the otolith core the otolith core was <2.5103, indicating that they all
was thought to correspond to the leptocephalus and had spent their early continental life in freshwater
early glass eel stages (Otake et al. 1994; Arai et al. habitats or culture ponds (Fig. 3). Although the Sr:Ca
1997), the mean Sr/Ca ratios at 150440 m were ratios of the eels for stocking were consistently low
used to estimate the environment of the habitat from the elver mark to the edge, those of the eels
where the eels had spent their early continental life. captured in Lake Mikata increased to the brackish
The otolith radius of 440 m should include a large water level (>2.5103). These results indicate that
enough number of measurement points (20) to in- the eels for stocking had stayed in the freshwater of
tegrate possible variation of Sr/Ca values caused by eel culture ponds until they were sampled, but the eels
measurement errors. captured in Lake Mikata had shifted or had been
Environ Biol Fish (2014) 97:921928 925

Fig. 2 Frequency histogram of total length classes of Japanese eels Anguilla japonica collected in Lake Mikata and its tributary rivers. The black
and gray bars indicate eels captured in Lake Mikata and its tributary rivers and cultured eels that were to be released for stocking, respectively

transferred from freshwater to the brackish environ- should be conducted in a future study. In the present
ment where they were captured. study though, we also investigated yellow eels in this
water system to examine whether they have recruited
from Japan Sea. Examining the yellow eels in this
Discussion system can provide information about their recruitment
patterns over a longer time scale.
Origin of eels in the Hayase River system If there were some eels smaller than the cultured
eels bought by fisheries cooperatives for the purpose of
By employing the exact same standardized procedure stocking, they could have come from the Japan Sea as
in this study, Aoyama et al. (2012) successfully cap- glass eels. However, we could not find any eels in Lake
tured recruiting glass eels at the Sagami River, which is Mikata or its tributary rivers that were smaller than the
facing to the Pacific Ocean. They captured glass eels smallest eel for stocking (144 mm TL). Yokouchi et al.
during December 2009 to July 2010, with a peak of 62 (2008) used the same technique of electro-fishing and
individuals caught in June 2010. Because the glass eel captured 65 eels (33 % of the total catch) smaller than
observations were conducted simultaneously in the 268 mm, and 15 eels (8 % of the total catch) smaller
Hayase River system, the method employed in this than 144 mm (the minimum size of the cultured eels for
study is appropriate to observe glass eel recruitment, stocking) in the Nishijinden River (the river mouth is at
not only in terms of observation/fishing method, but 3446N and 13732E). Therefore, at least one of the
also sampling season. However, no recruiting glass fishing gears we employed (electro-fishing) can catch
eels were observed in the present study, suggesting a eels smaller than the eels captured in this water system,
possibility that few or no glass eels recruited into this which would include eels much smaller than the min-
water system in 2010. Because our sampling effort for imum individual for stocking. The difference between
glass eel recruitment was limited in scope and within a the body sizes of the smallest eels captured in this study
single year, the possibility of some glass eel recruit- and in the previous one can be explained by a possible
ment into the Hayase River system cannot be excluded difference in glass eel recruitment. The Nishijinden
by our data, especially before or after 2010. In order to River is connected to the Pacific Ocean by Hamana
evaluate this further, longer time scale observations Lake, and Fukuda et al. (2009) have confirmed glass
926 Environ Biol Fish (2014) 97:921928

Fig. 3 Line transects of Sr:Ca ratios in the otoliths of Japanese 150 and 440 m, between which Sr/Ca ratio was averaged to
eels Anguilla japonica bought by a fisheries cooperative of Lake estimate the environment they used during their early continental
Mikata for stocking (a) and eels sampled in Lake Mikata (b). The life. The lateral dotted line indicates Sr:Ca *1,000=2.5, below
vertical dotted lines indicate distances from the otolith core of which indicates use of freshwater habitats

eel recruitment into the Hamana Lake-Nishijinden small, the sampling effort was adequate to determine
River system. Because the total length of recruiting that only a few large size eels were present in these
glass eels is about 60 mm (Fukuda et al. 2009), various rivers. This extremely low eel density is probably
sizes of eels larger than 60 mm can be captured in related to the pattern of eel stocking in this water
waters where glass eels are recruiting, but no eels system, because the fisheries cooperative of Lake
smaller than stocked ones will be found in waters Mikata and its tributary rivers has stocked eels into
without glass eel recruitment. Lake Mikata only, but not the nearby rivers (Torihama
In the tributary rivers of Lake Mikata, only two eels Fisheries Cooperative, pers. comm.).
were caught even though electro-fishing was conduct- The habitat use histories of the eels captured in the
ed intensively along these rivers for about 5 km in total. brackish water area of Lake Mikata was totally differ-
For the two largest rivers, Hasu River and Takase ent from other water systems and also provided no
River, approximately 33.5 % (3.7 km) of the total evidence of natural recruitment. Kaifu et al. (2012)
amount of river reaches was sampled by electro- showed that 79.8 % of eels captured in brackish water
fishing from the river mouths to the upper most areas. of Kojima Bay-Asahi River system had settled in
Even though the sample size of eels collected was brackish water habitat. In Hamana Lake, 35.5 % of
Environ Biol Fish (2014) 97:921928 927

silver eels had settled in brackish or sea water habitats of Japanese eel has decreased at the margins of its
(Yokouchi et al. 2012) and in Mikawa Bay (the open- species range in a similar way as for the American
ing is approximately at 3440N and 13702E), 83.3 % eel, Anguilla rostrata, which has experienced drastic
of silver eels were thought to have spent their growth decreases in recruitment to the St. Laurence River
phase in estuarine or marine areas (Kotake et al. 2005). system at the northern edge of its species range
These eels found to have settled in saline water in (Casselman 2003). In the last few decades the fisheries
previous studies must be wild eels, because eel culture catch of Japanese eels has been dramatically decreased,
ponds are usually filled with freshwater, as could be also suggesting a population decline (Tsukamoto et al.
seen in the otolith analyses of cultured eels in this 2009). It is likely that a decrease of their population
study. Therefore the results obtained from the otolith size might cause a decline in the number of larvae
analyses that found no evidence of natural recruitment recruiting to the northeastern edge of their distribution
are consistent with the lack of observations of any glass range, making it difficult to find naturally recruited
eel recruitment and the size comparison data. If some wild eels along the Japan Sea coast. Extensive research
recruitment was occurring in 2010 or the previous is required to investigate the current status and tempo-
several years, at least one of these methods should have ral trends of the species distribution range of the
detected at least a few naturally recruited eels, so there Japanese eel for the conservation and sustainable use
is no evidence to suggest that natural recruitment of of this species.
glass eels into the coastal areas of the Japan Sea is
presently occurring in this area of Japan.
Acknowledgments We thank the Torihama, Umiyama, Minami-
Saigo, and Mihama Fisheries Cooperatives for their cooperation
Possible decrease of the geographic distribution range with this study, and K Kodama and T Maezumi for helping our eel
sampling. This work was partly supported by the Environment
Research and Technology Development Fund (D-0910) of the
Japanese eels have been caught by commercial fisher-
Ministry of the Environment, Japan.
men in the waters connecting with Japan Sea
(Anonymous 19582008), however, as this study indi-
cates, recruitment into these water is questionable.
Because Japanese eel larvae are passively transported
by the North Equatorial Current and Kuroshio Current
to East Asia and they recruit into continental waters
randomly (Han et al. 2010), it is not likely that Anonymous (19582008) Statistical yearbook of ministry of
agriculture, forestry and fisheries, ministry of agriculture,
Japanese eels would recruit into waters along the forestry and fisheries, Tokyo
Japan Sea coast but not into the Hayase River system. Aoyama J (2009) Life history and evolution of migration in
For the Uono River that is located further north along catadromous eels (Genus Anguilla). Aqua Biosci Monogr
the Japan Sea coast, Miyai et al. (2004) reported that 2:142
Aoyama J, Miller MJ (2003) The silver eel. In: Aida K,
93.6 % of downstream migrating eels captured during Tsukamoto K, Yamauchi K (eds) Eel biology. Springer,
19961998 were non-native European eels, A. anguil- Tokyo, pp 107117
la. This indicates that at least 93.6 % of the silver eels Aoyama J, Shinoda A, Yoshinaga T, Tsukamoto K (2012) Late
in that study were eels that had been stocked into the arrival of Anguilla japonica glass eels at the Sagami River
estuary in two recent consecutive year classes: ecology and
river system. This suggests that few Japanese eels were socio-economic impacts. Fish Sci 78:11951204
recruiting into the waters connecting with the Japan Arai T, Otake T, Tsukamoto K (1997) Drastic changes in otolith
Sea about 10 years ago as well as in recent years. microstructure and microchemistry accompanying the onset
The Japan Sea coast has been thought to be the of metamorphosis in the Japanese eel Anguilla japonica.
Mar Ecol Prog Ser 161:1722
northeastern edge of its species distribution range Casselman JM (2003) Dynamics of resources of the American
(Tsukamoto et al. 2009). However, they appear to have eel, Anguilla rostrata: declining abundance in the 1990s. In:
been distributed naturally along the Japan Sea coast Aida K, Tsukamoto K, Yamauchi K (eds) Eel biology.
several hundred years ago (Kojima et al. 2012). This Springer, Tokyo, pp 255274
EASEC (2012) Statement of the East Asia Eel Resource
study, though, found that, they may not recruit very Consortium for the protection and conservation of the
much into these waters in recent years. This indicates Japanese eel. Emergency EASEC Symposium, 19 March
the possibility that the natural geographical distribution 2012.
928 Environ Biol Fish (2014) 97:921928

EASECdeclarations%28Final%29.pdf. Accessed 15 introduced European eels in the Uono River, Japan. Environ
November 2012 Biol Fish 71:105114
Fukuda N, Kuroki M, Shinoda A, Yamada Y, Okamura A, Nishi T, Kawamura G (2005) Anguilla japonica is already
Aoyama J, Tsukamoto K (2009) Influence of water temper- magnetosensitive at the glass eel phase. J Fish Biol 67:
ature and feeding regime on otolith growth in Anguilla 12131224
japonica glass eels and elvers: does otolith growth cease Otake T, Ishii T, Nakahara M, Nakamura R (1994) Drastic
at low temperatures? J Fish Biol 74:19151933 changes in otolith strontium/calcium ratios in leptocephali
Han YS, Hung CL, Tzeng WN (2010) Population genetic struc- and glass eels of Japanese eel Anguilla japonica. Mar Ecol
ture of the Japanese eel Anguilla japonica: panmixia at Prog Ser 112:189193
spatial and temporal scales. Mar Ecol Prog Ser 401:221 Tesch FW (2003) The eel biology and management of anguillid
232 eels. Blackwell Publishing, London
Kaifu K (2011) Natural resource ecology of Japanese eels in Tsukamoto K (1990) Recruitment mechanism of the eel,
Kojima Bay-Asahi River system, Okayama. Dissertation, Anguilla japonica, to the Japanese coast. J Fish Biol 36:
The University of Tokyo (in Japanese) 659671
Kaifu K, Tamura M, Aoyama J, Tsukamoto K (2010) Dispersal Tsukamoto K (1992) Discovery of the spawning area for
of yellow phase Japanese eels Anguilla japonica after re- Japanese eel. Nature 356:789791
cruitment in the Kojima Bay-Asahi River system, Japan. Tsukamoto K (2009) Oceanic migration and spawning of
Environ Biol Fish 88:273282 anguillid eels. J Fish Biol 74:18331852
Kaifu K, Miller MJ, Yada T, Aoyama J, Washitani I, Tsukamoto Tsukamoto K, Arai T (2001) Facultative catadromy of the eel
K (2012) Growth differences of Japanese eels Anguilla Anguilla japonica between freshwater and seawater habi-
japonica between fresh and brackish water habitats in rela- tats. Mar Ecol Prog Ser 220:265276
tion to annual food consumption in the Kojima Bay-Asahi Tsukamoto K, Aoyama J, Miller MJ (2009) Status of the
River system, Japan. Ecol Freshw Fish 22:127136 Japanese eel: resources and recent research. In: Casselman
Kawakami Y, Mochioka N, Kimura R, Nakazono A (1999) JM, Cairns DK (eds) Eels at the edge: science, status, and
Seasonal changes of the RNA/DNA ratio, size and lipid conservation concerns. American Fisheries Society
contents and immigration adaptability of Japanese glass- Symposium 58, Bethesda, Maryland, pp 2135
eels, Anguilla japonica, collected in northern Kyushu, Tsukamoto K, Chow S, Otake T, Kurogi H, Mochioka N, Miller
Japan. J Exp Mar Biol Ecol 238:119 MJ, Aoyama J, Kimura S, Watanabe S, Yoshinaga T,
Kim WS, Yoon SJ, Kim JW, Lee JA, Lee TW (2006) Metabolic Shinoda A, Kuroki M, Oya M, Watanabe T, Hata K, Ijiri
response under different salinity and temperature conditions S, Kazeto Y, Nomura K, Tanaka H (2011) Oceanic
for glass eel Anguilla japonica. Mar Biol 149:12091215 spawning ecology of freshwater eels in the western North
Kojima H, Kaifu K, Yokouchi K, Sudo R, Yoshida T, Tsukamoto Pacific. Nat Commun 2:19
K, Washitani I (2012) Historical changes of the Japanese eel Tzeng WN (1990) Relationship between growth rate and age at
distribution in the Mikata-Goko Lakes-Hayase River sys- recruitment of Anguilla japonica elvers in a Taiwan estuary
tem, Fukui prefecture. Zoo Archaeol 29:117 (in Japanese as inferred from otolith growth increments. Mar Biol 107:
with English abstract) 7581
Kotake A, Okamura A, Yamada Y, Utoh T, Arai T, Miller MJ, Tzeng WN, Tsai YC (1994) Changes in otolith microchemistry
Oka H, Tsukamoto K (2005) Seasonal variation in the of the Japanese eel, Anguilla japonica, during its migration
migratory history of the Japanese eel Anguilla japonica in from the ocean to the rivers Taiwan. J Fish Biol 45:671683
Mikawa Bay, Japan. Mar Ecol Prog Ser 293:213225 Yokouchi K, Aoyama J, Oka HP, Tsukamoto K (2008) Variation
Matsui I (1972) Eel biology (in Japanese) Kouseisha in the demographic characteristics of yellow-phase
Kouseikaku, Tokyo Japanese eels in different habitats of the Hamana Lake
McCleave JD, Robert CK (1982) Selective tidal stream transport system, Japan. Ecol Freshw Fish 17:639652
in the estuarine migration of glass eels of the American eel Yokouchi K, Fukuda N, Miller MJ, Aoyama J, Daverat F,
(Anguilla rostrata). J Cons int Explor Mer 40:262271 Tsukamoto K (2012) Influences of early habitat use on the
Miyai T, Aoyama J, Sasai S, Inoue JG, Miller MJ, Tsukamoto K migratory plasticity and demography of Japanese eels in
(2004) Ecological aspects of the downstream migration of central Japan. Estuar Coast Shelf Sci 107:132140