1423

Rainbow trout (Oncorhynchus mykiss) invasion in

Hokkaido streams, northern Japan, in relation to
flow variability and biotic interactions
Mikio Inoue, Hiroshi Miyata, Yousuke Tange, and Yoshinori Taniguchi
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14

Abstract: Factors controlling invasion success of rainbow trout (Oncorhynchus mykiss) and interactions with native masu
salmon (Oncorhynchus masou) were examined by field investigations. The results suggested that flow regime has a major
role in controlling rainbow trout invasion success. Although competitive relationships were found between the two species,
masu salmon were not likely to prevent trout invasion. Analyses on the distributions and abundances of the two species in-
dicated that the occurrence of rainbow trout was limited to streams with stable flows, and the density of each species was
negatively related with that of the other. Habitat use and diet analyses revealed a pattern of their resource partitioning;
rainbow trout occupy positions near the bottom or cover, foraging benthic prey, while masu salmon feed drifting prey at
the middle layers of the water column. This resource partitioning was suggested to reflect a diet shift by rainbow trout.
Our results also have a management implication that dams, which control flow regime, can be a key factor in managing
rainbow trout invasion and its effects.
Résumé : Des observations de terrain ont permis d’examiner les facteurs qui contrôlent le succès de l’invasion de la truite
arc-en-ciel (Oncorhynchus mykiss) et ses interactions avec le saumon masou (Oncorhynchus masou) indigène. Les résultats
font croire que le régime d’écoulement des eaux a un rôle majeur dans le contrôle du succès de l’invasion des truites.
Bien qu’il existe des relations de compétition entre les deux espèces, il est peu probable que les saumons masou empê-
For personal use only.

chent l’invasion des truites. Des analyses de la répartition et de l’abondance des deux espèces indiquent que la présence
de la truite arc-en-ciel est restreinte aux cours d’eau à écoulement stable; il y a aussi une relation négative entre les den-
sités des deux espèces. Des analyses de l’utilisation de l’habitat et du régime alimentaire montrent un patron de partage
des ressources : les truites arc-en-ciel se tiennent près du fond ou des abris et se nourrissent de proies benthiques, alors
que les saumons masou s’alimentent de proies en dérive dans les couches intermédiaires de la colonne d’eau. Ce partage
des ressources semble refléter un changement de régime alimentaire chez la truite arc-en-ciel. Nos résultats ont aussi des
conséquences sur la gestion, puisque les barrages, qui contrôlent le régime d’écoulement, peuvent être un facteur essentiel
pour gérer l’invasion des truites arc-en-ciel et ses effets.
[Traduit par la Rédaction]

Introduction affecting success and failure of invasion and variation in im-

Invasions by non-native species are a major threat to bio-
diversity and ecosystem health. Once non-native species es-
tablish self-reproducing populations, they can exert strong
impacts on native communities at various levels through
competition, predation, and indirect cascade effects (Crowl
et al. 1992; Townsend 2003). However, introduced species
cannot always become successful invaders; some species
fail to be established (e.g., Brown and Moyle 1997) or may
not have strong, detectable impacts on native communities
even if they are established. Therefore, understanding factors
Received 11 September 2008. Accepted 22 June 2009. Published
on the NRC Research Press Web site at cjfas.nrc.ca on
20 August 2009.
J20770
M. Inoue,1 H. Miyata, and Y. Tange. Department of Biology,
Faculty of Science, Ehime University, Bunkyo-cho 2-5,
Matsuyama 790-8577, Japan.
Y. Taniguchi. Department of Environmental Science and
Technology, Faculty of Science and Technology, Meijo
University, Tenpaku, Nagoya 468-8502, Japan.
1Corresponding author (e-mail: inom@sci.ehime-u.ac.jp).
pacts by invaders is an important subject in invasion biology
(Fausch et al. 2001; Light 2003; Marchetti et al. 2004).
Rainbow trout (Oncorhynchus mykiss), which is native to
the Pacific coast of North America and Kamchatka Peninsula
(Russian Far East), is one of the most widely introduced spe-
cies in cool-temperate regions of the world (Fausch et al.
2001), with evidence of its negative impacts on native biota
having been frequently reported (e.g., Crowl et al. 1992;
Weigel et al. 2003; Morita et al. 2004). Fausch et al. (2001)
analyzed geographic patterns in invasion success of rainbow
trout at a broad scale, using four regions where their inva-
sions varied from highly successful (southern Appalachians,
USA) to moderate (Colorado, USA, and Hokkaido, Japan) or
failed (Honshu, Japan). They showed that the region where
invasions failed has flow regimes that are not favorable for
reproductive life history of rainbow trout (e.g., frequent high
flows after their emergence period). The match between in-
vasion success and flow regime across the regions strongly
suggests that flow disturbance limits the trout’s invasion suc-
cess. However, because their comparison was made at a
large, regional scale, various factors may differ distinctly
among the regions (e.g., temperature regimes, biota), and
the differences in invasion success might be explained also

Can. J. Fish. Aquat. Sci. 66: 1423–1434 (2009) doi:10.1139/F09-088 Published by NRC Research Press
 1424
by similar other factors. Therefore, their flow disturbance
hypothesis needs to be examined at a smaller spatial scale
(i.e., within a region) so that possible confounding factors
that vary among regions can be ruled out. If presence or ab-
sence of rainbow trout among streams within a region can be
explained by differences in flow regimes, this provides
strong support for the flow disturbance hypothesis.
In Japan, rainbow trout has been stocked extensively over
the four main islands (Hokkaido, Honshu, Shikoku, and
Kyushu), with varying success of establishment among the
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14
regions (Kitano 2004). In Honshu, Shikoku, and Kyushu,
only a few cases of their establishment have been reported
(Kato and Yanagawa 2000), despite repeated stocking for
fisheries and recreational purposes in most river systems
through the last several decades (Kawanabe 1980; Kitano
2004). In Hokkaido, however, their self-sustaining popula-
tions are not rare (e.g., Kitano et al. 1993; Aoyama et al.
1999; Takami and Aoyama 1999); they have been success-
fully established in certain streams, but not in others. This
situation is ideal to test the flow disturbance hypothesis.
The objective of our study is to identify factors controlling
invasion success of rainbow trout in Hokkaido streams, con-
sidering the flow disturbance hypothesis and biotic interac-
tions with native salmonids.
It is important to examine interactions between native and
non-native salmonids, not only in terms of impacts of in-
For personal use only.
vaders but also of biotic resistance from native species. Be-
cause competition for food and space is common among
salmonids (Fausch 1988; Nakano 1995), native salmonids
may prevent invasion by non-native salmonids through com-
petitive interactions. On the Asian side of the Pacific coast,
masu salmon (Oncorhynchus masou) and whitespotted charr
(Salvelinus leucomaenis) are the two most widely distributed
native salmonids (Kawanabe 1989). Recently, a growing
number of studies, both empirical (Taniguchi et al. 2000;
Morita et al. 2004) and experimental (Hasegawa et al. 2004;
Hasegawa and Maekawa 2006), have revealed negative ef-
fects of non-native rainbow trout on whitespotted charr pop-
ulations in Japan. These studies suggest that whitespotted
charr is unable to prevent rainbow trout invasion. However,
studies on interactions with masu salmon are scarce and lim-
ited to small-scale, controlled experiments in artificial chan-
nels (Taniguchi et al. 2002; Hasegawa et al. 2004; Hasegawa
and Maekawa 2006); empirical data from the field are still
largely lacking (but see Nakano and Kaeriyama 1995).
Nakano and Kaeriyama (1995) described resource parti-
tioning among four salmonid species, including rainbow
trout (juvenile steelhead (i.e., sea-run rainbow trout)) and
masu salmon in a Kamchatkan stream, which provided a
unique case where both species were native. Their data sug-
gest that masu salmon occupied suitable feeding positions
despite the presence of rainbow trout, although their study
did not adopt a comparative approach in which competitive
effects can be examined (i.e., comparison of resource use
between sympatric and allopatric sites; e.g., Gatz et al.
1987). Recent experimental studies using laboratory chan-
nels reported that masu salmon matched rainbow trout in be-
havioral competitive ability (Hasegawa et al. 2004), and
rainbow trout did not affect habitat use of masu salmon (Ha-
segawa and Maekawa 2006). Such evidence suggests that ef-
fects of rainbow trout on masu salmon populations may be
Can. J. Fish. Aquat. Sci. Vol. 66, 2009
Fig. 1. Location of the study sites in the Iburi District, southwestern
Hokkaido, northern Japan. Fish distribution and abundance were
examined using 24 sites (solid circles and open squares); open
squares indicate six sites used for habitat use and diet analyses.
weak (but see Taniguchi et al. 2002). Rather, masu salmon
may offer biotic resistance to rainbow trout invasions.
In this study, we examined (i) distribution and abundance
of rainbow trout in relation to flow variability, masu salmon
abundance, and other habitat factors, and (ii) habitat use and
diet of rainbow trout and masu salmon in both sympatric
and allopatric sites. Because we assume that life history of
rainbow trout does not perfectly fit to the seasonal pattern
of stream discharge in Hokkaido (high flows in spring and
summer; see Fausch et al. 2001), we predict that rainbow
trout would fail to establish in streams with higher flow var-
iability, and therefore, the occurrence of rainbow trout
would be limited to streams with stable flows. If masu sal-
mon offers strong biotic resistance to rainbow trout invasion,
we predict that a complementary occurrence would be found
in their distributions. In addition, investigations on habitat
use and diet of the two species would provide information
on elaborating their interactions.
Materials and methods
Study area
The study was conducted using 13 streams in the Iburi
District in southwestern Hokkaido, Japan (Fig. 1). The mean
annual precipitation in this region is ~1200 mm, dominated
by summer rainfall (70%) and winter snowfall (25%). In
streams, high flows occur in the snowmelt season (April–
May) and between August and October by rainfall (Kishi et
al. 1999; Shibata et al. 2001). Although precipitation regime
and the seasonal pattern of stream discharge are similar
within the study area, the degree of fluctuation in stream dis-
charge differs between eastern and western parts of the area
owing to geological conditions. In the eastern side, the soil is
dominated by clastic pumice derived from volcanic ejecta of
Mt. Tarumae. Because this soil has high water permeability,
surface runoff during precipitation is negligible, and stream
discharge is derived mainly from deeper groundwater
(Shibata et al. 2001). As a result, streams on this volcanic
Published by NRC Research Press
 Inoue et al.
deposit have stable flow. Distribution of the thick deposits of
the Mt. Tarumae volcanic ejecta is limited to the eastern side
of Tarumae Stream, and thus flow fluctuation in the western
side streams is higher than in the eastern side.
Twenty-four study sites were established over the 13
streams to investigate fish distribution, abundance, habitat
use, and diet (Fig. 1). Streams at the study sites were gener-
ally small (first- to third-order), with low or moderate chan-
nel gradient (Table 1). All study sites were located in
deciduous broad-leaved forests dominated by oak (Quercus
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crispula) and maple (Acer spp.), and stream channels were
generally shaded by riparian canopy. Masu salmon or rain-
bow trout and Siberian stone loach (Barbatula barbatulus)
or sculpin (Cottus nozawae) were common among the study
sites. Two other salmonids, whitespotted charr and exotic
brown trout (Salmo trutta), were found at a few sites.
In Hokkaido, rainbow trout are strictly stream residents
that spawn during spring (late January – early May: Aoyama
et al. 1999; Taniguchi et al. 2000), whereas most masu sal-
mon are anadromous. Masu salmon adults ascend rivers and
spawn in small tributaries in autumn (late August – early
October). Fry emerge in spring (February–May), earlier than
rainbow trout (June–July). Juvenile masu salmon generally
spend 1 year as age-0 parr and descend to the sea in the fol-
lowing spring (i.e., most masu salmon in streams are age-0),
although some individuals stay in streams more than 1 year.
For personal use only.
Owing to these differences in life history traits, size structure
differs between the two species in streams (see Taniguchi et
al. 2002). Masu salmon (mostly age-0) are larger than age-0
rainbow trout but smaller than age-1 and older rainbow trout.
Distribution and abundance
Field surveys
A study reach including at least two sets of pool–riffle se-
quences (reach length: 50–150 m) was established at each
site to quantify fish abundance and habitat characteristics.
Fish abundance was estimated during June–July 2003 using
the removal method of DeLury (1951). Each study reach
was blocked with 5 mm mesh seines at the upper and lower
ends, and three removal passes were made using an electrof-
ishing unit (Model 12 Backpack Electrofisher, Smith-Root
Inc.). The fork length of captured fish was measured. Popu-
lation estimates of rainbow trout and masu salmon were cal-
culated separately for fry (age-0) and parr (age > 0) and
converted to density (N
100 m–2).
Summer water temperature, channel gradient, stream size,
and the area of habitat types (pool and cover) were quanti-
fied at each site during June–August 2003. A maximum–
minimum thermometer was placed in each study reach, and
the maximum temperature was recorded every 10–14 days
(four times) from 18 June to 6 August 2003. The channel
gradient, expressed as percent change in relative height to
reach length, was measured with a level and leveling rod.
Equally spaced transects were set along the study reaches
(3–5 m intervals depending on stream size), and bank-full
and wetted channel widths were measured. The boundaries
of bank-full channels were determined by scour lines, vege-
tation limits, or changes between developed organic soils
and inorganic alluvium. Each study reach was divided into
channel units, classified as riffle, glide, or pool. The length
1425
Table 1. Habitat variables characterizing each study site.
Habitat variable Mean (SD), range
Channel gradient (%) 1.0 (0.6), 0.2–2.1
Stream size (m)
Mean wetted width 4.1 (1.3), 1.5–7.1
Water temperature (8C)
Maximum in summer 13.5 (2.1), 9.3–18.5
Habitat component (%)
Pool area 45.4 (15.9), 15.7–80.7
Cover area 9.7 (6.5), 1.9–29.3
Flow variability index
Wetted/bank-full width ratio 0.74 (0.22), 0.24–1.00
of each channel unit was measured, and each channel unit
area was estimated by multiplying the length and the mean
wetted width of the transects within the channel unit. Cover
is generally considered as any object that conceals fish from
predators or provides shelter from strong current (e.g., Shir-
vell 1990). In this study, we considered coarse woody de-
bris, brush (often structured by twigs and grass stems),
undercut banks, and overhanging vegetation (instream or
within 40 cm of the water surface) as cover materials. The
length and several widths of each cover material were meas-
ured to calculate the occupied area within the wetted chan-
nel surface.
Habitat variables
Habitat characteristics of the study reaches were ex-
pressed by summer maximum water temperature (mean of
the four maximum temperatures recorded), channel gradient,
mean wetted width, the areal percentage of pools and that of
cover (Table 1), which can affect the abundance of rainbow
trout (Urabe and Nakano 1998, 1999) and masu salmon
(Inoue et al. 1997; Inoue and Nakano 1998). In addition to
these five variables, wetted/bank-full width ratio (W/B ratio),
calculated as the ratio of mean wetted width at low flow to
mean bank-full channel width, was used as an index of flow
variability. Bank-full channel is the region of wetted area at
high flows occurring once every few years (Gordon et al.
2004). Therefore, lower values of W/B ratio indicate greater
differences between peak and baseflow discharges, suggest-
ing higher flow variability. At locations near 9 of the 24
study sites, seasonal variations in discharge had been re-
corded (Hayakawa and Sato 1978). Our W/B ratio was neg-
atively correlated with coefficient of variation (SD/mean) of
the discharges (r = –0.79, P = 0.011, n = 9), suggesting that
this measure can be used as an index of flow variability in
this region.
Data analyses
Distribution and abundance of fishes were analyzed using
discriminant and regression analyses. First, a stepwise dis-
criminant function analysis was used to examine factors af-
fecting presence or absence of rainbow trout. The above
habitat variables and masu salmon density were used to dis-
criminate presence or absence of rainbow trout at the study
sites. The correct classification rate by the discriminant
model was estimated by a leave-one-out cross-validation
method (jackknife method in Olden et al. 2002). Second, ef-
fects of habitat and biotic interactions (rainbow trout vs.
Published by NRC Research Press
 1426
Fig. 2. Differences in habitat conditions (water depth and current velocity: a, 2003; c, 2004) and food availability (b, 2003; d, 2004) among
the study sites: rainbow trout allopatric sites (solid squares: TM, Tomakomai; UN, Uenae), masu salmon allopatric sites (open circles:
BB, Buhbetsu; BT, Betsubetsu), and sympatric sites (crosses: HN, Horonai; KI, Koitoi). Bars indicate ± standard deviation (SD).
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14
For personal use only.
masu salmon) on rainbow trout abundance were examined
by a stepwise multiple regression analysis, using study sites
inhabited by rainbow trout. Rainbow trout density was used
as a dependent variable and the habitat variables and masu
salmon density as independent variables. Similarly, effects
of rainbow trout on masu salmon abundance were examined
using rainbow trout density with the habitat variables as in-
dependent variables; rainbow trout density was expressed by
not only total density but also density of age-1 and older
trout (removing age-0 trout from total density), because
only larger trout may have significant effects on masu
salmon (Taniguchi et al. 2002). In addition, effects of habitat
variables on total salmonid density (rainbow trout and masu
salmon combined) were examined. Fish densities, channel
gradient, and mean wetted width were log10-transformed to
improve normality; the percentages of pool and cover and
W/B ratio were arcsin-square-root-transformed. All analyses
were performed using SPSS (version 13, SPSS Inc.,
Chicago, Illinois).
Habitat use and diet
Field observation and sampling
Habitat use and diet of rainbow trout and masu salmon were
surveyed using six study sites in autumn (September–October
2003) and early summer (June–July 2004) (habitat use survey
in autumn was conducted at only three sites) to examine their
interspecific interactions (Fig. 1). The six sites represented the
three cases of allopatric–sympatric situations (two sites per
case): sites inhabited only by rainbow trout (trout allopatric
site: Tomakomai, TM; Uenae, UN), only by masu salmon
(salmon allopatric site: Buhbetsu, BB; Betsubetsu, BT), and
by both species (sympatric site: Koitoi, KI; Horonai, HN).
Can. J. Fish. Aquat. Sci. Vol. 66, 2009
Some distinct differences were found among the study sites in
both habitat characteristics and food abundance. HN and BT
had slower current and deeper water, respectively, than other
sites (Fig. 2). HN was characterized also by abundant food.
Habitat use was surveyed by underwater observation.
Underwater observations were made by snorkeling in an up-
stream direction, with the focal points of fish being identi-
fied for all undisturbed individuals (rainbow trout or masu
salmon) encountered. Distance from the bottom to the focal
points (vertical position) and the fish total length were esti-
mated visually, and the focal points were marked by putting
a numbered flag on the streambed. In each site, the observa-
tion was made in a reach including at least two pool–riffle
sequences; in case the reach did not harbor sufficient num-
ber of fish, the observation reach was expanded so that data
of at least 30 individuals per species were obtained (length
of observation reaches: 80–500 m). After the observation,
water depth, current velocity at the surface, mid-depth (0.5
depth), bottom and fish vertical position, and the distance
from the nearest cover were measured at the marked focal
points. Focal points were then characterized by water depth,
focal point velocity, distance from cover, and the relative
vertical position (distance from the bottom/total depth). To
describe differences in habitat characteristics among the
study sites, water depth and current velocity were measured
over the observed reach using transects. Equally spaced
transects (3–6 m) with equally spaced measuring points
(0.3–0.5 m) were established, and water depth and the sur-
face, mid-depth, and bottom velocities were measured.
Fish were sampled using an electrofishing unit for analy-
sis of their diet. Captured fish were anesthetized with FA100
(Tamura-Seiyaku Pharmaceutical), and stomach contents
were sampled using a stomach pumping method. Fish were
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 Inoue et al. 1427

Table 2. Summary of stomach contents of rainbow trout and masu salmon in

allopatry and sympatry.

Rainbow trout Masu salmon

Prey category Allopatry Sympatry Allopatry Sympatry
Terrestrial invertebrates 63.0–78.0 7.6–73.7 41.8–85.4 62.1–89.1
Aquatic invertebrates
Gammaridae 1.3–27.3 0.4–87.3 0.–0.1 0.2–2.3
Other invertebrates 9.7–29.8 5.1–26.0 14.6–38.0 9.1–37.7
Others 0 0.–4.6a 0.–41.7b 0
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Note: Relative abundance (%) of each category by dry weight is expressed as the
range of four values from each site (two sites by two seasons).
a
Fish (rainbow trout fry) in early summer.
b
Eggs (masu salmon) in autumn.

then released alive. Benthic and drifting invertebrates were Fig. 3. The presence (solid squares) and absence (open circles) of
also sampled to examine differences in food availability rainbow trout in relation to wetted/bank-full width ratio and masu
among the study sites. Six samples of benthic invertebrates salmon density. The axes of masu salmon density and wetted/bank-
were sampled from riffles and pools (three samples per hab- full width ratio are logarithmic and arcsin-square-root scales, re-
itat type) in each site using a Surber sampler (25 cm  spectively.
25 cm quadrat, 0.3 mm mesh). Drift was sampled once at
each site using drift nets (25 cm  25 cm opening, 0.3 mm
mesh). Four drift nets were placed in a riffle for 30–40 min
during the daytime (1000–1500 h), with current velocity
measured at the center of net opening to estimate the vol-
ume of sieved water. Samples of stomach contents, benthic
For personal use only.

invertebrates, and drifts were preserved in 10% formalin in

the field and sorted (benthic and drift samples were washed
onto 1 mm mesh), identified to family or order, and weighed
(dry weight) in the laboratory.

Data analyses
Habitat and diet data were used to examine resource par-
titioning between rainbow trout and masu salmon and ef-
fects of interspecific interactions on it. To identify habitat
variables related to their habitat partitioning in the sympatric
sites, a stepwise discriminant function analysis was used.
Focal points of rainbow trout and those of masu salmon in
each sympatric site were discriminated using water depth,
focal point velocity, distance from cover, and the relative
vertical position as potential discriminant variables, with the
correct classification rate being assessed by a leave-one-out
cross-validation. On the basis of the results of this analysis,
key variables related to their habitat partitioning were iden-
tified. These variables were then compared among allopatric
rainbow trout and allopatric masu salmon using one-way
analysis of variance (ANOVA) with Tukey’s honestly sig-
nificant difference (HSD) multiple comparison test to exam-
ine whether their habitat partitioning in the sympatric sites
were mediated by habitat shift by either species. Water
depth, focal point velocity, and distance from cover were
log10- transformed to improve normality; the relative vertical
position was arcsin-square-root-transformed.
In the diet analysis, we focused on two prey categories,
terrestrial invertebrates and gammarids, which were major
components of stomach contents (Table 2) and may reflect
foraging behavior of fish. For stream fishes, terrestrial inver-
tebrates are provided as drifts, and thus their dominance in
stomach contents is indicative of drift feeding. On the other
hand, gammarids, which also made up large proportions in
some sites (Table 2), are a typical inhabitant of leaf litter
patches on the bottom (see Konishi et al. 2001). Therefore,
gammarids are likely consumed by benthic feeding rather
than drift feeding. To examine whether the food resource
use differs between rainbow trout and masu salmon in each
sympatric site, the percentage of terrestrial invertebrates and
that of gammarids in stomach contents (by dry weight) of
individual fish were compared between the two species us-
ing Mann–Whitney U test. Similarly, the percentage of ter-
restrial invertebrates and that of gammarids were compared
among allopatric rainbow trout and allopatric masu salmon
using Kruskal–Wallis test and subsequent pairwise compari-
sons (Mann–Whitney U test with Bonferroni correction). All
analyses were performed using SPSS (version 13, SPSS Inc.,
Chicago, Illinois).
Results
Distribution and abundance
Among the 24 study sites, rainbow trout occurred in 11
sites. The stepwise discriminant function analysis indicated
that the presence or absence of rainbow trout was discrimi-
nated by the W/B ratio but not by masu salmon abundance
(Fig. 3); the discriminant function, of which correct classifi-
cation rate was 91.7%, retained only W/B ratio as the dis-
criminant variable (Wilks’ l = 0.282, c2 = 27.19, P <
0.001). The study sites inhabited by rainbow trout were
characterized by a higher value of W/B ratio.

Published by NRC Research Press

 1428 Can. J. Fish. Aquat. Sci. Vol. 66, 2009

Table 3. Results of stepwise multiple regression analysis on the densities of rainbow trout,
masu salmon, and total salmonids (both species combined).

Significance
Standardized
Species Independent variable coefficient R2 F P
Rainbow trout Mean wetted width –0.64 0.75 11.72 0.004
Masu salmon density –0.45
Masu salmon Age-1 and older trout density –0.79 0.62 14.85 0.004
Total salmonids Cover area 0.67 0.45 7.38 0.024
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Note: Data were transformed prior to analysis.

The stepwise multiple regression analysis on the 11 sites Fig. 4. The density of rainbow trout (a, b), masu salmon (c), and
revealed negative relationships between rainbow trout and both combined (d) in relation to explanatory variables detected by
masu salmon densities (Table 3; Fig. 4). The variation of multiple regression analysis. The axes of fish densities and mean
rainbow trout density was best explained by a combination wetted width are logarithmic scale, while that of cover area is arc-
of mean wetted width and masu salmon density; rainbow sin-square-root scale. In panel (b), trout and salmon densities are
trout increased with decreasing wetted width and masu sal- expressed by residuals after removing effects of mean wetted width.
mon density. Masu salmon density was best modeled by a
negative effect of the density of age-1 and older rainbow
trout. For total salmonids, cover area was detected as the
single best predictor. The density of rainbow trout and
masu salmon combined increased with cover area.

Habitat use
In the sympatric sites, differences in the total body length
For personal use only.

between rainbow trout and masu salmon varied among the

sites and seasons (Fig. 5). In HN, rainbow trout was smaller
than masu salmon in autumn 2003, whereas rainbow trout
was larger than masu salmon in early summer 2004. In KI
in early summer, the sizes of the two species were compara-
ble, although the shape of frequency distribution differed
(bimodal vs. unimodal). The stepwise discriminant function
analysis showed that although variables discriminating be-
tween focal points of rainbow trout and those of masu sal-
mon in sympatry varied among the sites and seasons, focal
points of rainbow trout were generally characterized as
lower vertical positions or locations closer to cover (Table 4;
Fig. 6). In HN in autumn, focal points of the two species
were best discriminated by relative vertical position, with
rainbow trout occupying lower vertical position than masu
salmon. In HN in early summer, where rainbow trout was
larger in body size than masu salmon, their focal points
were best discriminated by water depth and distance from
cover. Rainbow trout occupied locations with deeper water
and closer to cover than masu salmon. In KI, where their
sizes were comparable, the discriminant function retained
all four variables, with the highest contribution of relative
vertical position followed by distance from cover; their hab-
itat partitioning in terms of the two variables was similar to
those in HN.
Relative vertical position and distance from cover were
considered as key variables in habitat partitioning between
the two species, and these variables were compared among
the allopatric sites (Fig. 7). In autumn 2003, relative vertical
position of allopatric rainbow trout (UN) was significantly
lower than that of allopatric masu salmon (BB) (one-way
ANOVA, F = 4.80, P < 0.032). Although the same result
was obtained from early summer 2004 (i.e., trout in UN <
salmon in BB; Tukeys’ HSD test, P = 0.021), relative verti-
cal position of rainbow trout in another allopatric site (TM)

Published by NRC Research Press

 Inoue et al. 1429

Fig. 5. Frequency distributions of body size for rainbow trout (solid Table 4. Results of stepwise discriminant function analysis dis-
columns) and masu salmon (open columns) in each sympatric site: criminating focal points of rainbow trout and those of masu sal-
HN (Horonai) in autumn 2003 (a), HN in early summer 2004 (b), mon in each sympatric site.
and KI (Koitoi) in early summer 2004 (c).
Autumn
2003 Early summer 2004
Variable–statistic HN HN KI
Variable
Water depth — –0.95 0.41
Focal-point velocity — — 0.40
Distance from cover — 0.81 0.56
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14

Relative vertical position 1.00 — 0.99

Statistics
Wilks’ l 0.590 0.513 0.703
c2 31.96 60.06 35.24
P <0.001 <0.001 <0.001
Correct classification rate 87.3 80.6 77.9
(%)
Note: Standardized coefficients, significance, and correct classification
rate based on a leave-one-out cross-validation are shown for each discri-
minant function. Data were transformed prior to analysis. HN, Horonai;
KI, Koitoi.

vertebrates, while many individuals of masu salmon had

terrestrial prey in their stomachs. In particular, in autumn
2003, more than 40% of masu salmon stomachs were
For personal use only.

filled with terrestrial prey by 90%–100%. In contrast with

was not significantly different from that of allopatric masu
salmon in BB and BT (Tukey’s HSD test, P = 0.198–
0.999). Distance from cover was not significantly different
between the two species in autumn 2003 (one-way AN-
OVA, F = 0.265, P = 0.609). However, in early summer
2004, distance from cover for allopatric rainbow trout in
both TM and UN was significantly shorter than that for al-
lopatric masu salmon in BB and BT (Tukey’s HSD test, P <
0.03).
Diet
The diet analysis indicated that in the sympatric sites,
stomach contents differed between rainbow trout and
masu salmon (Fig. 8). The percentage of terrestrial prey
in stomach contents of rainbow trout was significantly
lower than that of masu salmon in both sites in both sea-
sons (Mann–Whitney U test, see Fig. 8). From 20% to
50% of rainbow trout stomachs included no terrestrial in-
the case of terrestrial prey, the percentage of gammarids
in stomach contents was significantly higher for rainbow
trout than for masu salmon in all cases except KI in au-
tumn 2003 (Fig. 8). Most masu salmon had no gammarids
in their stomachs, whereas stomach contents of several in-
dividuals of rainbow trout were dominated by gammarids,
especially in HN.
In the allopatric sites (Fig. 9), contribution of terrestrial
prey to diet of rainbow trout was greater than that of
sympatric rainbow trout. In autumn 2003, the percentage
of terrestrial prey in rainbow trout stomachs in both TM
and UN was larger than that of masu salmon in BB
(Mann–Whtney U test, P = 0.003, 0.006, respectively;
see Fig. 9) and not significantly different from that of
masu salmon in BT (Mann–Whtney U test, P = 0.155,
0.467, respectively). This is attributable to lower contribu-
tion of terrestrial prey to masu salmon diet, as well as
higher contribution of terrestrial prey to rainbow trout
diet. In autumn 2003, the study sites in BB and BT were
used for spawning by adult masu salmon, and stomachs
of several masu salmon sampled were filled with salmon
eggs (see Table 2), which is a reason for the low contri-
butions of terrestrial prey to masu salmon diet in these
sites.
In early summer 2004, no significant difference was
found in the percentage of terrestrial prey among the species
and sites (Kruskal–Wallis test, P = 0.398). In every case of
rainbow trout and masu salmon under allopatric situations,
more than 30% of stomachs were filled with terrestrial prey
by >80%. Differences in gammarid contribution between the
two species in allopatric sites were similar to those in sym-
patric sites (Fig. 9). In both seasons, the percentage of gam-
marids in stomach contents of rainbow trout was higher than
that of masu salmon (Mann–Whtney U test, P < 0.005).

Published by NRC Research Press

 1430
Fig. 6. Differences in focal point characteristics between rainbow trout (solid squares) and masu salmon (open circles) in Horonai in autumn
2003 (a), Horonai in early summer 2004 (b), and Koitoi in early summer 2004 (c), based on the results of discriminant function analysis.
The axes of distance from cover and water depth are logarithmic scale, while that of relative vertical position is arcsin-square-root scale.
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14
Fig. 7. Comparisons of relative vertical position (a) and distance
from cover (b) for focal points held by rainbow trout (RT, solid
squares) and masu salmon (MS, open circles) among allopatric sites
(TM, Tomakomai; UN, Uenae; BB, Buhbetsu; BT, Betsubetsu).
Bars indicate ± standard deviation (SD). Data denoted by the same
letter (a, b; x, y, z) are not significantly different (P > 0.05) by one-
way analysis of variance (ANOVA) or Tukey’s honestly significant
difference (HSD) test.
For personal use only.
Discussion
Factors controlling invasion success
In our study area, the sites with and without rainbow trout
were clearly separated by W/B ratio, with sites inhabited by
rainbow trout having higher values. This result suggests that
stable discharge favors establishment of rainbow trout popu-
lations, supporting the flow disturbance hypothesis by
Fausch et al. (2001). Life history of rainbow trout is thought
to adapt to flow regimes in their native range (Fausch et al.
2001). For example, in Pacific Coast of North America,
where rainbow trout is native, flow regimes of streams are
characterized by winter high flows and summer low flows
Can. J. Fish. Aquat. Sci. Vol. 66, 2009
(see Fausch et al. 2001; Light 2003), and spawning and
emergence season of rainbow trout is from late winter to
early summer, the period of declining flow (Fausch et al.
2001). This match decreases the risk of mortality caused by
high flows at egg and fry stages. On the other hand, in our
study area in Hokkaido, northern Japan, high flows occur in
April–May by snowmelt and in August–October by rainfall
(Kishi et al. 1999; Shibata et al. 2001). Therefore, high flows
by snowmelt, coincident with the spawning and emergence
period of rainbow trout in this area (Kitano et al. 1993; Tani-
guchi et al. 2000), have potential to limit establishment of
their populations. In the eastern side of the study area, how-
ever, owing to the high water permeability of the soil, stream
flow variability is low, which was expressed by a higher W/B
ratio. For example, Shibata et al. (2001) reported the lack of
snowmelt peak in the hydrograph of Horonai Stream, a typi-
cal spring-fed stream in the eastern side. Probably, such a
moderate hydrological condition in the spawning and emer-
gence period is an important factor allowing rainbow trout
to establish their self-reproducing populations.
Presence or absence of rainbow trout was not related to
masu salmon density, suggesting that biotic resistance from
the native salmonid plays no role or a lesser role in control-
ling rainbow trout invasion. In general, where two species of
stream salmonids exhibit complementary distributions (oc-
currences or abundances), interference competition often
plays an important role (Fausch et al. 1994; Taniguchi and
Nakano 2000; Rieman et al. 2006). Superiority in interfer-
ence competition is largely determined by body size (e.g.,
Nakano 1995). In the case of competition between rainbow
trout and masu salmon, the latter has a clear advantage over
the former, because masu salmon fry emerge earlier than
rainbow trout. This advantage allows masu salmon to have
a great potential to prevent rainbow trout invasion. How-
ever, most masu salmon in Hokkaido are anadromous, stay-
ing in streams generally for only 1 year. Therefore, rainbow
trout that survive 1 year can grow larger than masu salmon,
resulting in a competitive advantage over masu salmon. An
experimental study by Taniguchi et al. (2002) demonstrated
that growth and survival of age-0 masu salmon were re-
duced by age-1 rainbow trout through aggressive interfer-
ence and predation. Anadromy of masu salmon may be one
reason why they cannot offer biotic resistance to rainbow
trout invasion in our study area.
Published by NRC Research Press
 Inoue et al. 1431

Fig. 8. Relative abundance of terrestrial prey and gammarids in the diet of rainbow trout (solid columns) and masu salmon (open columns)
individuals in each sympatric site (HN, Horonai; KI, Koitoi). Results (P value) of Mann–Whitney U test comparing between the two species
in each site are indicated in parentheses. An asterisk indicates statistically significant (P < 0.05).
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14

Fig. 9. Relative abundance of terrestrial prey and gammarids in the diet of rainbow trout (solid columns) and masu salmon (open columns)
individuals in each allopatric site (TM, Tomakomai; UN, Uenae; BB, Buhbetsu; BT, Betsubetsu). Significant differences among the four
sites based on Mann–Whitney U test after Kruskal–Wallis test are denoted by letters (a, b; x, y); sites denoted by the same letter are not
ignificantly different at P = 0.0083 (= 0.05/6; Bonferroni correction).
For personal use only.

Importance of abiotic factors, rather than biotic interac-
tions, in controlling invasion and its impacts has frequently
been reported in other regions. For example, in New
Zealand and Australia, both drought and spate are suggested
to limit invasion of brown trout (Salmo trutta) and prevent
them from eliminating native galaxiid fishes (Closs and
Lake 1996; McIntosh 2000; Leprieur et al. 2006). In
California, 5-year annual surveys of fishes in a regulated
stream revealed that the abundance of non-native fishes de-
creased with increasing stream flow, suggesting flushing ef-
fects on non-native fishes (Marchetti and Moyle 2001). Such
effects of high flows were reported also for non-native
crayfish in other streams of California (Light 2003). Our re-
sults are consistent with these previous studies and the view
that flow regime has a major role in controlling invasion
success.
Interactions with native masu salmon
In our study area, masu salmon were distributed regard-
less of the presence of rainbow trout. This lack of comple-
mentary occurrences between the two species indicates that
competitive exclusion is not likely to have occurred. How-
ever, the regression analysis on the abundance variations
showed significant negative effects of each density on the
other’s. Previous studies have shown that masu salmon
abundance is limited by the abundance of cover or pool in
northern Hokkaido streams where other salmonids were rare
(Inoue et al. 1997; Abe and Nakamura 1999; Abe 2007).

Published by NRC Research Press

 1432
Similarly for rainbow trout, importance of pool and cover in
determining their abundance has been shown in southern
Hokkaido streams, where rainbow trout numerically domi-
nated (Urabe and Nakano 1998, 1999). In the present study,
however, the abundance variation of each species was ex-
plained by the density of the opponent, rather than such hab-
itat factors, although the density of the two species
combined was explained by cover abundance. These results
suggest that rainbow trout and masu salmon have a compet-
itive relationship with each other and share the same carry-
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14
ing capacity that increases with cover abundance. In other
words, establishment of rainbow trout can decrease masu
salmon abundance by forcing to share the capacity, although
masu salmon is not likely inferior to rainbow trout in com-
petitive interactions (see discussion below).
The habitat use and diet analyses in the sympatric sites
describe resource partitioning between the two species. Ma-
jor variables related to their habitat partitioning were verti-
cal position and proximity to cover. Rainbow trout
generally occupied positions closer to the bottom or cover
than masu salmon. Further, the diet analysis showed that
contribution of terrestrial prey to the diet of rainbow trout
was lower than that of masu salmon. Instead, rainbow trout
tended to consume gammarids. These results illustrate that
in the sympatric sites, many individuals of rainbow trout oc-
cupied positions near the bottom or cover, foraging benthic
For personal use only.
prey, while the majority of masu salmon fed drifting prey at
the middle layers of the water column. Although informa-
tion on their interactions is limited, the pattern of their re-
source partitioning described here is largely consistent with
existing data from a field study (Nakano and Kaeriyama
1995) and an experimental work (Hasegawa and Maekawa
2006). Nakano and Kaeriyama (1995) described distinct dif-
ferences in diet between rainbow trout (juvenile steelhead)
and masu salmon, with the former characterized by aquatic
invertebrates of large body size (caddisfly larvae) and the
latter by terrestrial invertebrates. They also reported higher
selectivity for cover habitats by rainbow trout than by masu
salmon. Furthermore, an experiment using an artificial chan-
nel showed that rainbow trout occupied lower vertical posi-
tions relative to masu salmon when they were in sympatry
(Hasegawa and Maekawa 2006).
Although we also examined resource use by the two spe-
cies in allopatry, it is difficult to clarify whether the habitat
partitioning in sympatry is mediated by shift by either spe-
cies. Results of interspecific comparisons of habitat use
among the allopatric sites were equivocal, possibly owing to
among-site differences in habitat conditions and food avail-
ability (Fig. 2). For example, in autumn 2003, vertical posi-
tion of allopatric trout (in UN) was lower than that of
alloptaric salmon (in BB), suggesting that the vertical segre-
gation in the sympatric site is due to species-specific habitat
preference, rather than habitat shift by either species,
whereas the data from 2004 showed that vertical position of
allopatric rainbow trout in TM was as high as that of masu
salmon in BB and BT. Despite such equivocality, our data
suggest, at least, that masu salmon is unlikely to have shifted
their resource use under the influence of rainbow trout. The
pattern of resource partitioning we described is analogous to
that between masu salmon and whitespotted charr (Nakano
1995; Miyasaka et al. 2003), where masu salmon are com-
Can. J. Fish. Aquat. Sci. Vol. 66, 2009
petitively dominant and do not shift their resource use
(masu salmon occupy higher vertical positions than charr,
feeding on drifting terrestrial prey). That is, higher vertical
position is suitable for masu salmon. Therefore, the vertical
segregation between masu salmon and rainbow trout in our
sympatric sites suggests that masu salmon held their pre-
ferred position. This is supported by previous experiments
showing that competitive ability of masu salmon in interfer-
ence interactions was comparable to that of rainbow trout
(Hasegawa et al. 2004), and rainbow trout did not elicit hab-
itat shift by masu salmon (Hasegawa and Maekawa 2006).
In the experiment by Hasegawa and Maekawa (2006),
vertical position of allopatric rainbow trout was lower than
that of allopatric masu salmon, and they concluded that their
vertical segregation in sympatry simply reflects a difference
in species-specific habitat preference by each species. In ad-
dition, selective use of cover has often been reported for
rainbow trout in allopatric situation (Gatz et al. 1987;
Fausch 1993) and in streams where effects of other salmo-
nids are negligible (Kitano et al. 1993; Urabe and Nakano
1999). Considering these previous studies, the habitat parti-
tioning along vertical position and proximity to cover in our
sympatric sites is also not likely attributable to habitat shift
by rainbow trout. However, our analysis on stomach con-
tents suggests a diet shift by rainbow trout in the sympatric
sites. Contribution of terrestrial invertebrates to the diet of
sympatric rainbow trout was relatively low, whereas that of
allopatric rainbow trout was as high as that of both allopa-
tric and sympatric masu salmon. Such differences in diet
can be caused by differences in food availability, rather
than by the presence of opponent species. For example, it is
reported that stream salmonids shifted their foraging behav-
ior from drift interception to benthic foraging when drifting
prey abundance declined (Nakano et al. 1999a, 1999b). In
our study, however, the difference in terrestrial prey contri-
bution to rainbow trout diet between sympatric and allopa-
tric situations cannot be explained by differences in drift
availability. Drifting prey density in the sympatric sites (KI
and HN) was not always lower than that in the allopatric
sites (UN and TM). Rather, exceedingly high densities of
drifting prey were recorded in a sympatric site (HN) where
contribution of terrestrial prey to rainbow trout diet was
lowest. Although rainbow trout is thought to feed primarily
on drifting terrestrial prey (Nakano et al. 1999c; Baxter et
al. 2007), these data were obtained from streams where ef-
fects of other salmonids were negligible (very low density
of other salmonids or sympatry with competitively inferior
species). Our results suggest that rainbow trout shift their re-
source use from drifting prey to benthic prey when they are
sympatry with masu salmon. Although, in general, competi-
tive effects on resource use cannot be fully elucidated by
only comparative analyses as ours, our empirical data cor-
roborate results of previous controlled experiments on inter-
actions between rainbow trout and masu salmon (Hasegawa
et al. 2004; Hasegawa and Maekawa 2006).
Management implications
Overall results of our study suggest that (i) flow regime
has an important role in controlling success and failure of
rainbow trout invasion, as hypothesized by Fausch et al.
(2001), and (ii) masu salmon has competitive effects on
Published by NRC Research Press
 Inoue et al.
rainbow trout and vice versa, although native anadromous
masu salmon cannot offer strong biotic resistance to rainbow
trout invasion. Our results have important implications for
habitat management, especially associated with dams. In
general, natural flow regime is thought to favor native as-
semblages, and alterations of flow can facilitate invasions
by non-natives (Poff et al. 1997; Marchetti and Moyle
2001). In regions where flow regimes are different from
those in the native range of rainbow trout, altered flow may
facilitate invasion by rainbow trout, which is otherwise sup-
Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by TPE/MAIN LIBRARY on 04/10/14
pressed by natural flow regimes. In addition, reservoirs cre-
ated by dams may prevent rainbow trout that are flushed
from inlet streams from being lost downstream. Therefore,
dams may lead to the trout’s higher invasibility (see also
Marchetti et al. 2004). However, dams, as a flow-controlling
device, are usable to prevent or mitigate invasion effects by
manipulating discharge to mimic natural flow regimes. It is
important to consider both negative and positive aspects of
dams to seek better ways to manage non-natives and con-
serve native biota.
Acknowledgements
We are grateful to K.D. Fausch, C.V. Baxter,
M. Murakami, and H. Urabe for useful suggestions during
the fieldwork. We also thank the staff and graduate students,
especially M. Miura, at the Tomakomai Research Station,
For personal use only.
Hokkaido University, for help in conducting the fieldwork.
Critical and helpful comments from two reviewers and an
editor greatly improved the manuscript. This research was
planned on the basis of a preliminary survey on fish fauna
in the Iburi District conducted by Y. Shibata, H. Miyasaka,
H. Urabe, H. Onishi, Y. Taniguchi, Y. Kawaguchi, T. Saito,
Y. Miyake, T. Iwata, D. Kishi, H. Saito, K. Murakami, and
S. Nakano and was supported by a Grant-in-Aid for Scien-
tific Research from the Japan Society for the Promotion of
Science (No. 15770009).
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