41

Ecological Impacts of Rainbow, Brown and Brook Trout
in Japanese Inland Waters
Satoshi KITANO
Nagano Environmental Conservation Research Institute2054-120 Kitago, Nagano 381-0075, Japane-mail: kitano-satoshi@pref.nagano.jp

Abstract
Rainbow trout (Oncorhynchus mykiss), brown trout (Salmo trutta) and brook trout (Salvelinus
fontinalis) introduced to Japanese inland waters have significant impacts on native aquatic organisms
such as white-spotted charr (Salvelinus leucomaenis), Dolly Varden charr (Salvelinus malma), masu
salmon (Oncorhynchus masou) and Sakhalin taimen (Hucho perri), in coldwater habitats. Rainbow
and brown trout have successfully established reproducing populations in Japan, and especiallyexpanding their distribution in Hokkaido. Brook trout have shown relatively limited invasion in somespring-fed streams and ponds both in Honshu and Hokkaido. The introduced exotic trout have been
implicated in reducing populations of native fishes, especially stream salmonids, through predation,competitive interaction for resources, and interspecific hybridization. Monitoring, management,
including eradication, and evaluation of the effectiveness of these measures should be carried out tocontain the ecological risks of trout invasion.
Key words: brook trout, brown trout, competition, hybridization, non-native species, predation,
rainbow trout

1. Introduction
Invasion by non-native species is a serious threat
to the conservation of freshwater ecosystems (Fuller etal., 1999; Rahel, 2002). Of the non-native species
invading aquatic ecosystems, salmonid fishes (subfamily salmoninae: genera Oncorhynchus, Salmo andSalvelinus) are among the globally most widespreadtaxa (Welcomme, 1988). Many species of salmonids
have been introduced for recreational and commercial
purposes to regions throughout the world where they
did not occur naturally, with little regard to the effects
on native species (Fausch, 1988). In most cases,
salmonid introductions have been widely implicatedin declines of native biota (e.g., Polhemus, 1993;
Lassuy, 1995). In northern hemisphere freshwater
ecosystems, such as in North America and Europe,
which generally support one or more native salmonids,
the potential for impact of non-native salmonids onnative species are widely reported (e.g., Allendorf &Leary, 1988; Behnke, 1992). Because non-native
salmonids are biologically similar to native salmonids,
there is evidence of strong interspecific interactions
for resources (Fausch, 1988) and hybridization (Learyet al., 1995). Also in the southern hemisphere, suchas New Zealand and Australia, where no native
salmonids naturally occur, impact of introduced
Global Environmental Research8(1)/2004: 41-50
printed in Japansalmonids on native fish and invertebrates is widely
recognized (e.g., Townsend, 1996; Moyle & Light,
1996).
By the early 1990s, three salmonids, rainbow trout(Oncorhynchus mykiss), brown trout (Salmo trutta)
and brook trout (Salvelinus fontinalis), were among 12
fish species introduced to more than 30 countries(Welcomme, 1992). The former two salmonids are
the species with the biggest impact on biological diversity, being listed in ‘100 of the World’s Worst Invasive Alien Species’ by the IUCN (2000). Rainbow
trout, whose native range stretches along the Pacificcoast of North America and the Kamchatka Peninsula,
have been introduced to 87 countries worldwide(Welcomme, 1992) and all U.S. states outside theirnative range (Fuller et al., 1999), primarily to establish populations for recreational angling. Establishment of reproducing populations was highly successful in the Southern Appalachians of North America,
moderately successful in the Central Rocky Mountains of North America, the Andes Mountains in SouthAmerica, as well as in New Zealand and northern
Japan, but failed in Scandinavia and central Europe(Fausch et al., 2001). The rainbow trout hybridized
with other, more rare trout species, such as inland cutthroat trout (Oncorhynchus clarki), thereby affecting
their genetic integrity (e.g., Allendorf & Leary, 1988;©2004 AIRIES42

S. KITANO

Weigel et al., 2003). Stocking of hatchery rainbow
trout in rivers has led to introduction of whirlingdisease into open waters in some parts of the UnitedStates (Fuller et al., 1999). Brown trout have a large
native range that encompasses the natural boundaries
of Europe, extending south to North Africa and east to
the Ural Mountains and the Caspian Sea (Elliott,1994). The earliest known introduction beyond its
native range was made in Tasmania and New Zealandin the 1860s (MacCrimmon & Marshall, 1968).
They have been implicated in extirpating an endemic
grayling Prototroctes oxyrhynchus in New Zealand(MacDowall, 1990) and in reducing native salmonid
populations, through predation, displacement, andfood competition in North America (Taylor et al.,1984). Brook trout are native to northeastern North
America, but have become one of the most widespread non-native salmonid species in western North
America (Dunham et al., 2002) and have been introduced worldwide as a popular sport fish (Scott &Crossman, 1973). Introduced brook trout have replaced native cutthroat trout and golden trout (Oncorhynchus aquabonita) in some streams in western
North America (Moyle, 1976; Behnke, 1992).
Hybridization between native bull trout (Salvelinus
confluentus) and introduced brook trout is widespreadin North America (Howell & Buchanan, 1992; Learyet al., 1993).
Numerous studies have investigated the impact of
introduced trout on native biota (reviewed by Fausch,
1988; Moyle & Light, 1996; Townsend, 1996;
Dunham, 2002). However, most studies have been
conducted only in a few regions, primarily North
America, Australia and New Zealand. Studies ofother geographical regions with different biotic andabiotic conditions are needed for a general understanding of the impact of introduced trout.
In Japan, translocations of non-native salmonid
fishes from overseas began in the late 1800s for utilization as commercial or recreational fishery resources(Yuma et al., 1998). Rainbow and brook trout werefirst introduced into Japanese waters in 1877
(Kawanabe, 1980; EAGJ, 1982), and brown trout
were introduced in 1892 (Elliott, 1989). Since 1926,
when regulations for salmon and trout enhancement
came into enforcement by the Japanese government,
substantial efforts were put into importation andhatching of their fry, and this has continued until recent years (EAGJ, 1982). For instance, a total ofmore than 106 eggs of rainbow trout were imported
from the United States from 1877 to 1934 (Maruyama
et al., 1987). A large number of fry, juveniles andadults were introduced each year for decades, or escaped from fish hatcheries into Japan’s waters. Production and distribution of eggs or fry of rainbow,
brown and brook trout by trout hatcheries still continue throughout Japan (Maruyama et al., 1987).
Since these intensive introductions have resulted in
the establishment of reproducing populations in Japa-

nese inland waters, concerns have emerged about the
impact of introduced trout on native biota involving
endemic Asian salmonids such as masu salmon (Oncorhynchus masou), white-spotted charr (Salvelinus
leucomaenis) and Sakhalin taimen (Hucho perri),stream communities and ecosystems.
In this paper, I begin by reviewing the patterns andcauses of invasion of the three trout species, and thepotential impact of exotic trout on native fauna. Following this review, I offer suggestions for managing
the risk of exotic trout invasion and its impact on
Japanese inland waters.

2. Invasion of Rainbow Trout
There are records of rainbow trout occurrence from
Kyushu, Shikoku, Honshu and Hokkaido (Fig. 1(a)).
Donor stocks of rainbow trout introduced from North
America to Japan are of mixed lineage, including
stream resident and anadromous races, and hatcheryand wild stocks (e.g., various strains by the name of
Donaldson, Westalian and Steelhead trout are included: Maruyama et al., 1987; Fausch et al., 2001).
Rainbow trout were first introduced to Honshu, but
this resulted mostly in failure (Kawanabe, 1980;
Fausch et al., 2001), despite frequent stocking inmany locations by commercial fishermen’s unions andby the public during the last 50 years (e.g., 20 × 106
juveniles in 1988; see Yuma et al., 1998). Evidenceof only a few cases of natural reproduction have beenobtained in habitats in cold lakes with inlet systems orcold low gradient streams in Honshu (Kato, 1985;
Nakamura & Maruyama, 1994; Kitano et al., 2003).
Limited invasiveness presumed attributable to highangling pressure, bird predation at the time of release,high water temperatures and/or flood disturbance atthe time of fry emergence (Kawanabe, 1980). In
contrast, the rainbow trout invasion of the island of
Hokkaido was a significant success (Fausch et al.,2001), with records of occurrences gradually increaseing since 1970, ultimately resulting in a distribution of
72 sites, including every major river system, by 1996
(Takami & Aoyama, 1999). Most of these populations appear to be self-sustaining (Aoyama et al.,1999b; Taniguchi et al., 2000). Rainbow trout attainsexual maturity at 11 to 37 cm FL (fork length) at agesof 1 to 5+ in males and 17 to 33 cm FL at ages 2 to 4+
in females in spring-fed streams in Hokkaido (Kitano
et al., 1993). Rainbow trout typically spawn in late
winter and early spring, the date varying among populations. In Hokkaido, they spawn from April to June(range of water temperature 5 to 12°C: Aoyama et al.,1999b) in upper Shiribetsu River, but from January to
April in Horonai Creek (4 to 10°C: see Kitano et al.,
1993; Taniguchi et al., 2000). In central Honshu,
spawning takes place in April to June in the upper
Shinano River (Kitano et al., 2003) and in Lake
Nozori (Nakamura & Maruyama, 1994), but in December in the upper Tama River system (Kato, 1985).

Ecological Impacts of Invasive Exotic Trout in Japan

Rainbow trout prey primarily upon aquatic andterrestrial invertebrates, but eggs and larvae of thenative river sculpin (Cottus nozawae) constitute 10%of their diet in Horonai Creek, Hokkaido (Kitano et al.,1993). They also prey on the eggs and fry of chumsalmon (Oncorhynchus keta), the fry of masu salmon(O. masou) and, on rare occasions, voles in streams in
Hokkaido (Hikita et al., 1959; Taniguchi et al., 2002).

(a) Rainbow Trout(b) Brown Trout(c) Brook Trout

Fig. 143

Recorded location of occurrence (open circles) and natural reproduction (solid circles) of introduced(a) rainbow trout, brown trout (b) and brook trout (c). Occurrence data were obtained from the MEGJ
(2002) survey largely conducted from1960 to 2000. See text for reports of natural reproduction by introduced
trout.

44

S. KITANO

The fry of white-spotted charr (S. leucomaenis) werealso found in the diet of trout in the Honshu stream(Kitano et al., 2003). Rainbow trout occupy undercut banks that native fish normally use for refuge(Kitano et al., 1993; Urabe & Nakano, 1998), and may
be displacing native fish from preferred microhabitats
and pushing them into open water, making themvulnerable to predation.
Larger rainbow trout
dominate smaller masu salmon by aggressive interference, relegating them to less favorable foraging
positions downstream and reducing their foraging
frequency and growth in an experimental stream
channel (Taniguchi et al., 2002). The trout havebeen implicated in reducing native Dolly Varden charr(Salvelinus malma) on the Shiretoko Peninsula(Morita et al., 2003). Because habitat conditions ofspawning sites are quite similar among salmonid
fishes (i.e., gravel beds in shallows), competition for
spawning space may occur between rainbow trout andnative salmonids. Spring-spawning rainbow trout
reduce survival of eggs and alevins of fall-spawningsalmonids (white-spotted charr and Dolly Varden
charr) by dislodging the spawning redds (Taniguchi etal., 2000). They may potentially superimpose on theredds of the endangered Sakhalin taimen (Hucho
perryi) as an earlier spring-spawner in Shiribetsu
River, Hokkaido (Aoyama et al., 1999b).

3. Invasion of Brown Trout
Brown trout were first introduced into Japan fromthe United States, together with the eggs of rainbow
and brook trout in the late 1800s (Maruyama et al.,1987). Recently brown trout are distributed in
Honshu and Hokkaido (Fig. 1(b)). Natural reproduction has been reported in upper Azusa Creek (EAGJ,
1982) and in the inlets of Lake Chuzenji(Wakabayashi et al., 2002, 2003) in central Japan, andalso in Lake Shikotsu and its outlet stream in
Hokkaido (Kaeriyama, 2002; Aoyama et al., 2002),
but is not evident for most stocks in Japan. In
Hokkaido, the release of brown trout by sport anglers
has caused a rapid increase in records of occurrences
since 1980, ultimately reaching 48 sites distributed in
36 river systems (Takami & Aoyama, 1999;
Kaeriyama, 2002). In addition, the trout may extend
their distribution by means of sea-run migration(anadromous form) in Hokkaido (Aoyama et al.,1999a; Arai et al., 2002; Morita et al., 2003). Likeother salmonids, brown trout stocked in inlet-lakesystems have fluvial-lacustrine (lake-run form) life
histories (Misawa et al., 2001; Wakabayashi et al.,2002). The trout have been known to grow to over1 m in length and to live up to 18 years (McGinnis,
1984). The maximum total length is 84 cm in Lake
Shikotsu (Takayama et al., 2002) and 70 cm in Lake
Chuzenji (Wakabayashi et al., 2002). The trout
attain sexual maturity at 12 to 36 cm FL (ages 1 to 3+)
in males and 20 to 38 cm FL (ages 2 to 4+) in females

in a Hokkaido stream, with a growth pattern similar tothat in its native region (Aoyama et al., 2002).
Spawning takes place during mid-October to December in inlets of Lake Chuzenji (Wakabayashi et al.,2002) and during December to January in inlets ofLake Shikotsu (Kaeriyama, 2002).
Brown trout have displaced native white-spottedcharr in a tributary of the upper Ishikari River in
Hokkaido over the last 15 years (Takami et al., 2002).
The trout have been implicated in reducing the abundance of native goby (Rhinogobius sp. OR) in theshallow areas of Lake Shikotsu during the last decade(Takayama et al., 2002). In Lake Chuzenji, originally a fish-less lake, brown trout prey upon aquatic
invertebrates in spring, terrestrial insects in summer,and fish (Salmonidae, Cyprinidae and Gobiidae)
throughout the year (Shiraishi & Tanaka, 1967). The
trout, showing a piscivorous nature, consume
threespine stickleback (Gasterosteus aculeatus),
kokanee salmon (Oncorhynchus nerka) and whitespotted charr in Lake Shikotsu (Kaeriyama, 2002),
newly emerged masu salmon fry in Chitose River(Mayama, 1999) and a goby (Gymnogobious sp.) in a
stream on Shiretoko Peninsula (Morita et al., 2003).
Fall-spawning (or winter-spawning) brown trout havebeen implicated in the destruction of formerly spawning masu salmon, thereby reducing the breeding success of that species (Wakabayashi et al., 2002).
Although it rarely occurs, brown trout are known tohybridize with native white-spotted charr in natural
habitats of northern Honshu (Sugiyama, 1997).

4. Invasion of Brook Trout
Occurrences of brook trout have been recorded in
14 prefectures of Honshu and Hokkaido (Fig. 1(c)).
Judging from the records of occurrences, stocking ofbrook trout is not so popular compared with that of theother two trouts in Japan. Establishment of reproducing populations has been reported in Nijibetsu
Creek in Hokkaido (Hikita et al., 1959), inlets of Lake
Chuzenji (Wakabayashi et al., 2002) and Lake Yunoko(Shiraishi & Takagi, 1955), and in upper Azusa Creek(EAGJ, 1982) in central Honshu, sites characterizedby spring branches of low gradient streams or cold
mountain lakes. Brook trout spawn in early November
to December, attaining sexual maturity at 11 to 26 cm
FL (Kitano & Uehara, 2002).
Brook trout are carnivorous, and feed upon a wide
range of organisms (Kitano, 2002). Brook trout feed
upon benthic invertebrates more intensively than dorainbow trout in Lake Yunoko (Shiraishi & Takagi,1955), and upon eggs and fry of chum salmon in
Hokkaido (Hikita et al., 1959). Brook trout havehybridized with white-spotted charr, a congenericnative charr in an inlet of Lake Chuzenji (Suzuki &Kato, 1966) and in upper Azusa Creek (EAGJ, 1982).
Asymmetric hybridization in which male brook trout
cross with female white-spotted charr has beenEcological Impacts of Invasive Exotic Trout in Japan

implicated in reduction of native white-spotted charr(Kitano, 2002). Redd superimposition may also havedetrimental effects on the breeding success of native
charr (Kitano & Uehara, 2002).

5. Factors Affecting Invasion Success andEcological Impacts of Trout
The successful invasion of non-native trout may be
primarily associated with environmental factors suchas temperature, habitat size, stream flow, and human
influences in Japan, as already discussed in the case ofother countries (Dunham et al., 2002). AmongJapan’s three exotic trouts, rainbow trout are particularly germane to the analysis of the relationshipbetween success of invasion and environmental characteristics, because of more intensive stocking
attempts nationwide than for any other species.
Rainbow trout have been successful in Hokkaido, buthave failed in most of Honshu. Differential vulnerability to disturbance is often invoked as an explanation for the success or failure of non-native fish
invasions (Moyle & Light, 1996). Fausch et al.
(2001) concluded that flood disturbance regimes
strongly influence the success of rainbow trout invasions based on empirical evidence from five Holarctic
regions, including Honshu and Hokkaido, withsuitable water temperatures. Highly successful trout
invasions are observed in regions where trout fry
emerge during a period of low flood probability as in
Hokkaido, whereas a high probability of floods produced by summer monsoon rains followed by fall
typhoons was apt to wash away emergent trout fry in
Honshu streams. Similar mechanisms may explain
the successful invasion of brown and brook trout, although such an analysis is difficult to apply due tolack of broad, quantitative data. Given that suchmechanisms are real, Japanese stream environments,
altered by damming during these several decades,
have become hospitable to invasion by exotic trout.
In mountainous regions, small, rapid streams haveoften been changed into significantly slower ones by
the construction of small dams for irrigations andflood control (see Yuma et al., 1998). Storage reservoirs have also been built in the middle to upper
reaches of most of Japanese rivers for power production, agricultural, industrial water use and flow
regulation. These alterations may increase the invasive success of exotic trout because storage reservoirs
generally reduce peak flow (Raymond, 1988).
The number of introduced propagules, frequency
of introduction, characteristics of the donor populations as in term of genetic composition, and anglingpressure may also influence the success of invasion(e.g., Simberloff, 1989). Intensive introduction andrelatively low fishing pressure may relate to the rapid
expansions of rainbow and brown trout in Hokkaido.
The distribution of non-native trout has increased
since the 1970s when sports fishing using lures be-

45

came popular in Japan, with detailed information on
stockings by private individuals unavailable (seeTakami & Aoyama, 1999). Sea-migratory habits
have been suggested as another means of dispersion ofbrown trout in Hokkaido, even though the strains
introduced into Japan are considered to be nonmigratory (Aoyama et al., 1999a). Regardless oforiginal strain, transition from non-migratory to seamigratory has often been reported in North America(Dymond, 1963) and the Falkland Islands (McDowall
et al., 2001). It is still unclear whether introduced
rainbow and brown trout are reproducing naturally
even in Hokkaido, except in some streams. However,regardless of establishment of self-reproducing populations, large and long-lived trout can have detrimental
effects on native species for a long period.
Based on the available ecological information, I
conclude the three exotic salmonid fishes have significant impacts on native fauna in Japan, especially onnative freshwater fishes. However, the degree ofinvasive success and the processes and mechanisms of
ecological impact differ among trout species andamong regions. Predation on native fishes and interspecific competition with native salmonids has been
evident for the three exotic trout, but hybridization
with native species is almost specific to brook trout.
Native salmonid fishes with the potential to interact
with introduced trout differ between Honshu andHokkaido. In Honshu, landlocked masu salmon andwhite-spotted charr are native and may potentiallyinteract with exotic trout in cold water habitats,generally isolated areas in upper reaches or headwaters. In Hokkaido, four salmonids including anadromous forms: masu salmon, white-spotted charr,
Dolly Varden, and Sakhalin taimen, have the potential
to interact with introduced trout in low- to highaltitude streams and lakes.
The non-native trout have potential effects on
native fish species through predation and competition.
Serious impacts of brown trout have been noted onwhite-spotted charr (Takami et al., 2002) and native
goby (Rhinogobius sp. OR) in Hokkaido (Takayama etal., 2002), although extinction of native fish as a resultof invasion of these non-native trout has not been evident in Japan. Intensive predatory and competitive
impacts by brown trout have been implicated forNorth America, New Zealand and Falkland Islands(e.g., Taylor et al., 1984). For instance, invasive
brown trout preyed directly on native fishes and were
responsible for the local extinction and population
fragmentation of certain species of the family
Galaxiidae in New Zealand (McIntosh et al., 1992)
and in the Falkland Islands (McDowall et al., 2001).
Introduced rainbow trout also exerted a major
negative effect on the endangered humpback chub(Gila cypha) through predation in the Colorado River(Marsch & Douglas, 1997). Fausch and White(1981) revealed that adult brown trout displaced
native brook trout from the best habitats through46

S. KITANO

competition in a Michigan stream. Cutthroat trout
have been displaced from much of their native ranges
in the Rocky Mountains of the western United Statesby introduced brook trout, probably due to competitive interaction (Gresswell, 1988). Competition for
spawning resources (i.e., redd superimpositions or
dislodging by later-spawning species) seems to be
crucial for Japanese salmonids, because non-native
salmonids tend to spawn later than native species (e.g.,
Taniguchi et al., 2000).
Asymmetric hybridization is suspected to occur
between introduced brook trout and native whitespotted charr in a headwater stream in central Japan(EAGJ, 1982; Kitano, 2002). Introduced brook trout
have been displacing resident bull trout, an endangered salmonid species, in parts of northwestern North
America through asymmetric hybridization (Learyet al., 1993). According to mtDNA analysis, malebrook trout were thought to mate with female bull
trout, yet allozyme data indicated that about 97 % of
detected hybrids were first generation individuals.
Because brook and bull trout hybridization produces
mainly sterile progeny and no individuals beyond thebackcrosses have been detected, this hybridization
does not constitute a threat of introgression. Even
without substantial mixing, however, this hybridization can have serious consequences to the conservation of bull trout, because the more numerous brooktrout may have an advantage by wasting the smaller
reproductive potential of the bull trout with hybrid
production. On the other hand, hybridization between rainbow trout and native salmonids has notbeen reported in Japan, most likely due to differencesin spawning season, whereas introgressive hybridization with rainbow trout have often resulted in loss ofdiversity of native cutthroat trout in western U.S.
watersheds (Allendorf & Leary, 1988; Weigel et al.,2003).
There is also no doubt that native species suffer
more from the presence of two or more introduced
species in the same stream. Brown and brook trout
occur with native white-spotted charr in sympatry inupper Azusa Creek (EAGJ, 1982). Similarly, brown
and rainbow trout occur in inlets of Lake Shikotsu(Kikuchi et al., 2001) and in other Hokkaido lakes(Takami & Aoyama, 1999).
Since competitive
interaction between introduced trout generally leads todifferential resource utilization between them (seeKikuchi et al., 2001), inferior native species may beforced to utilize more depleted resources as theintroduced trout species increase (Nakano et al., 1998;
Nakano et al., 1999b; Taniguchi et al., 2002).
Parasite or disease transmission is another possible
influence on native species, but evidence for thismechanism is rarely available in Japan. However,since viral, bacterial and fungal diseases have frequently induced great losses in salmon and trout
hatcheries in Japan, diseases transmission could havecertain negative effects on native fishes (Nagasawa,1989). For instance, Yoshimizu et al. (1989) reported infectious hematopoietic necrosis (IHNV) in
wild chum and masu salmon in Hokkaido, which wasthought to have been imported with salmon eggs from
North America. This viral disease has become
widely distributed among most of the hatcheries from
Hokkaido to Kyushu (Nagasawa, 1989). Thus, transplantation of eggs and fishes may potentially transmit
disease nationwide. Inversely, whirling disease parasites (Myxobolus cerebralis) in the native European
range of brown trout are believed to be an importantfactor in their resistance to rainbow trout invasion(Lever, 1996).
Indirect effects are a common feature in animalcommunities (Sih et al., 1985; Pimm, 1991). The
impacts of invasive species on individual species are
expected to be propagated through other species in the
food-web. In relatively species-poor aquatic communities such as headwater streams, ‘top-down’trophic cascades are likely to occur (Strong, 1992).
Introduced brown trout profoundly affect the biotic
interactions in stream communities, reducing theabundance of grazing invertebrates and altering theirgrazing behavior so that algal biomass increases(Flecker & Townsend, 1994; McIntosh & Townsend,1994). More recently, Nakano et al. (1999a) havesuggested that invading rainbow trout also have strong
effects that could cascade through aquatic-terrestrial
food linkages and potentially affect forest birds living
near a Hokkaido stream. They showed that rainbow
trout actively selected terrestrial prey over aquaticprey, because of the larger size of the terrestrial preyand their peak flux into streams during the evening.
Consequently, rainbow trout consumed 77% of thetotal input of terrestrial prey into a stream reach
during mid-summer, and this constituted 73% of their
daily ration. Thus, if the input of terrestrial prey is
reduced by modification of riparian habitat, rainbow
trout will be forced to forage more intensively onaquatic invertebrates. Moreover, when drifting prey
are scarce, native Dolly Varden charr shift foraging
modes to benthos feeding (Fausch et al., 1997;
Nakano et al., 1999b). In either case, stream
salmonids have a potential for strong effects onaquatic invertebrates, their emergence as winged
adults and, ultimately, the flux of aquatic prey thatsubsidizes forest birds (Fausch et al., 2002).

6. Implications for Practical Management of
Trout
Japan’s domestic laws and regulations are insufficient to avoid ecological risks posed by exotic trout.
The rainbow trout is one of the most popular fish species in Japanese inland fisheries, being officially released to inland waters of 31 prefectures throughout
Japan (Fig. 2(a)). In contrast, legislation or regulations prohibiting stocking of trout are limited to only
three prefectures (Fig. 2(b)). The government ofEcological Impacts of Invasive Exotic Trout in Japan

Hokkaido, where rapid expansion of non-native trout
has been recognized, has prohibited stocking of brownand brook trout (but not rainbow trout) by regulations
with penalties of a maximum of six months’ penal
servitude and 100,000 yen since November 2003.
Under existing law, however, there are no regulations
to controlling the trout populations previously stocked
to the waters. Because catch-and-release fishing iscommon for sport anglers, stock depletion would not
be expected even where angling intensity is high.
There would seem to be an urgent need for additional
regulations encouraging catch-and-keep fishing of
non-native trout in special conservational areas suchas national parks or waters protected for fisheries resources. Furthermore, establishment of systematic
processes of screening, monitoring and feedback will
be required in order to contain ecological risks (Ham
& Pearson, 2001). Native salmon and charr should
be utilized as commercial resources more often than
non-native trout, since eggs and fry of native species
have become available recently at a reasonable cost.
In Japanese cold water streams and lakes, there areseveral endemic salmonid species or subspecies endangered due to human impacts such as habitat degradation or overfishing (MEGJ, 2003). For instance,
Sakhalin taimen, Dolly Varden charr and Miyabe charr(Salvelinus malma miyabei) in Hokkaido are listed as‘Critically Endangered taxa,’ and the southernmost
populations of Yamato charr (S. leucomaenis
japonicus) and Gogi charr (S. l. imbrius) in Honshu
are currently listed among ‘Threatened Local Populations.’ Yamamoto et al. (2004) have indicated thatwhite-spotted charr contain more pronounced genetic
variation at a population level based on molecularphylogenetic analyses of mtDNA. Habitats with endangered species or with populations of genetic
uniqueness should have a management priority.

Fig. 2

47

When there are adverse impacts from establishedpopulations of exotic trout, direct control can be applicable for their management. For instance, removal
by electrofishing, selective angling, or toxicants has
been used to control brook trout in non-native areas of
the United States (e.g., Bettoli & Maceina, 1996).
Eradication of invasive trout by electrofishing is more
likely to be effective in smaller streams, but not inlarge rivers or lakes. Toxicants can be applicable tocomplete eradication of invasive trout, and are sometimes used to control invasions in the United States(e.g., Gresswell, 1991). Dispersal barriers are installed at the down-stream end of the treated section of
streams to prevent re-invasion by non-native trout(Thompson & Rahel, 1996; Dunham et al., 2002;
Novinger & Rahel, 2003). There are, however, few
case studies about such direct control of invasive trout
in Japan. Eradication is most feasible for brook trout
with distribution restricted to headwater area, but isdifficult for rainbow and brown trout inhabiting wider
streams or lakes. Since spawning trout usually entersmall tributaries from larger streams or lakes andspawn in shallow areas with fine gravel suitable totrout, a precise detection of spawning sites andseasons may lead to effective management of brown
and rainbow trout.
Eradication of invasive trout may, on the other
hand, have short- and long-term unpredicted consequences for the native fauna and ecosystem.
Strong non-selective eradication may be detrimental
not only to invasive species but also to native species
in terms of loss of genetic diversity (e.g., Avise &Hamrick, 1996). In cases that invasive trout play amajor role in an aquatic community, rapid loss ofinvasive trout may seriously affect the viability ofnon-target native species. For example, parasitic
larvae of an endangered freshwater mussel (Marga-

Areas with permission to utilize the exotic trout as fisheries resources and (a) with regulations
on trout stocking (b). This information was provided by the Fisheries Agency, Government ofJapan, dated 2 December, 2003.

48

S. KITANO

ritifera laevis), utilizing invasive brook trout as hosts
instead of scarce native white-spotted charr (Kondo etal., 2000), may be threatened by the drastic removalof the invasive trout. Similarly, the piscivorous
Japanese water shrew (Chimarrogale platycephala)would be influenced by low food availability immediately after trout eradication (see Abe, 2003). To consider the full range of impacts associated with directcontrol, managers can evaluate short-term threats by
non-native trout and long-term considerations for persistence of native populations and communities.
Most recently, the number of artificial trout fishingponds and streams connected to natural streams havebeen increasing from southern to northern regions ofJapan (e.g., 98 sites were listed in the fishing guidebook edited by Murata, 2003). They have stocked
various types of exotic trout including a triploid typeof rainbow trout, hybrid trout (between brown trout
and brook trout or between brook trout and whitespotted charr) and arctic grayling (Thymallus arcticus)
in addition to rainbow, brown and brook trout.
Escapes of fish from the facilities may cause novel
ecological issues regarding the conservation ofJapanese freshwater fishes. In fact, accidental stocking of arctic grayling, originally cultured for the fishing ponds, to reservoirs has been detected in central
Japan (Kitano et al., 2004). Management of exotic
trout stocking sources is also important in controlling
the likelihood of stocking mishaps, thereby preventing
the problem from increasing.
Finally, our knowledge of invasive trout is still
limited in Japan. We need to know more about theinvasion process: how they arrive, how their arrival
can be blocked, how to handle problems when theyarise and how to evaluate the overall effectiveness of
our management programs. The feedback process of
research, monitoring, management action and evaluation may develop more robust theories for predicting
invasion success.

Acknowledgements
I thank Drs. Kentaro Morita, Satoshi Yamamotoand Shoichiro Yamamoto for their stimulating
discussion about ecology of introduced salmonid
fishes, and Azusa Kosaka for providing invaluable
information on the fisheries laws and regulations.
This work was partly funded by the Japanese Ministryof the Environment (F-3: 2001-2003).
References
Abe, H. (2003) Trapping, habitat, and activity of the Japanesewater shrew, Chimarrogale platycephala. Mammalogical
Science, 43: 51-65. (in Japanese with English Abstract)
Allendorf, F. W. and R. F. Leary (1988) Conservation anddistribution of genetic variation in a polytypic species, thecutthroat trout. Conservation Biology, 2: 170-184.
Aoyama, T., K. Naito and T. Takami (1999a) Occurrence of

sea-run migrant brown trout (Salmo trutta) in Hokkaido,
Japan. Scientific Report of Hokkaido Fish Hatchery, 53:
81-83.
Aoyama, T., T. Takami, M. Fujiwara and H. Kawamura (1999b)
Natural reproduction of rainbow trout, Oncorhynchus
mykiss, in the Shiribetsu River in Hokkaido, Japan.
Scientific Report of the Hokkaido Fish Hatchery, 53: 29-38.
(in Japanese with English Abstract)
Aoyama, T., T. Takami, K. Shimoda and T. Koyama (2002) Age,
growth and sexual maturity of brown trout, Salmo trutta, in
Hokkaido, Japan. Scientific Report of the Hokkaido Fish
Hatchery, 56: 115-123. (in Japanese with English Abstract)
Arai, T., A. Kotake, T. Aoyama, H. Hayano and N. Miyazaki(2002) Identifying sea-run brown trout, Salmo trutta, using
Sr:Ca ratios of otolith. Ichthyological Reasearch, 49:
380-383.
Avise, J. C. and J. L. Hamrick, eds. (1996) Conservation
genetics: case history from nature. Chapman & Hall, NewYork.
Behnke, R. J. (1992) Native trout of western North America.
American Fisheries Society Monograph 6, Bethesda,
Maryland.
Bettoli, P. W. and M. J. Maceina (1996) Sampling with
toxicants. In: B. R. Murphy and D. W. Willis, eds., Fisheries
tequniques, 2nd edition. American Fisheries Society,
Bethesda, Maryland, pp. 303-333.
Dunham, J. B., S. B. Adams, R. E. Schroeter and D. C.
Novinger (2002) Alien invasion in aquatic ecosystems:
Toward an understanding of brook trout invasions andpotential impacts on inland cutthroat trout in western North
America. Reviews in Fish Biology and Fisheries, 12:373-391.
Dymond, J. R. (1963) Family Salmonidae. In: H. Shultz, D.
Merriman and J. Tee-Van, eds., Fishes of the western North
Atlantic, Part 3. Sears Foundation for Marine Research, Yale
University, New Haven, pp. 455-502.
Elliott, J. M. (1989) Wild brown trout Salmo trutta: an
important national and international resource. Freshwater
Biology, 21: 1-5.
Elliot, J. M. (1994). Quantitative Ecology and the Brown Trout.
Oxford University Press, Oxford.
Environment Agency, Government of Japan (EAGJ) (1982)
Scientific Report on Conservation of Charrs in the
Headwater Drainage of Azusa River. Japan Environmental
Division, Tokyo. (in Japanese)
Fausch, K. D. and R. J. White (1986) Competition between
brook trout and brown trout for positions in a Michigan
stream. Canadian Journal of Fisheries and Aquatic Sciences,38: 1220-1227.
Fausch, K. D. (1988) Tests of competition between native andintroduced salmonids in streams: what have we learned?
Canadian Journal of Fisheries and Aquatic Science, 45:
2238-2246.
Fausch, K. D., S. Nakano and S. Kitano (1997) Experimentallyinduced foraging mode shift by sympatric charrs in a
Japanese mountain stream. Behavioral Ecology, 8: 414-420.
Fausch K. D., Y. Taniguchi, S. Nakano, G. D. Grossman and
C. R. Townsend (2001) Flood disturbance regimes influence
rainbow trout invasion success among five Holarctic regions.
Ecological Applications, 4: 798-807.
Fausch, K. D., M. E. Power and M. Murakami (2002) Linkages
between stream and forest food webs: Shigeru Nakano’s
legacy for ecology in Japan. Trends in Ecology and
Evolution, 17: 429-434.
Flecker, A. S. and C. R. Townsend (1994) Comminity-wide
consequences of trout introduction in New Zealand streams.
Ecological Applications, 4: 798-807.
Fuller, P. L., L. G. Nico and J. D. Williams (1999) Nonindigenous fishes introduced into inland waters of the UnitedEcological Impacts of Invasive Exotic Trout in Japan
States. American Fisheries Society, Special Publication 27,
Bethesda, Maryland.
Gresswell, R. E., ed. (1988) Status and Management of
Cutthroat Trout. American Fisheries Society Symposium 4,
Bethesda, Maryland.
Gresswell, R. E. (1991) Use of antimycin for removal of brooktrout from a tributary of Yellowstone Lake.
North
American Journal of Fisheries Management, 11: 83-90.
Ham, K. D. and T. N. Pearsons (2001) A practical approach forcontaining ecological risks associated with fish stocking
programs. Fisheries, 26: 15-23.
Hikita, T., S. Kameyama, A. Kobayashi and Y. Sato (1959)
Some biological observations on the rainbow trout Salmo
gairdneri irideus in the Nishibetsu River, with specialreference to the consumption of salmon eggs and fry by
predaceaous fishes. Bulletins of National Institute for
Salmon Hatchery, 14: 91-121. (in Japanese with English
Abstract)
Howell, P. J. and D. V. Buchanan, eds. (1992) Proceedings ofthe Gearhart Mountain Bull Trout Workshop. American
Fisheries Society, Oregon Chapter, Corvallis.
The World Conservation Union (IUCN)(2000) Booklet on 100of the World’s Worst Invasive Alien Species. (Available fromthe internet URL http://www.issg.org/database)Kaeriyama, H. (2002) Salmo trutta. In: The ecological societyof Japan, ed., Handbook of Alien Species in Japan.
Chijinshokan Press, Tokyo, pp. 113. (in Japanese).
Kato, K. (1985) Natural spawning of rainbow trout in the upper
Tama River, Tokyo. Bulletin of the Japanese Society of
Scientific Fisheries, 51: 1947-1953. (in Japanese withEnglish Abstract)
Kawanabe, H. (1980) Rrainbow trout, Oncorhynchus mykiss.
In: T. Kawai, H. Kawanabe and N. Mizuno, eds., Freshwater
Organisms in Japan: Their Ecology of Invasion andDisturbance. Tokai University Press, Tokyo, pp. 44-48. (inJapanese).
Kitano, S., S. Nakano, M. Inoue, K. Shimoda and S. Yamamoto(1993) Feeding and reproductive ecology of exotic rainbow
trout Oncorhynchus mykiss in the Horonai Stream in
Hokkaido, Northern Japan. Nippon Suisan Gakkaishi, 59:
1837-1843. (in Japanese with English Abstract)
Kitano, S. (2002) Salvelinus fontinalis. In: The ecologicalsociety of Japan, ed., Handbook of Alien Species in Japan.
Chijinshokan Press, Tokyo, p. 114. (in Japanese).
Kitano, S. and T. Uehara（2002）Nest superimposition by exotic
brook, brown trout and hybrids on native charrs in a
Japanese stream．In: Proceeding of the VIII International
Congress of Ecology (INTECOL), Seoul, Korea, 138p.
Kitano, S., Y. Kawai and H. Ida (2003) Ecological aspects ofrainbow trout and white-spotted charr in a headwater
tributary of Zako Stream, central Japan. Bulletin of theInstitute of Nature Education in Shiga Heights, 40: 9-13. (inJapanese with English Abstract)
Kitano, S., S. Kobayashi, T. Yamagata and T. Uehara (2004)
Potential ecological impacts by arctic grayling Thymallus
arcticus stocked to a tributary of upper Kiso River System,
Nagano Prefecture. Bulletins of Nagano Nature Conservation Research Institute. (in Japanese)
Kondo, T., M. Yamada, Y. Kusano and K. Sakai (2000) Fishhosts of the freshwater mussel Margaritifera laevis(Bivalvia: Margaritiferidae) in the Furebetsu River,
Hokkaido. Venus, 59: 177-179.
Lassuy, D. R. (1995) Introduced species as a factor in extincttion and endangerment of native fish species. American
Fisheries Society Symposium, 15: 391-396.
Leary, R. F., F. W. Allendorf and S. H. Forbes (1993) Conservation genetics of bull trout in the Columbia and Klamath
River drainages. Conservation Biology, 7: 856-865.
Leary, R. F., F. W. Allendorf and G . K. Sage (1995)

49

Hybridization and introgression between introduced andnative fishes. American Fisheries Symposium, 15: 91-101.
Lever, C. (1996) Naturalized Fishes of the World. Academic
Press, San Diego, California, USA.
MacCrimmon, H. R. and T. L. Marshall (1968) World distribution of brown trout, Salmo trutta. Journal of the Fisheries
Research Board of Canada, 25: 2527-2548.
Marsh, P. C. and M. E. Douglas (1997) Predation by introduced
fishes on endangered humpback chub and other native
species in the Little Colorado River, Arizona. Transactionsof the American Fisheries Society, 126: 343-346.
Maruyama, I., K. Fujii, T. Kimura and H. Maeda (1987) A
Historical Report on Introductions of Non-native Fish
Species into Japan. Fisheries Agency, Government ofJapan, Tokyo, pp. 1-157. (in Japanese)
Mayama, H. (1999) Predation of juvenile masu salmon(Oncorhynchus masou) and brown trout (Salmo trutta) on
newly emerged masu salmon fry in the Chitose River.
Bulletins of the National Salmon Resources Center, 2: 21-27.
(in Japanese with English Abstract)
McDowall, R. M., R. M. Allibone and W. L. Chadderton (2001)
Issues for the conservation and management of Falkland
Islands freshwater fishes. Aquatic Conservation: Marine andFreshwater Ecosystems, 11: 473-486.
McGinnis, S. M. (1984) Freshwater Fishes of California.
University of California Press, London.
McIntosh, A. R., C. R. Townsend and T. A. Crowl (1992)
Competition for space between introduced brown trout
Salmo trutta L. and a native galaxiid Galaxias vulgaris
Stockell in a New Zealand stream. Journal of Fish Biology,
41: 63-81.
McIntosh, A. R. and C. R. Townsend (1994) Interpopulation
variation in mayfly anti-predator tactics: differential effectsof contrasting predatory fish. Ecology, 75: 2078-2090.
Ministry of the Environment, Government of Japan (MEGJ)(2002) The National Survey on the Natural Environment
Report of the Distributional Survey of Japanese Animals(Freshwater fishes). Biodiversity Center of Japan, Yamanashi.
(The data are partly available from the internet URL
http://www.biodic.go.jp)
Ministry of the Environment, Government of Japan (MEGJ)(2003) Threatened Wildlife of Japan: Red Data Book 2ndedition. Vol. 4, Pisces, Brackish and Freshwater Fishes.
Japan Wildlife Research Center, Tokyo. (in Japanese)
Misawa, K., M. Kukuchi, H. Nozawa and M. Kaeriyama(2001) Effects of rainbow and trout on the Lake Shikotsu
watershed. Research Report from the National Institute forEnvironmental Studies, Japan, 167: 125-132. (in Japanesewith English Abstract)
Morita, K., D. Kishi, J. Tsuboi and S. H. Morita (2003)
Rainbow trout and brown trout in Shiretoko Peninsula,
Hokkaido, Japan. Bulletins of the Shiretoko Museum, 24:
17-26. (in Japanese with English Abstract)
Moyle, P. B. (1976) Fish introduction in California: history andimpact on native fishes. Biological Conservation, 9: 101-118
Moyle, P. B. and T. Light (1996) Biological invasions of fresh
water: empirical rules and assembly theory. BiologicalConservation, 78: 149-161.
Murata, M., ed. (2003) A Guide to Artificial Fishing Ponds andStreams in Japan. Naigai publications, Tokyo. (in Japanese)
Nagasawa, K. (1989) Microval and fungal diseases in masu andamago salmon and charr cultured in Japan: a review.
Physiology and Ecology Japan, Special Volume 1: 547-558.
Nakamura, T. and T. Maruyama (1994) Spawning of rainbow
trout, Oncorhynchus mykiss in the Lake Nozori, central Japan.
Suisanzoshoku, 42: 7-13. (in Japanese with English Abstract)
Nakano, S., S. Kitano, K. Nakai and K. D. Fausch (1998)
Competitive interactions for microhabitat among introduced
brook charr, Salvelinus fontinalis, and native bull charr, S.

50

S. KITANO

confluentus, and westslope cutthroat trout, Oncorhynchus
clarki lewisi, in a Montana stream. Environmental Biologyof Fishes, 52, 345-355.
Nakano, S., Y. Kawaguchi, Y. Taniguchi, H. Miyasaka, Y.
Shibata, H. Urabe and N. Kuhara (1999a) Selective foraging
on terrestrial invertebrates by rainbow trout in a forested
headwater stream in northern Japan. Ecological Research,
14: 351-360.
Nakano, S, K. D. Fausch and S. Kitano (1999b) Flexible niche
partitioning via a foraging mode shift: a proposed mechanism for coexistence in stream-dwelling charr. Journal ofAnimal Ecology, 68: 1079-1092.
Novinger, D. C. and F. J. Rahel (2003) Isolation managementwith artificial barriers as a conservation strategy for cutthroat
trout in headwater streams. Conservation Biology, 17:772-781.
Pimm, S. L. (1991) The Balance of Nature: Ecological Issues
in the Conservation of Species and Communities. Universityof Chicago Press, Chicago.
Polhemus, D. A. (1993) Conservation of aquatic insects:
worldwide crisis or localized threats? American Zoology, 33,
588-598.
Rahel, F. J. (2002) Homogenization of freshwater faunas.
Annual Reviews in Ecology and Systematics, 33: 291-315.
Raymond, H. L. (1988) Effects of hydroelectric development
and fisheries enhancement on spring and summer Chinook
salmon and steelhead in the Columbia Basin. North
American Journal of Fisheries Management, 8: 1-24.
Scott, W. B. and E. J. Crossman (1973) Freswater fishes ofCanada. Bulletin 184, Fisheries Research Board of Canada.
Ottawa.
Shiraishi, Y. and M. Takagi (1955) Food habit of trout in Lake
Yunoko, with reference to the morphology of body anddigestive organs. Bulletin of the Freshwater Fisheries
Research Laboratory, 5: 11-30. (in Japanese with English
Abstract)
Shiraishi, Y. and M. Tanaka. (1967) Feeding habit of brown
trout, in Lake Chuzenji. Bulletin of the Freshwater Fisheries
Research Laboratory, 17: 87-95. (in Japanese with English
Abstract)Sih, A., P. Crowley, M. McPeek, J. Petranka and K. Strohmeier(1985) Predation, competition and prey communities: areview of field experiments. Annual Review of Ecology and
Systematics, 16: 269-311.
Simberloff, D. (1989) Which insect introductions succeed andwhich fail? In: J. A. Krake, H. A Mooney, F. di Castri, R. H.
Groves, F. J. Kruger, M. Rejmanek and M. Williamson, eds.,
Biological Invasions: A Global Perspective. John Wiley andSons, Chichester, UK, pp. 61-75.
Strong, D. R. (1992) Are trophic cascades all wet? Differentiation and donor-control in speciose ecosystems. Ecology,
73: 747-754.
Sugiyama, H. (1997) Freshwater Fishes of Akita Prefecture.
Mumyosha Press, Akita, Japan. (in Japanese)
Suzuki, R. and T. Kato (1966) Hybridization in nature between
salmonid fishes, Salvelinus pluvius x Salvelinus fontinalis.
Bulletin of the Freshwater Fisheries Research Laboratory,
17: 83-90.
Takami, T. and T. Aoyama (1999) Distributions of rainbow
trout and brown trouts in Hokkaido, northern Japan.
Wildlife Conservation Japan, 4: 41-48. (in Japanese withEnglish Abstract)
Takami, T., T. Yoshihara, Y. Miyakoshi and R. Kuwabara(2002) Replacement of the white-spotted charr Salvelinus
leucomaenis by brown trout Salmo trutta in a branch of the
Chitose River, Hokkaido. Nippon Suisan Gakkaishi, 68:

24-28. (in Japanese with English Abstract)
Takayama, H., M. Kikuchi and I. Wakana (2002) Lakes in
Hokkaido: expansions of introduced brown trout and crayfish.
In: The ecological society of Japan, ed., Handbook of Alien
Species in Japan. Chijinshokan Press, Tokyo, pp. 254-257.
(in Japanese).
Taniguchi, Y., Y. Miyake, T. Saito, H. Urabe and S. Nakano(2000) Redd superimposition by introduced rainbow trout onnative charrs in a Japanese stream. Ichthyological Research,
47: 149-156.
Taniguchi, Y., K. D. Fausch and S. Nakano (2002) Sizestructured interactions between native and introduced
species: can intraguild predation facilitate invasion by stream
salmonids? Biological Invasion, 4: 223-233.
Taylor, J. N., W. R. Courtenay, Jr. and J. A. McCann (1984)
Known impact of exotic fishes in the continental UnitedStates. In: W. R. Courtenay, Jr. and J. R. Stauffer, eds.,
Distribution, Biology, and Management of Exotic Fish.
John Hopkins University Press, Baltimore, Maryland, pp.
322-373.
Thompson, P. D. and F. J. Rahel (1996) Evaluation of
depletion-removal electrofishing of brook trout in small
Rocky Mountain streams. North American Journal of
Fisheries Management, 16: 332-339.
Townsend, C. R. (1996) Invasion biology and ecologicalimpacts of brown trout Salmo trutta in New Zealand.
Biological Conservation, 78: 13-22.
Urabe, H. and S. Nakano (1998) Contribution of woody debris
to trout habitat modification in small streams in secondary
deciduous forest, northern Japan. Ecological Research, 13:335-345.
Wakabayashi, T., T. Nakamura, H. Kubota and T. Maruyama(2002) Comparison of spawning ecology of three salmonids
in the inlet streams of Lake Chuzenji, central Japan.
Japanese Journal of Ichthyology, 49: 133-141. (in Japanesewith English Abstract)
Wakabayashi, T., T. Nakamura, H. Kubota and T. Maruyama(2003) Comparison of microhabitats of two sympatricunderyearling salmonids in an inlet stream of Lake Chuzenji,
central Japan. Japanese Journal of Ichthyology, 50: 123-130.
(in Japanese with English Abstract)
Welcomme, R. L. (1988) International introductions of inland
aquatic species. Food and Agriculture Organization of theUnited Nations, Fisheries Technical Paper, 294: 1-318.
Rome, Italy.
Welcomme, R. L. (1992) A history of international introductions of inland aquatic species. ICES (International Council
for Exploitation of the Sea) Marine Science Symposium, 194:
3-14.
Weigel, D. E., J. T. Peterson and P. Spruell (2003) Introgressive hybridization between native cutthroat trout andintroduced rainbow trout. Ecological Applications, 13:
38-50.
Yamamoto, S., K. Morita, S. Kitano, K. Watanabe, I. Koizumi,
K. Maekawa and K. Takamura (2004) Phylogeography ofwhite-spotted charr (Salvelinus leucomaenis) inferred frommitochondrial DNA sequences. Zoological Science.
Yoshimizu, M., T. Nomura, T. Awakura, Y. Ezura and T.
Kimura (1989) Prevalence of pathogenic fish viruses inanadromous masu salmon (Oncorhynchus masou) in thenorthern part of Japan, 1976-1987. Physiology and Ecology
Japan, Special Volume 1: 559-576.
Yuma, M., K. Hosoya and Y. Nagata (1998) Distribution of the
freshwater fishes of Japan: a historical overview. Environmental Biology of Fishes, 52: 97-124.
(Received 26 January 2004, Accepted 26 April 2004)