Global introductions of salmon and trout in the genus Oncorhynchus: 1870–2007

Abstract

The purpose of this review is to provide a global perspective on Oncorhynchus salmonine introductions and put-and-take fisheries based on modern stocking programs, with special emphasis on freshwater
ecosystems. We survey the global introductions of nine selected salmonines of the genus Oncorhynchus: golden trout, cutthroat trout, pink salmon, chum salmon, coho salmon, masu/cherry salmon, rainbow trout/steelhead, sockeye
salmon/kokanee, and chinook salmon. The information is organized on a geographical basis by continent, and then by species
and chronology. Two different objectives and associated definitions of ‘success’ for introductions are distinguished: (a)
seed introduction: release of individuals with the purpose of creating a wild-reproducing, self-sustaining population; and
(b) put-and-take introduction: release of individuals with the purpose of maintaining some level of wild population abundance,
regardless of wild reproduction. We identify four major phenomena regarding global salmonine introductions: (1) general inadequacy
of documentation regarding introductions; (2) a fundamental disconnect between management actions and ecological consequences
of introductions; (3) the importance of global climate change on success of previous and future introductions; and (4) the
significance of aquaculture as a key uncertainty in accidental introductions. We conclude this review with a recognition of
the need to terminate ongoing stocking programs for introduced salmonines worldwide.

Full text

RESEARCH PAPER
Global introductions of salmon and trout in the genus
Oncorhynchus: 1870–2007
Stephen S. Crawford ÆAndrew M. Muir
Received: 5 May 2007 / Accepted: 19 October 2007
ÓSpringer Science+Business Media B.V. 2007
Abstract The purpose of this review is to provide a
global perspective on Oncorhynchus salmonine intro-
ductions and put-and-take fisheries based on modern
stocking programs, with special emphasis on fresh-
water ecosystems. We survey the global introductions
of nine selected salmonines of the genus Oncorhyn-
chus: golden trout, cutthroat trout, pink salmon, chum
salmon, coho salmon, masu/cherry salmon, rainbow
trout/steelhead, sockeye salmon/kokanee, and chi-
nook salmon. The information is organized on a
geographical basis by continent, and then by species
and chronology. Two different objectives and asso-
ciated definitions of ‘success’ for introductions are
distinguished: (a) seed introduction: release of indi-
viduals with the purpose of creating a wild-
reproducing, self-sustaining population; and (b) put-
and-take introduction: release of individuals with the
purpose of maintaining some level of wild population
abundance, regardless of wild reproduction. We
identify four major phenomena regarding global
salmonine introductions: (1) general inadequacy of
documentation regarding introductions; (2) a funda-
mental disconnect between management actions and
ecological consequences of introductions; (3) the
importance of global climate change on success of
previous and future introductions; and (4) the signif-
icance of aquaculture as a key uncertainty in
accidental introductions. We conclude this review
with a recognition of the need to terminate ongoing
stocking programs for introduced salmonines
worldwide.
Keywords Salmoninae Exotics 
Introductions Stocking History
Introduction
The purpose of this review is to provide a global
perspective on salmonine introductions and put-and-
take fisheries based on modern stocking programs,
with special emphasis on freshwater ecosystems. In
addition to providing a general perspective on the
distribution and intensity of stocking efforts, we also
compile and organize important references as a lead
for researchers who wish to pursue specific ecological
or socio-economic aspects of salmonine introduc-
tions. Due to the broad nature of this topic, we do not
attempt to explore the complex patterns in ecological
effects, changing stakeholder values, or the role of
S. S. Crawford (&)
Department of Integrative Biology, University
of Guelph, Guelph, ON, Canada N1G 2W1
e-mail: scrawfor@uoguelph.ca
A. M. Muir
Department of Forestry and Natural Resources, Purdue
University, 195 Marsteller Street, West Lafayette,
IN 47907, USA
e-mail: ammuir@purdue.edu
123
Rev Fish Biol Fisheries
DOI 10.1007/s11160-007-9079-1

ecosystem-based management in shaping policies on
introductions of exotics species.
Specifically, we survey the global introductions
of 9 Pacific salmonines of the genus Oncorhynchus
(Table 1): golden trout (O. aguabonita); cutthroat
trout (O. clarki); pink salmon (O. gorbuscha); chum
salmon (O. keta); coho salmon (O. kisutch); masu/
cherry salmon (O. masou); rainbow trout/steelhead
(O. mykiss); sockeye salmon/kokanee (O. nerka);
and chinook salmon (O. tshawytscha) (see Robins
et al. 1991: Common and Scientific Names of
Fishes, Fifth Edition 1991, American Fisheries
Society Special Publication 20). The information
presented in this review is organized on a geo-
graphical basis by continent, and then by species
and chronology.
We classify salmonine introductions on the basis
of objectives for hatchery stocking programs. At the
most general level, there are two different objectives
and associated definitions of ‘success’ for introduc-
tions that must be clearly distinguished: (1) seed
introduction: release of individuals with the purpose
of creating a wild-reproducing, self-sustaining popu-
lation; and (2) put-and-take introduction: release of
individuals with the purpose of maintaining some
level of wild population abundance, regardless of
wild reproduction (if any). Based on the information
that we have reviewed, it appears that most of the
global salmonine introductions were originally (usu-
ally implicitly) undertaken as seed introductions.
Proponents often wanted to stock fish into a new
ecosystem with hopes that the individuals would
survive in the wild, and reproduce at levels that
would maintain a population large enough to support
some degree of harvest. In many situations where
seed introductions were unsuccessful, stocking efforts
were terminated altogether. However, in other cases
where seed introductions failed, stocking efforts
continued with the (usually implicit) objective of
put-and-take introductions; the proponent wanted to
maintain a hatchery-based artificial population in the
wild to support some degree of harvest. ‘Success’ of a
put-and-take stocking program would then be defined
as the survival of sufficient fish in the wild, long
enough to support harvest. In this review we will
attempt to clearly distinguish between the two kinds
of introductions (seed, put-and-take), and we will
focus on the stocking programs that have been
maintained to the present.
Salmonine introductions
North & Central America
There are numerous reviews of the history, geopolit-
ical distribution and ecological effects of intra-
continental Pacific salmonine introductions in North
America, including: Dymond (1947); Dymond
(1955); MacCrimmon (1965); Moyle (1986); Krueger
and May (1991); Leach and Lewis (1991); Mills et al.
(1993); Crossman and Cudmore (1999); Dextrase and
Coscarelli (1999); Fuller et al. (1999); Kocik and
Jones (1999); Nico and Fuller (1999); Pearsons and
Hoply (1999); Epifanio (2000); Crawford (2001);
Ham and Pearsons (2001); Dunham et al. (2002);
Dunham et al. (2004); and USGS (2004). While it has
not been uncommon for North American seed
introductions to establish self-sustaining populations,
levels of wild recruitment have often been replaced or
heavily supplemented by stocking programs for put-
and-take recreational fisheries (Horak 1995; Epifanio
2000).
The North American (and global) experience with
large-scale Pacific salmonine seed introductions
began in the late 19th century, when the U.S. Fish
Commission directed the construction of hatcheries
and rail systems to transport salmonines from their
native range to locations throughout continental
North America and abroad. Western and eastern
states of the USA received the majority of intentional
salmonine seed introductions (Rahel 2000). Cur-
rently, put-and-take recreational fisheries for Pacific
salmonines in the USA are dispersed across 50 states
(Figs. 1–9), most beyond the native range of the
target species (Waddington and Laughland 1996).
More than half of recreational fishing for these
species in the USA occurs in six states (California,
Pennsylvania, Colorado, New York, Washington, and
Oregon), while seven states (previous six states, plus
Maine) annually receive more than 50% of USA
‘trout’ fishing effort (Epifanio 2000).
Golden trout (Oncorhynchus aguabonita)
Golden trout are native to western North America
where they are endemic to the Kern Plateau, Tulare
and Kern Counties, California (Table 1; Fig. 1;
Behnke 1992). There is a long, yet largely
Rev Fish Biol Fisheries
123

Table 1 Introduced Pacific salmonines (Family Salmonidae, Subfamily Salmoninae, Genus Oncorhynchus) and their native distributions, based on continent and major
watershed
Species Common Continent Native distribution Major watershed(s)
O. aguabonita Golden trout North America Upper Kern River basin, Tulare and Kern counties, California,
USA
Sacramento
O. clarki Cutthroat trout North America Pacific Coast drainages from Prince William Sound, Alaska, to Eel
River, northern California. Freshwater populations range
through Rocky Mountains to Hudson Bay, Mississippi River,
Great (including Lahontan, Bonneville, and Alvord basins), and
Pacific basins from southern Alberta to Rio Grande drainage,
New Mexico.
Fraser, Columbia, MacKenzie
O. gorbuscha Pink salmon Asia North America Arctic and Pacific drainages from Mackenzie River delta,
Northwest Territories, to Sacramento River drainage, California;
occasionally as far south as La Jolla, southern California.
Northeastern Asia, Korea, Japan, Okhotsk and Bering Sea.
Amur, Yukon, Fraser, Columbia,
MacKenzie, Sacramento
O. keta Chum salmon Asia North America Arctic and Pacific drainages from Anderson and Mackenzie rivers,
Northwest Territories, through much of Alaska south to
Sacramento River drainage, California; rarely to San Lorenzo
River, southern California. Northeastern Asia, Korea, Japan,
Okhotsk and Bering Sea.
Amur, Yukon, Fraser, Columbia,
MacKenzie, Sacramento
O. kisutch Coho salmon Asia North America Arctic and Pacific drainages from Point Hope, Alaska, to Monterey
Bay, California, infrequently as far south as Chamalu Bay, Baja
California. Northeastern Asia, Japan, Russia.
Anadyr, Yukon, Fraser, Columbia,
MacKenzie, Sacramento
O. masou Cherry salmon Asia Marine and freshwaters of Japan, Korea, and Russia, Okhotsk and
Bering Sea.
Amur
O. mykiss Rainbow trout/
steelhead
Asia, North America Pacific Slope from Kuskokwim River, Alaska, to (at least) Rio
Santa Domingo, Baja California; upper MacKenzie River
drainage (Arctic basin), Alberta and British Columbia;
endorheic basins of southern Oregon.
Amur, Yukon, Fraser, Columbia,
MacKenzie, Sacramento
O. nerka Sockeye salmon Asia, North America Arctic and Pacific drainages from Point Hope, Alaska, to
Sacramento River drainage, California. Landlocked populations
in Alaska, Yukon, British Columbia, Washington, and Oregon.
Northeastern Asia, Japan, Kamchatka Peninsula, Okhotsk and
Bering Sea.
Yukon, Fraser, Columbia,
MacKenzie, Sacramento
O. tshawytscha Chinook salmon Asia, North America Arctic and Pacific drainages from Point Hope, Alaska, to Ventura
River, California. Occasionally strays south to San Diego,
California. Northeastern Asia, Japan, Okhotsk and Bering Sea.
Yukon, Fraser, Columbia,
MacKenzie, Sacramento
Sources: Beacham et al. 2006; Behnke 1992; Behnke 2002b; Berg 1977; Coad 1996a; Coad 1996b; Froese and Pauly 2004; Morrow 1980; Page and Burr 1991; Popov 1933; Scott
and Crossman 1973; Svetovidov 1984; USGS 2004; Welcomme 1988
Rev Fish Biol Fisheries
123

undocumented history of golden trout introductions
in North America. Beginning in the mid 1800s, this
species was intentionally seeded in hundreds of
typically fishless mountain watersheds in the Rocky
Mountain states and provinces (Page and Burr 1991;
Bradford et al. 1993,1998; Epifanio 2000; Knapp
et al. 2001). In the late 1950s, golden trout were
introduced (mostly unsuccessfully) from California to
the lower Fraser River, Arctic drainages, and high
altitude lakes of southern Alberta (Crossman 1984;
Welcomme 1988; Crossman 1991; McPhail and
Carveth 1993; Coad 1995; Lever 1996). Put-and-take
introductions of golden trout have continued in Idaho,
Montana, Oregon, and Washington (Fig. 1).
Cutthroat trout (Oncorhynchus clarki)
The cutthroat trout (i.e., including O. clarki and the
sub-species recognized by Behnke (1992) is the most
widely distributed of the American western trout
species (Behnke 1992). This species is native to
coastal and mountain watersheds in Alaska, British
Columbia, Washington, Oregon, California, and
extends as far inland as Alberta in Canada, central
Colorado (extirpated), Idaho, Montana, Nevada,
northwestern New Mexico, Texas (extirpated), Utah,
and Wyoming in the USA (extirpated; Table 1;
Fig. 2). However, it is only in BC and Alaska where
healthy populations persist (Scott and Crossman
1973; Morrow 1980; Epifanio 2000).
Introductions began in the 1890s when the State of
Michigan received Yellowstone cutthroat trout from
an American federal fish hatchery in Colorado
(Worth 1895); these fish were seeded into the Pere
Marquette River of Lake Michigan (Emery 1985;
Crawford 2001). Similar efforts were undertaken by
various jurisdictions to introduce cutthroat trout to
the Great Lakes; however, by 1958 this species was
considered rare or extinct in the Great Lakes basin
(Hubbs and Lagler 1958). During the period 1910-
1970, cutthroat trout were seeded, often with rainbow
trout, to many western USA mountain lakes and
streams where they did not previously exist,
Fig. 1 Native distribution of golden trout (Oncorhynchus
aguabonita; black stipple) and countries, states or provinces
into which this species has been introduced (red stipple);
sources for native distribution: Behnke (1992); sources for
introductions are provided within the text
Rev Fish Biol Fisheries
123

including the many fishless alpine lakes of the Sierra
Nevada (Kucera et al. 1985; Drake and Naiman
2000; Dunham et al. 2004). Cutthroat trout were also
introduced to a relatively few number of lakes in
eastern North America (Froese and Pauly 2004) and
southwestern USA, often for put-and-take fisheries in
private ponds (Morrow 1980; Nico and Fuller 1999).
Unsuccessful seed introductions to Canadian water-
sheds (Table 1) include the Similkameen and lower
Peace Rivers (McPhail and Carveth 1993) and
various lakes and streams in the Northwest Territo-
ries, Alberta, Saskatchewan, Manitoba, Ontario,
Quebec (Coad et al. 1995; Crossman 1991; Froese
and Pauly 2004).
Pink salmon (Oncorhynchus gorbuscha)
Within North America, the native distribution of pink
salmon extends from the Mackenzie River basin in
the Canadian Arctic west and south through the
Bearing Sea, Aleutian Islands, Alaska, British
Columbia, Washington, Oregon, northern California,
and occasionally straying as far as La Jolla, southern
California (Table 1; Fig. 3; Epifanio 2000). Spawn-
ing populations were recently recorded as far north as
Banks Island, NT (Babaluk et al. 2000; Stephenson
2005).
According to available records there have only
been a handful of attempted seed introductions of
pink salmon outside of their native distribution
(Table 1; Fig. 3). In the late 1950s, 15 million pink
salmon embryos were transported from the Skeena
watershed in British Columbia and seeded to tribu-
taries from Labrador to the north shore of Quebec
where the survivors gradually died out (Scott and
Crossman 1973; Page and Burr 1991; van Zyll de
Jong et al. 2004). There are also reports that pink
salmon were unsuccessfully introduced to tributaries
in Nova Scotia, New Brunswick and Hudson Bay
tributaries in northern Ontario (Crossman 1991).
Perhaps the most surprising introduction of pink
salmon in North America is the accidental, yet
successful, seeding of the Laurentian Great Lakes. In
Fig. 2 Native costal (black diagonal) and inland (black
stippled) distribution of cutthroat trout (Oncorhynchus clarki)
and countries, states or provinces into which this species has
been introduced (red stipple); Sources for native distribution:
Scott and Crossman (1973), Behnke (1992,2002b); sources for
introductions are provided within the text
Rev Fish Biol Fisheries
123

1955, pink salmon embryos were transported from
British Columbia to an Ontario hatchery for incuba-
tion before being air-lifted for release in Goose Creek
in the Hudson Bay drainage basin (Ricker and Loftus
1968; Crawford 2001). Through a combination of
accidents in technical operations, and by the inten-
tional discharge of excess stock into the Current
River, the Ontario hatchery had inadvertently intro-
duced pink salmon to the Great Lakes (Collins 1975).
After 1958, small but increasing numbers of pink
salmon were taken by commercial and recreational
fishermen (Scott and Crossman 1973). The pink
salmon in Lake Superior soon spread to Lake Huron
where spawning in tributaries was first observed in
1969 (Collins 1975), and subsequently populations
were established in each of the remaining Great
Lakes–Michigan in 1973, Lakes Erie and Ontario by
1979 (Crawford 2001). This unintentional introduc-
tion resulted in colonization of all five Great Lakes,
and even further to the St. Lawrence River, without
supplemental stocking within a mere 10 generations
(Scott and Crossman 1973; Emery 1985; Kwain
1987; Cudmore-Vokey and Crossman 2000).
Chum salmon (Oncorhynchus keta)
Chum salmon have the widest natural geographic and
spawning distribution of any Pacific salmonine
(Table 1; Fig. 4). Within the Americas, the native
distribution of chum salmon extends west from
Kugluktuk, Nunavut throughout the Mackenzie drain-
age and coastal Yukon south through the Bering Sea,
Aleutian Islands, coastal Alaska, British Columbia,
Washington, Oregon, northern and southern Califor-
nia (Aro and Shepard 1967; Hallock and Fry 1967;
Hart 1973; Stephenson 2005).
Given the chum salmon’s broad native range, it is
somewhat surprising that the history of North Amer-
ican seed introductions was as brief as it was
unsuccessful. Despite a handful of failed attempts in
USA and Canada (Crossman 1991; Epifanio 2000;
Fig. 3 Native distribution of pink salmon (Oncorhynchus
gorbuscha; black diagonal) and countries, states or provinces
into which this species has been introduced (red stipple). Black
bars represents the primary distribution, but spawning fish have
been reported throughout the entire range; Sources for native
distribution: Berg (1948); Hart (1973), Scott and Crossman
(1973), Babaluk et al. (2000), Behnke (2002b), Stephenson
(2005); sources for introductions are provided within the text
Rev Fish Biol Fisheries
123

Page and Burr 1991), no wild populations were
established or maintained by put-and-take stocking.
From approximately 1910 to the early 1940s, hatch-
eries in Michigan, Pennsylvania and Wisconsin
provided juvenile chum salmon for release into
waters of Lake Superior and Lake Huron; however,
none of these efforts led to the establishment of self-
sustaining populations. By 1945, these chum salmon
stocking programs were terminated (MacCrimmon
1977; Crawford 2001). In the mid-1950s, there was
an unsuccessful attempt to take chum salmon from
Puget Sound (Washington) and introduce them to
Hudson Bay tributaries in northern Ontario, including
the Winisk and Attawapiskat Rivers (Fig. 4; Scott
and Crossman 1973).
Coho salmon (Oncorhynchus kisutch)
Within the Americas, the native distribution of coho
salmon extends south from Point Hope Alaska,
through the Aleutian Islands, coastal British Colum-
bia, Washington, Oregon, to the Sacramento drainage
in California, occasionally as far as La Jolla, southern
California, inland to Idaho (extirpated), and Nevada
(extirpated; Table 1; Fig. 5; Hart 1973; Scott and
Crossman 1973; Epifanio 2000; Behnke 2002b;
Kaczynski and Alvarado 2006). During the period
1930–1950, fish culturists unsuccessfully attempted
to seed coho salmon to a large number of states and
provinces where it did not previously exist (Fig. 5;
Walker et al. 1961; Crossman 1991; Epifanio 2000;
USGS 2004).
The first attempt to seed coho salmon to the Great
Lakes drainage basin occurred in the 1870’s when
government hatcheries in Ontario, Ohio, and Mich-
igan released thousands of juveniles into Lake Erie
and its tributaries (Crawford 2001). However, no self-
sustaining populations were established from this
early round of introductions, and stocking was
terminated in 1935 (Mills et al. 1993; Scott and
Crossman 1973). An aggressive coho salmon
Fig. 4 Native distribution of chum salmon (Oncorhynchus
keta; black diagonal) and countries, states or provinces into
which this species has been introduced (red stipple). Sources
for native distribution: Berg (1948), Hart (1973), Scott and
Crossman (1973), Babaluk et al. (2000), Behnke (2002b),
Stephenson (2005); sources for introductions are provided
within the text
Rev Fish Biol Fisheries
123

introduction program was re-initiated in the Great
Lakes by the States of Michigan and Ohio in the mid-
1960s (Parsons 1973). Unlike previous attempts, the
initial return from these plantings was described as
immediate and spectacular (Crossman 1968). From
1966 to 1998, a total of more than 148 million coho
salmon were introduced to the Great Lake ecosystem,
many of which were stocked on a put-and-take basis
(Crawford 2001).
Masu salmon (Oncorhynchus masou)
Masu or cherry salmon are not native to any North or
Central America watersheds (Fig. 6) and according to
available records, there have only been a few
attempts to seed this species outside of its native
Asian distribution. In 1929 the State of Michigan
unsuccessfully attempted to seed a tributary of Lake
Michigan, and in 1965 Ontario stocked 5,500 juve-
niles in Lake Westward in Algonquin Park ‘‘as part
of failed attempt to enhance Great Lakes fisheries’’
(Westerman 1930; Christie 1970; Crossman 1984;
Welcomme 1988; Crawford 2001). Masu salmon
were also unsuccessfully seeded into Hewitt Lake in
Washington in the 1970s (Courtenay et al. 1984;
USGS 2004).
Rainbow trout (Oncorhynchus mykiss)
Within North America, rainbow trout (including
redband and steelhead trout) are native to the Pacific
northwest (Table 1; Fig. 7). Their coastal distribution
extends from the Bering Sea and Bristol Bay,
Aleutian Islands, Alaska, south throughout British
Columbia, Washington, Oregon, northern and south-
ern California. Non-migratory rainbow and redband
trout are also native to southwestern Alaska, the
Stikine, Skeena, Fraser, and Peace River drainages in
BC and extended as far inland as the Kootenay River,
Montana (Behnke 1992), the headwaters of the
Athabasca in Alberta, and the lower Columbia, San
Joaquin, Sacramento, and Klamath drainages in
Fig. 5 Native distribution of coho salmon (Oncorhynchus
kisutch; black diagonal) and countries, states or provinces into
which this species has been introduced (red stipple). Sources
for native distribution: Berg (1948), Hart (1973), Scott and
Crossman (1973), Behnke (2002b), Stephenson (2005); sources
for introductions are provided within the text
Rev Fish Biol Fisheries
123

California (Behnke 1992; Epifanio 2000; Hart 1973;
Scott and Crossman 1973). It is possible that rainbow
trout are also native to high elevation tributaries in
northwest Mexico (Needham and Gard 1959; Behnke
1972), however this record has been questioned
(MacCrimmon et al. 1970).
As for other continents, rainbow trout is one of the
most widely introduced salmonines in North and
Central America (Moyle 1986; Crossman 1991;
Crossman and Cudmore 1999; Dextrase and Coscar-
elli 1999). During the early period of seed
introductions, transcontinental railways shipped east-
ern fishes to western watersheds, and returned with
shipments of rainbow trout that were stocked in the
prairies or plains, the Mississippi drainage and
throughout the eastern watersheds (Trautman 1981;
Nico and Fuller 1999; Moring 2000). For example,
Pister (2001) wrote:
As the ‘‘fish cars’’ pulled into railroad sidings
along the way, they would be met by different
groups (sportsmen and so on) who would ask
for (as an example) two cans of eastern brook
(Salvelinus fontinalis) two of brown trout
(Salmo trutta), and two of rainbow. The hatch-
ery personnel would dutifully procure six cans
of steelhead, label two of them ‘‘eastern brook,’’
two of them ‘‘brown,’’ and two of them
‘‘rainbow,’’ and the sportsmen would take it
from there, happier but never the wiser. This
example may be a bit extreme, but it is true and
underscores the chaos of early-day trout distri-
bution. No one really knew what went where,
when, or why. (Pister 2001).
In a recent survey of introductions to US states,
rainbow trout was determined to be the fourth most
frequently seeded species, ranking only after (1)
common carp Cyprinus carpio, (2) goldfish Carassius
auratus, and the European salmonine (3) brown trout
(Rahel 2000).
Rainbow trout were first seeded to the Great Lakes
in 1876 when individuals were released to Michigan
tributaries of Lake Huron (Smedley 1938; Kocik and
Jones 1999). The early American and Canadian
introductions of rainbow trout to the Great Lakes
Fig. 6 Native distribution of masu salmon (Oncorhynchus
masou; black diagonal) and countries, states or provinces into
which this species has been introduced (red stipple). Sources
for native distribution: Berg (1948), Hart (1973), Machidori
and Kato (1984); sources for introductions are provided within
the text
Rev Fish Biol Fisheries
123

basin met with a general, yet limited degree of
success (MacCrimmon 1977). In the 1930s, fish
culturists in Minnesota began a new initiative in
rainbow trout stocking in the Great Lakes drainage
basin by successfully manipulating the life-history of
the typical spring-spawning rainbow trout to become
a fish that would spawn between November and
January (MacKay 1969; MacCrimmon 1977). During
the period 1966–1998, a total of 174 million rainbow
trout had been stocked to the Great Lakes ecosystem
(Crawford 2001).
Between 1870 and 1970, attempts were made to
introduce rainbow trout to virtually every other North
American state and province/territory with the excep-
tions of Florida and Northwest Territories (Table 2).
These introductions led to the establishment of self-
sustaining populations in 39 of the 42 states including
Hawaii (Courtenay et al. 1984; Rinne and Janisch
1995; Hayes et al. 1996; Thurow et al. 1997; Nico and
Fuller 1999; Epifanio 2000; Rahel 2000; Adams et al.
2001; Knapp et al. 2001; Weigel et al. 2003; Dunham
et al. 2004; McHugh and Budy 2006), and 8 of the 9
provinces/territories outside the native range (Mac-
Crimmon 1971; Scott and Crossman 1973; Crossman
1984;1991; Leach and Lewis 1991; Kerr and Grant
2000; Kerr and Lasenby 2000; Crawford 2001; DFO
2003; van Zyll de Jong et al. 2004). It is highly likely
that in all of these cases, wild reproduction is heavily
supplemented with ongoing introduction in the form
of put-and-take hatchery stocking.
In a recent survey of North American hatcheries,
Barnes and Whelan (2004) found that 65% of all US
(state: 76%, federal: 49%) and Canadian (17%)
hatcheries ranked rainbow trout as the primary
species for production. Although this survey did not
decipher whether the hatchery production was for
release into native waters (i.e. supplementation) or
non-native waters (i.e. introduction), more than 80%
of the responding hatcheries were located in states/
provinces outside of rainbow trout native distribution
(Table 2; Fig. 7).
Beyond the USA and Canada, rainbow trout were
also seeded to a variety of Central American water-
sheds. Between 1920 and 1950, rainbow trout were
Fig. 7 Native distribution of rainbow trout (Oncorhynchus
mykiss; black diagonal) and countries, states or provinces into
which this species has been introduced (red stipple). Sources
for native distribution: Berg (1948), Hart (1973), Scott and
Crossman (1973), Behnke (1992,2002b); sources for intro-
ductions are provided within the text
Rev Fish Biol Fisheries
123

Table 2 Summary of salmonine introductions in North America based on species and consequences of attempted introduction
Species Seed introduction Put-and-take introduction
Fail Success
Golden trout (O. aguabonita) AB, AZ, BC, CO, ID, MT, NV, NM, OR, WY CA, UT, WA ID, MT, OR, WA
Cutthroat trout (O. clarki) AB, AR, AZ, CA, CO, CT, KS, MB, MD, MI,
MN, MT, NB, NJ, NM, NV, ND, NT, ON,
OR, QC, SD, SK, TN, UT, VT, VA, WA, WI,
WV, WY
AZ, CA, CO, MT, NV, OR, UT, WA AR, AZ, MD
Pink salmon (O. gorbuscha) MD, ME, ON, NB, NL, NS, QC Great Lakes [IL, IN, MI, MN, NY, OH,
ON, PA, WI]
none
Chum salmon (O. keta) BC, ID, ME, MI, MX, NV, ON, UT none? none
Coho salmon (O. kisutch) AK, AZ, BC, CA, CO, CT, DE, IA, ID, IL, IN,
KT, MA, MD, ME, MI, MN, MT, NB, ND,
NH, NV, NM, NY, OH, ON, OR, PA, SD, TX,
TN, VG, UT, WA, WI, WV, WY, YT
Great Lakes? ND and Great Lakes [IL, IN, MI, MN,
NY, OH, ON, PA, WI]
Masu salmon (O. masou) MI, ON, WA none none
Rainbow trout/steelhead (O.
mykiss)
AL, AK, AR, AZ, CA, CO, CT, DE, FL, GA, HI,
IA, ID, IL, IN, KS, KY, LA, MA, MD, ME,
MI, MN, MO, MS, MT, NB, NC, ND, NH, NJ,
NL, NM, NY, OH, OK, ON, OR, PA, PR, SC,
SD, TN, TX, UT, VG, VT, WI, WV, WY,
Puerto Rico, Dominican Republic, Mexico?,
Cuba?
‘‘established in many states [and
provinces]’’ Costa Rica, Panama
AL, AZ, AR, CO, CT, DE, GA, HI, IL,
IA, KS, KY, ME, MD, MA, MI, MN,
MO, NB, NH, NJ, NM, NY, NC, ND,
OH, OK, PA, RI, SC, SD, TN, TX, UT,
VT, VG, WV, WI, WY
Sockeye salmon/kokanee (O.
nerka)
AB, AZ, BC, CA, CO, CT, ID, MA, ME, MI,
MN, MO, MT, NB, NC, ND, NM, NV, NY,
ON, OR, PA, SD, TN, UT, VT, WA, WI, WV,
WY
‘‘established locally in some areas’’ CO,
NM, NY, SD, UT, WY
CO, NM, SD, UT, WY
Chinook salmon (O.
tshawytscha)
BC, CO, CT, DE, HI, IA, IL, IN, KS, MA, MD,
ME, MI, MN, MT, MX, NB, ND, NH, NJ, NS,
NV, NY, OH, ON, PA, QC, SD, TX, VG, VT,
WI, WV
‘‘most attempts to establish populations
failed’’ Great Lakes? ND, SD
ND, SD and Great Lakes [IL, IN, MI,
MN, NY, OH, ON, PA, WI]
Sources: Epifanio (2000), Crawford (2001), USGS (2004); references cited within the text; abbreviations for states and provinces as presented by Canada Post
(http://www.canadapost.com, cited May 2007)
Rev Fish Biol Fisheries
123

introduced from the USA to Panama and Puerto Rico,
with the former establishing abundant self-sustaining
population in high elevation ([1,000 m) streams
(MacCrimmon 1971; Erdman 1984; Welcomme
1988). In the 1950s, rainbow trout from the USA
were also successfully introduced to high elevation
localities in Costa Rica and Honduras (Welcomme
1988; Bussing 1998). During the 1980s, rainbow
trout were introduced to Guatemala then the Domin-
ican Republic where there was a recent report of an
established wild population (MacCrimmon 1971;
Welcomme 1988; FAO 1997; Sartorio 2003). Mac-
Crimmon (1971) described the seeding of rainbow
trout in Mexico and Cuba with unknown, yet
doubtful, consequences.
Sockeye salmon (Oncorhynchus nerka)
Within North America, sockeye salmon and kokanee
(its freshwater form) are native to the Pacific
northwest (Table 1; Fig. 8). According to Behnke
(1972), it is probable that all lacustrine populations of
O. nerka were independently derived from anadro-
mous populations in postglacial times. Although
Babaluk et al. (2000) reported reproductively mature
individuals as far north as the Sachs River, Banks
Island, Northwest Territories, this species is primarily
concentrated from Point Hope, Alaska south through
the Bering Sea, Aleutians, Yukon Territory, British
Columbia, Washington, Oregon to the Klamath River
system in northern California (Taft 1937; Scott and
Crossman 1973; Crossman 1984; Moyle 1986;
Burgner 1991; Epifanio 2000).
Sockeye salmon and kokanee have been seeded to
a variety of states and provinces in North America
(Crossman 1984; Epifanio 2000; Moyle 1986; Scott
and Crossman 1973). Kokanee were unsuccessfully
seeded to Lake Tahoe (Moyle 1976) and various
mountain lakes of the Sierra Nevada, many of which
were previously fishless (Moyle 1986; Knapp and
Matthews 2000). Beginning in 1917 and continuing
Fig. 8 Native distribution of sockeye salmon (Oncorhynchus
nerka; black diagonal) and countries, states or provinces into
which this species has been introduced (red stipple). Black bars
represents the primary distribution, but spawning fish have
been reported throughout the entire range; Sources for native
distribution: Berg (1948), Hart (1973), Scott and Crossman
(1973), Behnke (2002b), Stephenson (2005); sources for
introductions are provided within the text
Rev Fish Biol Fisheries
123

on to the late 1940s, kokanee were also introduced to
Lake Washington (Hendry et al. 1996; Hendry and
Quinn 1997), ultimately leading to a complex pop-
ulation which is a mixture of both native and
introduced sockeye salmon (Hendry et al. 1998).
Kokanee have been introduced to Canadian provinces
from Alberta, Saskatchewan, Manitoba, Ontario to
Quebec (Crossman 1984,1991).
The first documented introduction of kokanee to
the Great Lakes occurred in 1950, when the New
York government seeded kokanee juveniles in Lake
Ontario tributaries (Parsons 1973; Emery 1985;
Crawford 2001). In 1964, Ontario received 1.5 mil-
lion embryos from British Columbia (MacCrimmon
1977; Scott and Crossman 1973), and began releasing
them in tributaries and open waters of Lakes Huron
and Ontario, with some considerable success in
returning adults (Collins 1971; Parsons 1973). How-
ever, by the early 1970s, the spawning runs had
effectively ceased due to a combination of reduced
stocking and their inability to establish self-sustaining
populations through wild reproduction (MacKay
1969; Emery 1985). All kokanee stocking programs
were terminated by the Province of Ontario in 1972
(Mills et al. 1993).
Chinook salmon (Oncorhynchus tshawytscha)
Within North America, chinook salmon are also
native to the Pacific northwest (Table 1; Fig. 9; Scott
and Crossman 1973; Epifanio 2000). Their North
American native range extends from the Coppermine
River, Nunavut in the Canadian Arctic, west and
south throughout costal Northwest and Yukon Terri-
tories, the Bering Sea, Aleutian Islands, British
Columbia, Washington, Oregon, to the Ventura River
system in southern California (Aro and Shepard
1967; Atkinson et al. 1967; Hart 1973; Healey 1991).
Beginning in the 1870s, the first shipments of
chinook embryos were collected from California egg-
taking facilities—notably the Baird fish station on the
McCloud River (Sacramento watershed), and trans-
ported along with rainbow trout (see above) by
Fig. 9 Native distribution of chinook salmon (Oncorhynchus
tshawaytscha; black diagonal) and countries, states or prov-
inces into which this species has been introduced (red stipple).
Sources for native distribution: Berg (1948), Hart (1973), Scott
and Crossman (1973), Behnke (2002b), Stephenson (2005);
sources for introductions are provided within the text
Rev Fish Biol Fisheries
123

railway throughout the USA, Canada and beyond
(Nico and Fuller 1999; Moring 2000; Yoshiyama and
Fisher 2001). As soon as they became available in the
1870s, chinook salmon were introduced to each of the
Great Lakes by both American and Canadian agen-
cies and station operators (Crawford 2001). Hundreds
of thousands of chinook salmon continued to be
released to each of the Great Lakes until approxi-
mately 1880 (MacCrimmon 1977; Scott and
Crossman 1973). By the early 1880s chinook salmon
in the Great Lakes had largely died out, leading both
US and Canadian introduction plans to be terminated
(Dymond et al. 1929; Peck et al. 1999). In the 1960s,
Michigan stimulated another intensive round of
chinook salmon introductions to the Great Lakes
by most of the other states and provinces, this time
at stocking levels that dwarfed all previous efforts.
As a result of these intensive efforts, 336 million
chinook salmon were introduced to the Great Lakes
ecosystem between 1966 and 1998 (Crawford 2001).
By the 1980s, chinook salmon in the Great Lakes
had begun to exhibit symptoms of overstocking such
as disease outbreaks, declining body condition and
massive mortalities (Keller et al. 1989; Stewart and
Ibarra 1991), after which stocking rates were
significantly reduced. Currently, the degree of wild
reproduction by chinook salmon remains a key
uncertainty, largely due to the fact that all wild
chinook populations in the Great Lakes are still
heavily supplemented with put-and-take stocking
programs. Recent estimates based on creel and river
surveys have indicated that 30–50% of Lake Ontario
chinook salmon are of wild origin (Berends 2004;
Gross et al. 2004); however, it is doubtful that these
populations would persist without the ongoing
supplementation provided by the put-and-take stock-
ing programs.
Europe
Salmonines of the genus Onchorynchus are not native
to any of the 29 major European watersheds, but there
is a long and complex history of introductions in this
region, which has not received the kind of scholarly
attention that might be expected (Table 1; Figs. 1–9).
To date, the only major work that has focused on the
history, distribution and effects of European fish
introductions was presented by Holcı
´k(1991).
Golden trout (Oncorhynchus aguabonita)
There have been a few, unsuccessful attempts to seed
golden trout to European watersheds, including:
1960s transfers from the USA to Denmark then
Sweden (Muus and Dahlstro
¨m1968; Welcomme
1988; Kullander 1999), and also a 1980 attempt from
the USA to a Cyprus via fish farm releases (Fig. 1;
Welcomme 1988).
Cutthroat trout (Oncorhynchus clarki)
Cutthroat trout were introduced in the 1950s and
1960s from the USA to Denmark then Sweden
(Fig. 2; Welcomme 1988). As with golden trout,
cutthroat trout, were also shipped from the USA to
the Cyprus fish farm and were subsequently released
to the wild (Welcomme 1988). None of these
attempted seed introductions was successful.
Pink salmon (Oncorhynchus gorbuscha)
There is a somewhat more varied history of pink
salmon seed introductions to European watersheds
(Fig. 3). In the 1950s, pink salmon were introduced
from the Kola River of the former USSR to Finland,
Norway, Sweden, and Poland (Berg 1977; Bakshtan-
sky 1980; Welcomme 1988; Holcı
´k1991; Lever
1996; Kullander 1999). Also in the 1950s, a few pink
salmon were captured on the West Greenland Shelf
and in Icelandic waters; these were hypothesized to
have strayed across the Atlantic from the attempted
Newfoundland introduction in 1956 (Svetovidov
1984; Jonsson 1992; Nielsen and Bertelsen 1992).
During the 1960s, wild pink salmon, presumably
from the Scandinavian stocking efforts, migrated to
other tributaries of the Baltic Sea, in Germany and
Latvia, as well as English, Scottish, and Irish waters
(Welcomme 1988; Swaby and Potts 1990; Holcı
´k
1991; Winkler et al. 2000). Pink salmon were also
unsuccessfully seeded during the 1960s from
unknown sources (perhaps the former USSR) to
tributaries of the Baltic Sea and Israel (Coad 1996a).
Chum salmon (Oncorhynchus keta)
There have been suggestions that east-west intro-
ductions of chum salmon were attempted by the
Rev Fish Biol Fisheries
123

former USSR as early as 1932, and that up until the
1960s, individuals from these transplants were
reported to have migrated to Norway, Finland,
Latvia, and many other tributaries of the Baltic
Sea (Welcomme 1988; Holcı
´k1991; Lever 1996;
Appleby 1999; Winkler et al. 2000). Later in the
1960s, chum salmon were seeded from an unknown
source to the Caspian Sea, where contested rumours
claim that it is still occasionally captured in the wild
(Walczak 1972; Magomedov 1978; Holcı
´k and
Razavi 1992).
Coho salmon (Oncorhynchus kisutch)
Seed introductions of coho salmon to European
waters began relatively late in comparison to other
Pacific salmonines. The first recorded coho salmon
introduction was in 1971 when fish from the USA
were imported by French fish culturists and acciden-
tally released to local inland watersheds where they
persisted for only a short time (Fig. 5; Welcomme
1988; Keith et al. 1992; Lever 1996). Shortly there-
after, coho salmon were also introduced from Canada
to a hatchery in Greece, and then unsuccessfully
introduced to local lakes and reservoirs (Welcomme
1988; Economidis 1991; Crivelli 1995; Lever 1996;
Economidis et al. 2000).
During the 1980s coho salmon in France were
introduced to Belgium (Lever 1996), the Netherlands
and Sweden (de Groot 1985; Welcomme 1988; Lever
1996; Kullander 1999). In the 1980s, and again in the
1990s, coho salmon from the USA were introduced to
the Jordan River, Lake Kinneret, and Upper Galilee
of Israel (Golani and Shefler 1985; Golani and Mires
2000; Froese and Pauly 2004). Largely undocu-
mented reports also exist for introductions of coho
salmon from the USA and former USSR to tributaries
of the Baltic Sea (Winkler et al. 2000).
Masu salmon (Oncorhynchus masou)
A single report exists for masu salmon being
introduced to European waters, and that is a 1976
unsuccessful attempt to establish a recreational
fishery in Germany using fish from Japan (Fig. 6;
Welcomme 1988; Holcik 1991).
Rainbow trout (Oncorhynchus mykiss)
During the early days of salmonine introductions
(1880–1900), rainbow trout were extensively intro-
duced from the USA to various European nations,
including: Great Britain (Frost 1974; Welcomme
1988; Lever 1996); France (Welcomme 1988; Lever
1996; Keith and Allardi 1998); Germany (Lever
1996; Muus and Dahlstro
¨m1968; Welcomme 1988;
Winkler et al. 2000); Portugal (FAO 1997; Welcom-
me 1988); Spain (Welcomme 1988; Elvira 1998);
Belgium (Lever 1996; Welcomme 1988); Estonia;
Finland (Koli 1990; FAO 1997; Winkler et al. 2000);
Sweden (Landergren 1999; Holmlund 2000; Holml-
und and Hammer 2004); Hungary (FAO 1997;
Welcomme 1988); Turkey (Innal and Erk’akan
2006); Yugoslavia (Nijssen and de Groot 1974;
Welcomme 1988; Lever 1996); Italy (Gandolfi
et al. 1991; Bianco and Ketmaier 2001); Cyprus
(Lever 1996; Welcomme 1988); and possibly the
Netherlands and Liechtenstein (Holcı
´k1991; Wel-
comme 1988; Fig. 7). These early seed introductions
from the Pacific northwest were quickly followed by
an intense round of secondary intra-European trans-
fers of rainbow trout, especially from German
hatcheries, to Austria, Bulgaria, Denmark, Poland,
Sweden, Switzerland, Russia, Slovakia, Czech
Republic, and Estonia (Welcomme 1988; Koli
1990; Holcı
´k1991; Frier 1994; Lever 1996; FAO
1997; Flajshans et al. 1999; Kullander 1999; Winkler
et al. 2000; Blanchet et al. 2007).
During the period 1900–1940, rainbow trout were
introduced from Denmark to Norway, probably
unsuccessfully, despite intensive stocking efforts
(Holcı
´k1991; Welcomme 1988). Swiss stocks were
used as the source for the original rainbow trout
introductions to Israel (then Mandatory Palestine),
including the Dan and Jordan Rivers (Golani and
Mires 2000; Welcomme 1988). Rainbow trout intro-
ductions were repeated in most tributaries (including
mountain streams) of Bulgaria during 1920s (Lever
1996; Welcomme 1988). Similarly, repeated attempts
were made to introduce rainbow trout from other Great
Britain tributaries and USA to Ireland (Went 1957;
Welcomme 1988; Lever 1996; Griffiths 1997) and
from USA to Slovaki and the Czech Republic (Lever
1996; Petr Hartvich and Pavel Vra
´na pers. comm.).
During the period 1940–1975, rainbow trout were
seed introduced from Denmark to Lebanon, Iceland,
Rev Fish Biol Fisheries
123

and Luxembourg (Welcomme 1988; Jonsson 1992).
They were then introduced from Lebanon and
Germany to Syria in 1950s and 1960s (Welcomme
1988), and from Switzerland to many locations in
Greece (Economidis 1991; Economidis et al. 2000;
Welcomme 1988). Rainbow trout were successfully
introduced to northern Mediterranean watersheds of
Italy, Spain, France, Slovenia, Albania, Montenegro,
Portugal, and Greece (Crivelli 1995). There are also
reports of rainbow trout stocking from primary
sources in North America (and secondary sources in
Africa, Europe, India, and Japan) in Iraq, Lebanon
Saudi Arabia, and Jordan (Welcomme 1988; Krupp
and Schneider 1989; Coad 1996a). Repeated rainbow
trout intra-continental transfers also took place after
unsuccessful initial attempts throughout Europe,
including: stocking in Denmark from sources in
Austria, France, USA, Canada, and New Zealand
(Welcomme 1988); in Israel from Switzerland
(Sklower 1951; Golani and Shefler 1985; Golani
and Mires 2000); in Cyprus from the USA (Lever
1996; Welcomme 1988); and in Slovakia and the
Czech Republic from Denmark. In the last case, it is
interesting to note that the stocking took place after
‘extirpation’ of wild populations by soldiers during
WWII (Lever 1996; Flajshans et al. 1999).
There are less well-documented records of ongoing
put-and-take rainbow trout introductions where the
sources are not identified. These include: England,
Wales, and Scotland in Great Britain (SNH 2003;EA
2004); Poland (Welcomme 1988; Winkler et al.
2000); Baltic tributaries of Estonia (Lever 1996;
FAO 1997; Anon 1999); the Netherlands (Raat 2003);
Finland (FAO 1997; Koli 1990; Winkler et al. 2000);
and notably in Austria where put-and-take stocking to
support recreational fisheries is ‘‘obligatory’’. Accord-
ing to Weiss and Kummer (1999):
Such stocking laws remain, in part, because
managers and fishermen believe that stocked
trout move or are washed downstream by high
flow events. Thus obligatory stocking ensures
that each party renting a river stretch contrib-
utes equally to the overall stock in the river.
In addition to these relatively well-documented
European introductions, there were also a host of
undated initial European transfers of rainbow trout
from unknown sources to Turkey (Kırkan et al.
2003), Albania (Holcı
´k1991; Welcomme 1988), the
Kola Peninsula in Russia (Lukin 1998), many differ-
ent tributaries in Spain, including River Ebro and
River Duero (de Sostoa and Lobon-Cervia 1989),
Sweden (Holmlund 2000), and formerly Soviet
waters of the Caspian Sea (Coad 1996a). Many
records of European rainbow trout introductions
simply indicate that the species was stocked, but
contained no other relevant information. These
include: Armenia (Gabrielyan 2001); Baltic tributar-
ies in Lithuania (Winkler et al. 2000); Serbia/Monte
Negro (Welcomme 1988); tributaries of Gulf of
Trieste, Slovenia (Marceta 1999); Turkey (Welcom-
me 1988; Coad 1996b); and Ukraine (Pavlov 1980).
Sockeye salmon (Oncorhynchus nerka)
There have only been a handful of unsuccessful
attempts to seed sockeye salmon from the USA to
Europe. These include introductions to Finland in the
early 1930s and again in the 1960s (FAO 1997;
Holcı
´k1991), to Sweden in the 1950s (Holcı
´k1991;
Kullander 1999), and to Denmark in the 1960s
(Fig. 8; Welcomme 1988; Holcik 1991).
Chinook salmon (Oncorhynchus tshawytscha)
European chinook salmon seed introductions date to
the earliest days, however these attempts were never
successful, and culturists soon abandoned their efforts
with this species (Fig. 9). During the period 1870–
1895, chinook salmon from the USA were seeded in
France (Holcı
´k1991; Keith et al. 1992; Billard 1997;
Keith and Allardi 1998) and the Netherlands (de
Groot 1985; Welcomme 1988). There are undated
reports of chinook salmon introductions from the
USA to England, Ireland, and Italy (Holcı
´k1991;
Welcomme 1988). Chinook salmon from Canada
were also unsuccessfully introduced to Germany, and
then from Germany back to Denmark (Scott and
Crossman 1973; Welcomme 1988; Holcı
´k1991;
Gerstmeier and Romig 1998; Monnerjahn 1999).
Oceania
Salmonines of the genus Onchorynchus are not
native to Oceania, but there is a long and well
Rev Fish Biol Fisheries
123

documented history of introductions within this
region (Table 1; Figs. 1–9). For major reviews on
the history, geopolitical distribution, and ecological
effects of introductions the reader is referred to: (1)
‘‘The origin of rainbow trout, Salmo gairdneri
Richardson, in New Zealand’’ (Scott et al. 1978);
(2) ‘‘Origin of New Zealand’s sockeye’’ (Hardy
1983); (3) ‘‘Origin of the New Zealand sockeye
salmon, Oncorhynchus nerka (Walbaum)’’ (Scott
1984); (4) ‘‘Salmon at the Antipodes’’ (Clements
1988); (5) ‘‘New Zealand freshwater fishes: a natural
history and guide’’ (McDowall 1990a); (6) ‘‘Game-
keepers for the nation: the story of New Zealand’s
acclimatisation societies, 1861–1990’’ (McDowall
1994a); (7) ‘‘Overview of the impacts of introduced
salmonids on Australian native fauna’’ (Cadwallader
1996); (8) ‘‘Impacts of introduced salmonids on
native galaxiids in New Zealand upland streams: a
new look at an old problem’’ (McDowall 2003); and
finally (9) ‘‘Crying wolf, crying foul, or crying
shame: alien salmonids and a biodiversity crisis in
the southern cool-temperate galaxioid fishes?’’
(McDowall 2006).
For all intents and purposes, stocking has ceased in
New Zealand except for some lakes where natural
reproduction is considered inadequate for harvest
demands (Bob McDowall, pers. comm.). Non-native
salmonines have been stocked into most Australian
waters thought to be suitable for them, and have now
established self-sustaining populations in many
waters, particularly in Tasmania and the cooler parts
of Victoria and New South Wales (Cadwallader
1996). The Australian states of Victoria, New South
Wales, Australian Capital Territory, South Australia,
Western Australia, Queensland, and to a lesser extent
Tasmania are responsible for conducting or licensing
ongoing stocking programs (Cadwallader 1996).
There have also been efforts to introduce salmonines
to numerous islands in the Southern Ocean, including
Iles Kerguelen, Ile de la Possession, Iles Crozet,
South Georgia, Marion Island, and the Prince Edward
Islands (Headland 1984; Watkins and Cooper 1986;
Davaine and Beall 1997; Ayllon et al. 2004a; Ayllon
et al. 2004b). Despite the long history of introduc-
tions in Oceania, golden trout, pink salmon, chum
salmon, and masu salmon were either not introduced
or documentation of such introductions outside of the
native range does not exist.
Cutthroat trout (Oncorhynchus clarki)
There is a single report of an unsuccessful attempt to
seed cutthroat trout in New Zealand during the period
1870–1890 (Fig. 2; McDowall 1990a).
Coho salmon (Oncorhynchus kisutch)
The only record of coho salmon seed introduction in
Oceania is the ongoing attempt to establish popula-
tions into the rivers of Marion, Crozet, and Kerguelen
Islands (French Austral Territories) in the southern
Indian Ocean (Davaine and Beall 1982; Cooper et al.
1992; Davaine and Beall 1997). Coho salmon
embryos continue to be introduced from the USA,
and then released into rivers of the Kerguelen
archipelago where wild progeny have also been
collected and alternately reared and released (Fig. 5;
Davaine 1991).
Rainbow trout (Oncorhynchus mykiss)
The origins and long history of rainbow trout in
Oceania have been well described (Roughley 1961;
Scott et al. 1978; Clements 1988). The first rainbow
trout were transported to New Zealand in 1877 from
the USA to the Acclimatization Society (to make
wildlife more familiar to European colonisers) who
seeded them in a lake of the volcanic plateau of the
central North Island (Fig. 7; Welcomme 1988;
McDowall 1994a; For detailed description of the
initial seed sources see Behnke 1972). In those days,
embryos were wrapped in moss on large blocks of ice
and carried by sailing clippers for a voyage that could
take several months (Aquatas 2004). Subsequent seed
introductions through the 1880s and 1890s took place
in the upper reaches of North Island and South Island
rivers and lakes, where they are now dominant
predators (McDowall 1990a; Swales 1990; Chadder-
ton et al. 2003). Currently, almost all of New
Zealand’s rainbow trout populations are self-sustain-
ing, and management stocking is undertaken only in
rare instances, such as the Waingongoro River and
the Rotorua Lakes (Bob McDowall, Rob Pitkethley,
Allen Stancliff pers. comm.).
In the 1890s, rainbow trout were successfully
introduced from the New Zealand stocks to most
Rev Fish Biol Fisheries
123

alpine and subalpine waters in Queensland, New
South Wales Victoria, South Australia, Western
Australia, and Tasmania (Roughley 1961; Fletcher
1986; Paxton et al. 1989; Kailola et al. 1993; Lever
1996; Arthington and McKenzie 1997). Ongoing
stocking programs are licensed by each of the
Australian States, and maintain put-and-take popu-
lations in warmer rivers/lakes/reservoirs, in
marginal habitats lacking suitable spawning condi-
tions, and where fishing pressure is considered too
high for local populations to sustain (Cadwallader
1996; Fulton 2004). The success of rainbow trout in
Australian waters has been attributed to the simi-
larity of Australian and ancestral habitats, and the
absence of large, coldwater predatory native fish
(Moyle and Light 1996; Molony 2001). Currently,
Australian rainbow trout are still considered to be
solely derived from the New Zealand source strains
of the early US introductions (Farrington et al.
2004).
In 1920, rainbow trout were introduced from
Oregon, USA to Hawaii, where self-sustaining
populations were subsequently established in Kauai.
Limited populations are also maintained by ‘put-
and-take’ stocking in some other high altitude
streams (Maciolek 1984; Yamamoto 1992; Kido
et al. 1999). In the 1950s, rainbow trout were
introduced from New Zealand to Papua New
Guinea (and possibly Tahiti) where they established
numerous self-sustaining populations in highland
reaches (West and Glucksman 1976; Allen 1991;
Eldredge 1994). As with coho salmon (see above),
there are also ongoing efforts to establish rainbow
trout into the rivers of Marion, Crozet, and
Kerguelen Islands in the southern Indian Ocean
(Davaine and Beall 1982; Cooper et al. 1992;
Davaine and Beall 1997).
Sockeye salmon (Oncorhynchus nerka)
The origins of New Zealand’s sockeye were
described by Hardy (1983) and Scott (1984). In
1901 and 1902 sockeye salmon were introduced from
Canada to various watersheds in New Zealand
(Fig. 8). However, these efforts only resulted in the
establishment of one self-sustaining population in
headwater lakes of the Waitaki River system of South
Island (McDowall 1984; Welcomme 1988; Paulin
et al. 1989; McDowall 1990a;1990b; Chadderton
et al. 2003).
Chinook salmon (Oncorhynchus tshawytscha)
Despite a great deal of interest and activity, the actual
source(s) of chinook salmon in Oceania remain
unclear, largely due to lack of documentation
(McDowall 1994b; Quinn et al. 1996). Beginning in
1876, chinook salmon were seeded from California,
USA to New Zealand, notably in coastal waters and
tributaries of eastern South Island, and some tribu-
taries along the west of South Island, and in North
Island (Fig. 9; McDowall 1984; McDowall 1990a;
McDowall 1994b; Chadderton et al. 2003). As an
interesting aside, it should be noted that chinook
salmon shipped from California in the same year
were introduced to Hawaii in an unsuccessful intro-
duction to the Kauai watershed (Maciolek 1984;
Welcomme 1988b).
Since the early 1920s, rivers along the east coast of
New Zealand’s South Island have represented the
only known anadromous wild populations of chinook
salmon in Oceania—many of which now show
significant population-specific phenotypic variation
(McDowall 1994b; Unwin and Glova 1997; Kinnison
et al. 1998; Unwin and James 1998). Beginning in the
1970s, chinook salmon became the focus of increas-
ing interest in commercial aquaculture in New
Zealand, and this has led to the development of
many hatcheries on the same tributaries (Crowe 1985;
McDowall 1990a; Unwin 1991).
In 1877, the year after they were introduced to New
Zealand, chinook salmon were introduced from the
New Zealand stocks and additional USA sources to
many rivers in Australia, but none of these introduc-
tions established self-sustaining populations
(MacKinnon 1987; Paxton et al. 1989; Cadwallader
1996). Beginning in the 1930s, chinook salmon were
introduced from New Zealand and the USA for
repeated releases, leading to the establishment of
fisheries in Lakes Purrumbete and Bullen Merri—two
deep, crater lakes which represent the only sites of
ongoing chinook salmon releases in Australia
(Butcher 1947; Barnham 1977; Cadwallader 1996).
Currently, there is no evidence that chinook salmon
have established self-sustaining populations anywhere
in their limited Australian range (Cadwallader 1996).
Rev Fish Biol Fisheries
123

Asia
Six Pacific salmonines are native to Asia; however,
with the exception of the rainbow trout, limited
attempts have apparently been made to seed the other
five Pacific salmonines to Asian watersheds. The
golden trout (O. aguabonita) and the cutthroat trout
(O. clarki) are not native to Asia and documentation
associated with introductions is scarce or incomplete
(Froese and Pauly 2004). Although the sockeye
salmon (Oncorhynchus nerka) is native to Asia,
where its distribution extends from Anadyr south
through the Bering Sea and Kamchatka region, the
northern Sea of Okhotsk as far south as Hokkaido
(Fig. 3; Popov 1933;Berg1948; Hanamura 1967;
Hart 1973), there are no documented introductions
outside of its native Asian range.
Pink salmon (Oncorhynchus gorbuscha)
Within Asia, pink salmon are distributed throughout
the Siberian Arctic to the Lena River, south along the
eastern Asian coast, Bering Sea, Anadyr, Kamchatka,
Sea of Okhotsk, North Korea, Hokkaido, and Hondo
Islands, Japan (Fig. 3; Berg 1948; Hart 1973). The
earliest introduction efforts were apparently made in
the 1930s to transfer pink salmon between unspec-
ified Arctic watersheds within the former USSR
(Holcı
´k1991; Welcomme 1988). Pink salmon were
unsuccessfully seeded during the 1960s from
unknown sources (perhaps USA and USSR) to
tributaries of Caspian Sea, notably the fresh waters
of Iran (Fig. 3; Coad 1995; Froese and Pauly 2004).
Chum salmon (Oncorhynchus keta)
Within Asia, chum salmon are native to the Lena
River, east to the Chukchi Sea, and south along the
Asian coast to Anadyr, Kamchatka, Sea of Okhotsk,
and Honshu, Japan (Fig. 4; Popov 1933; Berg 1948;
Hart 1973). In the 1960s, chum salmon were intro-
duced from an unknown source to the Caspian Sea and
particularly the freshwaters of Iran, where contested
rumours claim that it is still occasionally captured in
the wild (Coad 1995; Coad 1996a). Incomplete
records of attempted seed introductions of chum
salmon in Japan also exist (Molony et al. 2003).
Coho salmon (Oncorhynchus kisutch)
Within Asia, coho salmon are native to the Asian
coast south from Anadyr, throughout Kamchatka, the
northern Sea of Okhotsk, Kuril Islands, Hokkaido
Japan, northern Honshu, and Korea (Fig. 5; Berg
1948; Hart 1973). During the 1960s, coho salmon
were introduced with chum salmon from an unknown
source to the Caspian Sea and particularly the
freshwaters of Iran (Coad 1995;1996a).
Masu salmon (Oncorhynchus masou)
The cherry or masu salmon is the only Pacific
salmonine that is exclusively native to the western
Pacific. It ranges south from Kamchatka, through the
Sea of Okhotsk along the Russian mainland to the
Japan sea, including Hokkaido and Honshu (Fig. 6;
Berg 1948; Augerot 2004). During the decade 1975–
1985, there were several unsuccessful attempts to
seed masu salmon from its native distribution in
Japan to Nepal (Shrestha 1994) and Thailand (Wel-
comme 1988; Piyakarnchana 1989).
Rainbow trout (Oncorhynchus mykiss)
Within Asia, rainbow trout are native to the Kam-
chatka region (Fig. 7, Berg 1948). Behnke (1992)
suggested that the Russian forms may be a recent
derivative of costal rainbow trout that migrated from
North America during the late Pleistocene when the
Bering land bridge existed. As with most other
continents, the Asian introduction of Pacific salmo-
nines has been dominated by rainbow trout. In the
1870s and 1880s and again in the 1920s, rainbow
trout were introduced from the USA to Japan with
wild reproduction confirmed in Hokkaido; however,
it was not until the 1970s when seed stocking resulted
in a significant increase in population range and
associated recreational fisheries (Welcomme 1988;
Chiba et al. 1989; Takami and Aoyama 1999; Pietsch
et al. 2000; Hasegawa and Maekawa 2006). Early
successful rainbow trout introductions also included:
from Great Britain to Sri Lanka (MacCrimmon 1971;
Pethiyagoda 1991); and from Sri Lanka, Great
Britain, Germany, and New Zealand to India (Mac-
Crimmon 1971; Welcomme 1988; Shetty et al. 1989;
Rev Fish Biol Fisheries
123

Talwar and Jhingran 1991; Gopi 2000; Shaji et al.
2000).
The 1920s saw introductions of rainbow trout from
the Japanese stock and USA sources to northern
tributaries in India and Pakistan, especially Azad
Jammu, and Kashmir. These introduction efforts were
repeated in the 1970 and 1980s leading to the
widespread establishment of rainbow trout in streams
and reservoirs above 1370 m altitude (MacCrimmon
1971; Akhtar 1991a,b; Gopi 2000; Mirza 2003).
During the 1930s, there were introductions of rain-
bow trout from New Zealand, Scotland, and England
to Malaysia for recreational fisheries in mountain
streams. The Malaysian introduction attempt was
repeated in the late 1960’s, but was terminated due to
the lack of economic return (Welcomme 1988; Ang
et al. 1989).
After the Second World War, an intensive new
round of rainbow trout introductions occurred
throughout Asia. In the late 1950s, rainbow trout
were introduced from Japan to Taiwan primarily for
aquaculture, but with establishment of some self-
sustaining, wild populations (Welcomme 1988; Liao
and Lia 1989; Liao et al. 2001). Shortly afterward,
rainbow trout were introduced from Korea (unre-
ported origin, probably from the USA) to China
primarily for aquaculture, but with resulting estab-
lishment of self-sustaining populations in more than a
dozen Chinese provinces (Chyung 1977; Welcomme
1988; Tan and Tong 1989; Walker and Yang 1999;
Xie et al. 2001). During the 1960s, rainbow trout
were introduced from an unknown source to the
Salang/Panjsher watersheds of Afghanistan and
northward flowing tributaries in the Hindu Kush
region for aquaculture and recreational fisheries
(Coad 1981; Welcomme 1988). There are reports
that rainbow trout were introduced from various
European sources, including Denmark, to many
Iranian watersheds, notably the Tigris River (Coad
and Abdoli 1993; Coad 1995; Coad 1996b). Rainbow
trout from Czechoslovakia were introduced to
Kazakhstan (especially the Irtysh watershed), leading
to a commercial fishery in Lake Balkhash by 1967
(Mitrofanov and Petr 1999). During the early 1970s,
rainbow trout were unsuccessfully introduced for
aquaculture from India to the Godwari River in
Nepal, later from Japan to Nepal (FAO 1997;
MacCrimmon 1971; Shrestha 1994), and from Can-
ada to the highlands of northern Thailand for food
fisheries (Piyakarnchana 1989; Welcomme 1988).
Finally, rainbow trout were introduced for aquacul-
ture or food from undocumented sources to Chirchik
River and Charvak Reservoir of Uzbekistan (Kamilov
and Urchinov 1995; Khurshut 2001) and Lake Issyk-
kul in Kyrgyzstan (Savvaitova and Petr 1999).
Chinook salmon (Oncorhynchus tshawytscha)
Within Asia, chinook salmon are native to the Anadyr
River, Kamchatka, Maritime territories, Sea of Okh-
otsk, Amur River, and are found as far south as
Hokkaido Japan (Fig. 9; Berg 1948; Hart 1973).
Unsuccessful seed introductions of chinook salmon
from the USA and Canada were attempted in Japan
during the periods 1880–1905 and 1960–1970 (Wel-
comme 1988; Chiba et al. 1989), and from various
sources in the former Soviet Union to new watersheds
elsewhere in the Union (Morrow 1980; Reshetnikov
et al. 1997; Pietsch et al. 2000; Bogutskaya and
Naseka 2002). The Japanese broodstock of chinook
salmon also served as a donor for failed seed
introduction to the Nilgiri watershed and other Indian
tributaries (Talwar and Jhingran 1991; FAO 1997;
Kapoor et al. 2002).
Africa
Pacific salmonines are not native to the African
Continent (Table 1; Figs. 1–9); however, the vast
majority of the African experience with introduced
salmonines has focussed on rainbow trout. General
reviews of Pacific salmonine introductions to Africa
are included in literature published by de Moor and
Bruton (1988) and Cambray (2002;2003). Golden
trout (Oncorhynchus aguabonita), pink salmon (O.
gorbuscha), chum salmon (O. keta), coho salmon (O.
kisutch), masu salmon (O. masou), and sockeye
salmon (O. nerka) were either not introduced to
Africa, or records of introductions do not exist or
were not readily available.
Cuthroat trout (Oncorhynchus clarki)
Only a single record of cutthroat trout being intro-
duced from the USA to Morocco existed, where they
Rev Fish Biol Fisheries
123

reportedly hybridized with rainbow trout (Mouslih
1987; Welcomme 1988).
Rainbow trout (Oncorhynchus mykiss)
Rainbow trout were introduced in the late 1890s from
Great Britain to South Africa for recreational fisher-
ies; they survived in the colder tributaries of the south
coast and the Olifants system and were implicated in
the decline of a number of local species (Fig. 7;
Skelton 1987; de Moor and Bruton 1988; Welcomme
1988). During the period 1905–1910, South Africa
exported rainbow trout for the establishment of
recreational fisheries in Kenya, Swaziland, and
Zimbabwe. The Kenyan introductions included the
Athi-Galana-Sabaki watershed and other mountain
streams (Welcomme 1988; FAO 1993; Okeyo 1998).
The only surviving Swaziland introductions are the
Komati and the Usutu in the highveld of the country;
the Malolotja Reserve, as well as stocked populations
in dams, such as the Forbes Reef Dam (Clay 1976;
Welcomme 1988; Skelton 1993). The Zimbabwe
introductions occurred in cooler upland tributaries of
the Lower Zambezi and Pungwe River systems;
current stocking of rainbow trout persists in Haroni
River and the Buzi River system (Bell-Cross and
Minshull 1988; Welcomme 1988). Great Britain also
exported rainbow trout for repeated introduction
attempts in Malawi over a period from 1906 to
1960, ultimately leading to establishment of self-
sustaining populations for recreational fisheries (FAO
1997; Welcomme 1988).
During the 1920s and 1930s, a variety of seed
introductions established self-sustaining populations
of rainbow trout, including transfers from France to
Madagascar then to Reunion (Welcomme 1988;
Stiassny and Raminosoa 1994; Fricke 1999; Keith
et al. 1999), from France and Switzerland to moun-
tain streams in Morocco (Welcomme 1988; Azeroual
et al. 2000), and from South Africa to Mauritius
(FAO 1997; Welcomme 1988). Shortly after, rainbow
trout were also seeded from Scotland to Tanzania,
where they evidently established self-sustaining pop-
ulations in cool highland streams of the Mbeya and
Moshi areas (Welcomme 1988; Eccles 1992).
The period from 1940–1950 was largely charac-
terized by unsuccessful introductions of rainbow trout
among African nations, including: South Africa to
Congo (FAO 1997); South Africa to Zambia (Thys
van den Audenaerde 1994); and Kenya to Sudan
(Welcomme 1988). Also during this time, rainbow
trout were reportedly introduced from Italy to Lake
Aba Samuel of Ethiopia. Ethiopian tributaries (Lake
Wonchi, the Sibilo, Chacha, Danka and Beressa
Rivers) were re-seeded with rainbow trout in the late
1960s and 1970s by UNESCO and others. These
efforts evidently led to the establishment of wild
populations (Tedla and Meskel 1981; Welcomme
1988). Rainbow trout were also introduced in 1967
from France to Tunisia for aquaculture and recrea-
tional fisheries (FAO 1997). Other successful
introductions of rainbow trout to Africa include:
South Africa; Ethiopia (at least to 1970); Malawi;
Zimbabwe (then Rhodesia); Sudan; Tanzania;
Uganda; Madagascar and La Re
´union (MacCrimmon
1971). MacCrimmon (1971) and Welcomme (1988)
also reported a rather long list of undated rainbow
trout introductions, as follows: South Africa-Lesotho;
Great Britain-Guyana; Germany-Cameroon; Zimba-
bwe-Mozambique; Lesotho; Cameroon; Congo;
Mozambique; Zambia; Mauritius; Eritrea; and
Uganda. Finally, ongoing put-and-take rainbow trout
stocking introductions in Africa include: South
Africa; Kenya; Malawi; Morocco; Zimbabwe; Sudan;
Tanzania; Tunisia; Uganda; Zambia; and the Azores
(MacCrimmon 1971).
Chinook salmon (Oncorhynchus tshawytscha)
The only record we were able to find was a report of a
1955 introduction of chinook salmon to Madagascar
from an unknown source (Stiassny and Raminosoa
1994).
South America
None of the Pacific salmonines considered in this
review are native to South America (Figs. 1–9).
Pacific salmonines introductions in South America
have primarily been limited to Argentina and Chile.
Introductions of golden trout (O. aguabonita), cut-
throat trout (O. clarki), chum salmon (O. keta), coho
salmon (O. kisutch), and masu salmon (O. masou)
were either not undertaken in South America or were
poorly documented.
Rev Fish Biol Fisheries
123

Pink salmon (Oncorhynchus gorbuscha)
There are only two undocumented records for seed
introductions of pink salmon from the USA to South
America: Argentina and the southern waters of Chile
(Fig. 3; Welcomme 1988; Lever 1996). It is pre-
sumed that neither of these attempts was successful.
Coho salmon (Oncorhynchus kisutch)
In the first decade of 1900, coho salmon were seeded
from the USA to Argentina and Chile to ‘‘fill
[undefined] ecological niches’’ Fig. 5; Welcomme
1988; Lever 1996; Soto et al. 2001). More recently,
coho salmon from California (and possibly Ontario,
Canada) have been the subject of renewed aquacul-
ture and fisheries interests at Chiloe Island and in the
Lake Llanqihue region of Chile, where escapees have
evidently become established in the wild (Lever
1996;Pe
´rez et al. 2001). In 1994/95, heavy storms
destroyed farm pens in southern Chile, releasing
millions of coho salmon with yet undetermined status
of wild reproduction and population maintenance
(Soto et al. 2001). To date, coho salmon populations
in Argentina have not shown evidence of self-
sustaining reproduction, and appear to be maintained
by ongoing stocking programs and/or accidental
releases from aquaculture operations.
Masu salmon (Oncorhynchus masou)
A single record of masu introductions to South
America exists (Fig. 6). Citing Nagasawa and Aguil-
era 1974, Machidori and Kato (1984) reported that
approximately 85, 000 juvenile masu salmon were
released in the Clara River, a tributary of the Simpson
River in Aisen province of Chile in 1973. It is likely
that these seed introductions were unsuccessful,
because no recovery of adult fish was ever recorded.
Rainbow trout (Oncorhynchus mykiss)
In 1904, rainbow trout were introduced from the USA
to Argentina for aquaculture and recreational fisheries
(Welcomme 1988; McDowall et al. 2001; Pascual
et al. 2002). During the period 1930–1960, rainbow
trout were also introduced to Argentina from France,
Germany, and especially Denmark (Riva Rossi et al.
2004). Rainbow trout eventually became a dominant
self-sustaining species in the Patagonian watersheds
(Pascual et al. 2001; Pascual et al. 2002; Riva Rossi
et al. 2004). Rainbow trout in the Santa Cruz River of
Patagonia have shown evidence of a migratory life
history with both marine (anadromous) and freshwa-
ter (non-anadromous) forms (Behnke 2002a; Riva
Rossi et al. 2004).
In 1905, rainbow trout were introduced from
Germany to Chile, where self-sustaining populations
were established in southern watersheds (Welcomme
1988). Rainbow trout were also introduced from
Great Britain to the southern highlands of Brazil in
1913, with repeated, and apparently successful,
attempts made in the 1940s and 1950s (Welcomme
1988). In the 1960s, rainbow trout seed introductions
were promoted and maintained by US-Chilean fish-
eries partnerships (Davidson and Hutchinson 1938;
Soto et al. 2001). As was described for coho salmon,
heavy storms in the mid-1990s resulted in the release
of millions of rainbow trout, with yet undetermined
status of wild reproduction (Soto et al. 2001).
Currently, put-and-take stocking in Chile also sup-
plements unknown levels of wild reproduction by
rainbow trout to support large recreational fisheries
(Soto et al. 2001).
The 1920s witnessed an increase in rainbow trout
introductions to South America with fish from the
USA shipped to Columbia and Ecuador for aquacul-
ture—the latter introduction was repeated in the
1960s, apparently with establishment of wild popu-
lations (MacCrimmon 1971; Welcomme 1988). In
1927, rainbow trout were also introduced from the
USA to Lake Titicaca, Peru where it formed the basis
of a large commercial fishery that collapsed due to
overexploitation (Welcomme 1988). During the
period 1930–1960, rainbow trout were introduced
from the USA to Venezuela, and reintroduced from
the USA and Chile to Bolivian waters of Lake
Titicaca (FAO 1997; MacCrimmon 1971; Welcomme
1988). In the 1950s and 1960s, rainbow trout from
Denmark and the USA were introduced to the Rio de
Janeiro in Brazil, where self-sustaining populations
were established (MacCrimmon 1971). Rainbow trout
were also unsuccessfully introduced from Argentina
to the Falkland Islands, Trinidad, Uruguay, Guyana
and Paraguay (MacCrimmon 1971; Lever 1996).
Rev Fish Biol Fisheries
123

Sockeye salmon (Oncorhynchus nerka)
During the first decade of 1900, sockeye salmon were
evidently introduced from the USA along with
chinook salmon (see below) to Argentina to ‘‘fill
[another] undefined niche’’ (Fig. 8; Welcomme 1988;
Pascual et al. 2002).
Chinook salmon (Oncorhynchus tshawytscha)
The largely undocumented history of chinook salmon
in South America has its origins with most of the
other introduced salmonines—at the turn of the
previous century. Chinook salmon from the USA
were first introduced to Argentina and Chile between
1900 and 1910 (FAO 1993; Welcomme 1988).
Subsequent stocking and escapes from salmon ranch-
ing operations have taken place in these countries,
and it appears that in a manner similar to that
observed for rainbow trout, wild, self-sustaining
populations may have been established (Zamorano
1991; Pascual et al. 2001; Soto et al. 2001).
Discussion and conclusions
Synthesizing the available literature and providing a
quantitative historical synopsis of global Pacific
salmonine introductions has proven to be a challeng-
ing task. Based on the complexity of the accounts
presented above, it would be fair to say that such a
broad goal could only be humbly satisfied. There are
several factors which generally limited our ability to
provide a comprehensive and quantitative review of
historical global salmonine introductions. First and
foremost, while detailed records of introductions
existed for North America, Oceania and western
Europe, they were rarely in an electronic form that
was readily accessible. Elsewhere in the world, hard
data on any aspect of salmonine introductions was a
rare commodity indeed. As a consequence of these
limitations, we could not really offer the kind of
quantitative measures of introduction that we sought,
but rather we had to strive for reasonably meaningful,
qualitative descriptions. With this caveat in mind,
there are several important observations and trends
that emerged from our review of the available
literature, which we summarize and discuss in the
following paragraphs.
Currently, the United Nations (UN 2007) lists 192
countries worldwide: 23 in North America; 43 in
Europe; 14 in Oceania; 53 in Africa; 47 in Asia; and
12 in South America. Based on our review, Pacific
salmonines have been introduced to approximately
65% of these countries. Pacific salmonines are native
to approximately 13 of the worlds’ major watersheds
(WRI 2003), but through a combination of intentional
and accidental introductions, and dispersal from seed-
point-sources, have become exotic species in nearly
all of the remaining 93 major watersheds in the
world. The incredible success of these introductions–
or failure depending on which side of the coin one is
looking at–can be attributed to a complex combina-
tion of social, cultural, economic, biological, and
ecological factors.
Of the Oncorhynchus species reviewed, the rain-
bow trout (O. mykiss) is far-and-away the most
widely introduced and the most successful in terms of
establishing self-sustaining populations outside of its
native range. The global success of the rainbow trout
can be ascribed to a combination of its social
importance as a recreational game-fish in western
culture, its importance as a food-fish in south-Asian
culture, the persistence of initial seed efforts, manip-
ulation of life history traits through selective
breeding, and the ongoing maintenance of artificial
populations through hatchery cultivation. However,
the plasticity of the salmonine life history in general,
and their physiology and behaviour in particular,
make them highly adaptable and have facilitated their
current global distribution.
In general, we recognize three primary factors
influencing the natural geographic distribution of
fishes: (1) physical opportunity; (2) constitutional
opportunity; and (3) ecological opportunity. For
fishes to colonize new habitats, the physical oppor-
tunity (i.e., connections between water-bodies) must
be present and navigable for successful dispersal to
new environments. Constitutional opportunity refers
to the physiological ability of the individual fish to
withstand the chemical, physical, and biological
environments through which it disperses and in
which it persists. Finally, ecological opportunity
refers to the availability of an ecological opportunity
for survival. While we acknowledge difficulties in the
concept of the pre-Hutchinsonian (1957) ‘vacant
niche’, many of the early salmonine introductions
were explicitly justified on this basis. More
Rev Fish Biol Fisheries
123

specifically, ecological opportunity refers to the
availability of sufficient spawning, nursery and
feeding habitat as well as adequate trophic resources
within the receiving water-body.
In the case of intentionally introduced Oncorhyn-
chus salmonines, geographic barriers to dispersal are
not an issue as they are mitigated by anthropogenic
means (i.e., transportation infrastructures). Similarly,
constitutional opportunity has been taken care of with
the development of modern hatchery facilities and
stocking practices. However, the expressed pheno-
typic physiology of the fish remains the primary
factor in determining the introduced fish’s ability to
survive and reproduce in a receiving water-body. For
example, the broad geographic distribution of rain-
bow trout suggests that they are extremely tolerant
and highly adaptable to new environments. The
adaptive nature of rainbow trout has also been
augmented through selective breeding of hatchery
strains. Anadromy, the life history trait of migrating
from salt water into freshwater to spawn (see
McDowall 1999 for refined definitions), also provides
Pacific salmonines with enhanced constitutional
opportunity to disperse from seeding sites via oceanic
migration (McDowall 1999). Pacific salmonines also
have the plasticity to exhibit potamodromous pheno-
types, which complete their life-cycle entirely within
freshwater, migrating to rivers or suitable near-shore
freshwater habitat for reproduction. Therefore, life
history and physiological plasticity predisposes On-
corhynchus salmonines for dispersal from seeding
point-sources, and provides them with greater con-
stitutional and ecological opportunity to succeed in
new habitats.
Another general observation that was apparent in
our review was that the type of introduction (i.e., seed
versus put-and-take) was determined and maintained
by logistic and economic factors. Not surprisingly,
the original intent of stocking was often driven by
social and cultural ideals (Johnson et al. 1995). With
the exception of southeast Asia, most of the global
salmonine introductions were undertaken with the
explicit intent of developing recreational fisheries
rather than providing human sustenance. To meet
angler demand, the inertia of recreational stocking
almost always led to the development of put-and-take
fisheries that were maintained through intensive
hatchery-based stocking efforts (MacCrimmon
1971; McDowall 1984; Crossman 1991; Coad and
Abdoli 1993; Mills et al. 1993; Coad 1996a,b;
Crawford 2001). While these social values certainly
played an important role in the nearly ubiquitous
distribution of Pacific salmonines outside of their
native range, attitudes are changing and a more
globally- and ecologically-minded management ethic
is definitely emerging. Changing management ethics
may have influenced the global decline in the
frequency of salmonine seed introductions. Perhaps
the simplest hypothesis to account for this phenom-
enon would be that humans have already attempted to
introduce all of the candidate salmonines into all of
the candidate watersheds, and these seedings were
either (a) successful, (b) unsuccessful and the propo-
nents gave up, or (c) unsuccessful and the proponents
converted the program to ongoing put-and-take
stocking.
MacCrimmon (1971) identified water temperature
and precipitation as the two most important environ-
mental constraints that could define the natural limits
of rainbow trout population establishment and main-
tenance, and ultimately the success of salmonine seed
or put-and-take introductions. As we grapple with
global climate change in the new millennium, we
should consider that constraints to salmonine intro-
ductions may change markedly; habitats that
supported previously successful introductions may
no longer support populations, while habitats that
were previously inhospitable for salmonines may
now allow for wild colonization, or even a new round
of seed introductions. From a global perspective, we
suspect that the most significant environmental
changes favouring salmonines (native or introduced)
will be detected in sub-Arctic tributaries such as
those in northern Canada, Alaska, Russia and Scan-
dinavia (Babaluk et al. 2000; Stephenson 2005).
As previously stated, we did not attempt to explore
the complex patterns in ecological effects, changing
stakeholder values, or the role of ecosystem-based
management in shaping policies on introductions of
exotics species. That being said, we would be remiss
in failing to acknowledge some of the major ecolog-
ical issues that are currently threatening global
ecosystems as a consequence of Pacific salmonine
introductions. Indeed, the rainbow trout is currently
considered one of the 100 ‘worst global invasive
species’ (ISSG 2007).
There can be no doubt that one of the most
important developments in the story of global
Rev Fish Biol Fisheries
123

salmonine introductions has been the expansion of
aquaculture over the past three decades. The scale of
commercial production and the accidental releases
that are always associated with salmon farming,
combine to form an unintentional but dominant factor
that affects global salmonine distribution in the wild.
The hypothesized ecological effects of such uninten-
tional introductions (especially for transgenic fish)
are hotly debated by proponents, opponents, scien-
tists, managers, and the public. These debates will
continue to escalate as the global demand for food
production increases, and the longer-term ecological
consequences of introduced salmonines become more
apparent. As alluded to previously, one factor that has
the potential to dramatically change the future of
salmonine introductions by farm escapes is the
maturation of an ecological ethic in fisheries man-
agement that explicitly recognizes the consequences
of introducing an exotic species into an ecosystem
that evolved in the absence of that species. From our
perspective, the key uncertainty regarding the future
of global salmonine introductions will be whether the
economic incentives of commercial aquaculture and
artificial recreational fisheries will outweigh the value
that humans place on any degree of ‘naturalness’ in
the aquatic ecosystems that support them. At the rate
things are changing, the next twenty years should
reveal which of these forces will prevail.
Finally, and perhaps most importantly, we found a
host of strong and repeated warnings about the
serious ecological dangers associated with salmonine
introductions. In our opinion, McDowall (2006)
presents the single most rigorous, scholarly and
passionate example of the dangers posed by salmo-
nine introductions in his review: ‘‘Crying wolf,
crying foul, or crying shame: alien salmonids and a
biodiversity crisis in the southern cool-temperate
galaxioid fishes?’’ McDowall argues that enduring
and pervasive information clearly demonstrates that
survival of the entire group of galaxoid species has
been seriously threatened by intentionally introduced
salmonine fishes. From a global perspective, we are
not likely to receive, nor should we expect, any better
warnings of the clear and present ecological danger
posed by introductions of non-native salmonines. For
those with an appreciation of ecological risk, there is
no reason to wait—ongoing stocking of non-native
salmonines should be terminated. If we do not
change, then we fully deserve to answer our children
and grandchildren when they will ask us the painful
question: ‘‘Why didn’t you do something when it was
possible to make a difference and when you knew
what was happening?’’
Acknowledgements This general review builds on a long
series of previous historical and ecological/socio-economic
reviews of introduced fishes in general, and introduced
salmonines in particular (MacCrimmon and Marshall 1968;
MacCrimmon 1971; McDowall 1984; Welcomme 1988;
Holcı
´k1991; McDowall 1994a,b; Lever 1996; Cowx 1999;
Crawford 2001). Internet sources proved to be especially
valuable in piecing together the diverse and often complex
history of global salmonine introductions. We would like to
especially recognize the primary and technical literature that
was cited by Fishbase the Nonindigenous Aquatic Species
(NAS) information resource (USGS 2004). Finally, we
gratefully acknowledge the host of international colleagues,
too numerous to thank individually, who responded to our
requests for information on salmonine introductions in their
region. One person who deserves special recognition is Dr.
Juraj Holc
ˇı
´k (Slovak Academy of Sciences, Bratislava,
Slovakia) who was kind enough to introduce us to many of
his global contacts with special insight on this issue.
References
Adams SB, Frissell CA, Rieman BE (2001) Geography of
invasion in mountain streams: consequences of headwater
lake fish introductions. Ecosystems 4:296–307
Akhtar N (1991a) The Northern areas (Pakistan). Fisheries
profile, feasible sites for trout culture and an overall sec-
toral development perspective. Report for Project PAK/
91/008. FAO, Rome, 29 pp
Akhtar N (1991b) Azad Jammu and Kashmir. Fisheries profile,
feasible sites for trout culture and an overall sectoral
development perspective. Report for Project PAK/88/048.
FAO, Rome, 25 pp
Allen GR (1991) Field guide to the freshwater fishes of New
Guinea. Christensen Research Institute, Madang, 268 pp
Ang KJ, Gopinath R, Chua TE (1989) The status of introduced
fish species in Malaysia, exotic aquatic organisms in Asia:
introduction of exotic aquatic organisms in Asia Spec
Publ 3, Manila, Philippines, Asian Fisheries Society, p 71
Anon (1999) Systematic list of Estonian fishes. World Wide
Web Electronic Publication, 14 January 2000. http://
www.bio.edu.ee/animals/Kalad/kalalist2.htm Accessed 26
October 2007
Appleby C (1999) List of Norwegian common names of fishes.
Unpublished manuscript – cited in Fishbase
Aquatas (2004) Tasmanian salmon industry – history of the
salmon industry. http://www.aquatas.com/industry.html.
Accessed 28 November 2004
Aro KV, Shepard MP (1967) Salmon of the north Pacific
Ocean: Part IV. Spawning populations of north Pacific
salmon. Int N Pac Fish Comm Bull 23:225–327
Arthington AH, McKenzie F (1997) Review of impacts of
displaced/introduced fauna associated with inland waters.
Rev Fish Biol Fisheries
123

Australlia: State of the environment technical paper series.
Department of the Environment, Canberra, 69 pp
Atkinson EE, Rose JH, Duncan TO (1967) Salmon of the north
Pacific Ocean, Part IV. Spawning populations of north
Pacific salmon. Int N Pac Fish Comm Bull 23:43–223
Augerot X (2004) Salmon stocks and habitat in the Russian far
east. In: Gallaugher P, Wood L (eds) Proceedings of the
world summit on salmon, Simon Fraser University, Brit-
ish Columbia, p 77
Ayllon F, Davaine P, Beall E, et al (2004a) Bottlenecks and
genetic changes in Atlantic salmon (Salmo salar L.)
stocks introduced in the subantarctic Kerguelen Islands.
Aquaculture 237:103–116
Ayllon F, Martinez JL, Davaine P et al (2004b) Interspecific
hybridization between Atlantic salmon and brown trout
introduced in the subantarctic Kerguelen Islands. Aqua-
culture 230:81–88
Azeroual A, Crivelli AJ, Yahyaoui A et al (2000) L’ichtyo-
faune des eaux continentales du Maroc. Cybium
24(Suppl. 3):17–22
Babaluk JA, Reist JD, Johnson JD et al (2000) First records of
sockeye (Oncorhynchus nerka) and pink salmon (O.
gorbuscha) from Banks Island and other records of Pacific
salmon in the Northwest Territories, Canada. Arctic
53:161–164
Bakshtansky EL (1980) The introduction of pink salmon into
the Kola Peninsula. In: Thorpe JE (ed) Salmon ranching.
Academic Press, London, p 245
Barnes ME, Whelan GE (2004) Visitor policies and practices at
public trout and salmon hatcheries in North America.
Fisheries 29:16–21
Barnham CA (1977) Ten-year effort to acclimatise quinnat
salmon pays off. Aust Fish 36:14–15
Behnke RJ (1972) The systematics of Salmonid fishes of
recently glaciated lakes. J Fish Res Board Can 29:639–671
Behnke RJ (1992) Native trout of western North America.
American Fisheries Society Monograph 6, Bethesda,
275 pp
Behnke RJ (2002a) Comment: first documented case of
anadromy in a population of introduced rainbow trout in
Patagonia, Argentina. Trans Am Fish Soc 131:582–585
Behnke RJ (2002b) Trout and salmon of North America.
Simon and Schuster, New York
Bell-Cross G, Minshull JL (1988) The fishes of Zimbabwe.
Copeia 1989:809–810
Berends M (2004) Identification by scales of hatchery and wild
chinook salmon (Oncorhynchus tshawytscha) in the
Credit River, Ontario. Ph.D. Thesis, University of
Toronto, Ontario
Berg LS (1948) Freshwater fishes of the U.S.S.R. and adjacent
countries, vol I. Academy of Sciences of the U.S.S.R.
(Translation by Israel Program for Scientific Translations,
1962), 493 pp
Berg M (1977) Pink salmon, Oncorhynchus gorbuscha (Wal-
baum) in Norway. Report of the Institute for Freshwater
Research, Drottningholm 56:12–17
Bianco PG, Ketmaier V (2001) Anthropogenic changes in the
freshwater fish fauna of Italy, with reference to the central
region and Barbus graellsii, a newly established alien
species of Iberian origin. J Fish Biol 59(Suppl.A):
190–208
Billard R (1997) Les poissons d’eau douce des rivie
`res de
France. Identification, inventaire et re
´partition des 83 es-
pe
`ces. Delachaux & Niestle
´, Lausanne, 192 pp
Blanchet S, Loot H, Grenouillet G, Brosse S (2007) Compet-
itive interactions between native and exotic salmonids: a
combined field and laboratory demonstration. Ecol
Freshw Fish 16:133–143
Bogutskaya NG, Naseka AM (2002) An overview of nonin-
digenous fishes in inland waters of Russia.
http://www.zin.ru/animalia/pisces/eng/taxbase_e/intro_e/
intro_e.htm. Accessed 26 October 2007
Bradford DF, Tabatabai F, Graber DM (1993) Isolation of
remaining populations of the native frog, Rana muscosa,
by introduced fishes in Sequoia and Kings Canyon
National Parks, California. Conserv Biol 7:882–888
Bradford DF, Cooper SD, Jenkins TM et al (1998) Influences
of natural acidity and introduced fish on faunal assem-
blages in California alpine lakes. Can J Fish Aquat Sci
55:2478–2491
Burgner RL (1991) Life history of sockeye salmon (On-
corhynchus nerka). In: Groot C, Margolis L (eds) Pacific
salmon life histories. UBC Press, Vancouver
Bussing WA (1998) Peces de las aguas continentales de Costa
Rica [Freshwater fishes of Costa Rica], 2nd edn. Editorial
de la Universidad de Costa Rica, San Jose
´, 271 pp
Butcher AD (1947) Quinnat salmon in Victorian inland waters.
Fisheries and Game Department, Fisheries Pamphlet
No. 4, Victoria, 28 pp
Cadwallader PL (1996) Overview of the impacts of introduced
salmonids on Australian native fauna. Report prepared for
the Australian Nature Conservation Agency, Canberra,
68 pp
Cambray JA (2002) The global impact of alien trout species – a
review; with reference to their impact in South Africa. Afr
J Aquat Sci 28:61–67
Cambray JA (2003) Impact on indigenous species biodiversity
caused by the globalisation of alien recreational fresh-
water fisheries. Hydrobiologia 500:217–230
Chadderton WL, Grainger N, Dean T (2003) Proceedings of a
workshop hosted by Department of Conservation:
appendix 1 – prioritising control of invasive freshwater
fish, Managing invasive freshwater fish in New Zealand,
Hamilton, 10–12 May 2001, New Zealand Department of
Conservation, p 171
Chiba K, Taki Y, Sakai K et al (1989) Present status of aquatic
organisms introduced into Japan. Exotic aquatic organ-
isms in Asia proceedings of the workshop on introduction
of exotic aquatic organisms in Asia Spec Publ 3, Manila,
Philippines, Asian Fisheries Society, p 63
Christie WJ (1970) Introduction of the cherry salmon On-
corhynchus masou in Algonquin Park, Ontario. Copeia
1970:378–379
Chyung MK (1977) The fishes of Korea. Il Ji Sa Publishing Co,
Seoul, 727 pp
Clay D (1976) An investigation into the distribution of fish in
Swaziland. Rev Zool Afr 90:547–558
Clements J (1988) Salmon at the antipodes: a history and
review of trout, salmon and char and introduced coarse
fish in Australasia. Skipton Eels, 391pp
Coad BW (1981) Fishes of Afghanistan, an annotated check-
list. Publ Zool Natl Mus Can 14:23
Rev Fish Biol Fisheries
123

Coad BW (1995) Freshwater fishes of Iran. Acta Sci Nat Acad
Sci Brno 29:1–64
Coad BW (1996a) Exotic and transplanted fishes in southwest
Asia. Publ Espec Inst Esp Oceanogr 21:81–106
Coad BW (1996b) Exotic fish species in the Tigris-Euphrates
basin. Zool Middle East 13:71–83
Coad BW, Abdoli A (1993) Exotic fish species in freshwater
fishes of Iran. Zool Middle East 9:65–80
Collins JJ (1971) Introduction of kokanee salmon (Oncorhyn-
chus nerka) into Lake Huron. J Fish Res Board Can
28:1857–1871
Collins JJ (1975) Occurrence of pink salmon (Oncorhynchus
gorbuscha) in Lake Huron. J Fish Res Board Can 32:402–
404
Cooper J, Crafford JE, Hecht T (1992) Introduction and
extinction of brown trout (Salmo trutta L.) in an impov-
erished Subantarctic stream. Antarct Sci 4:9–14
Courtenay WR Jr, Hensley DA, Taylor JN et al (1984) Dis-
tribution of exotic fishes in the continental United States.
In: Courtenay WR, Stauffer JR (eds) Distribution, biology
and management of exotic fishes. Johns Hopkins Uni-
versity Press, Maryland, p 41
Cowx IG (1999) An appraisal of stocking strategies in the light of
developing country constraints. Fish Manag Ecol 6:21–34
Crawford SS (2001) Salmonine introductions to the Laurentian
Great Lakes: an historical review and evaluation of eco-
logical effects. Can Spec Publ Fish Aquat Sci 132:205
Crivelli AJ (1995) Are fish introductions a threat to endemic
freshwater fishes in the northern Mediterranean region?
Biol Conserv 72: 311–319
Crossman EJ (1968) Changes in the Canadian freshwater fish
fauna. In: Loftus KH (ed) A symposium on introductions
of exotic species. Ontario Department of Lands and For-
ests, Research Report (Fisheries) 82:1–20
Crossman EJ (1984) Introduction of exotic fishes into Canada.
In: Courtenay WR Jr, Stauffer JR Jr (eds) Distribution,
biology and management of exotic fishes. Johns Hopkins
University Press, Baltimore, p 78
Crossman EJ (1991) Introduced freshwater fishes: a review of
the North American perspective with emphasis on Can-
ada. Can J Fish Aquat Sci 48(Suppl. 1):46–57
Crossman EJ, Cudmore BC (1999) Summary of fishes inten-
tionally introduced in North America. In: Claudi R, Leach
JH (eds) Nonindigenous freshwater organisms. Lewis
Publishers, Louisiana, p 99
Crowe AW (1985) The N.Z. Salmon Co. ocean ranching
venture. In: Taylor JL, Ogilvie RM, Todd PR (eds) Pro-
ceedings of the salmon farming conference, New Zealand
Ministry of Agriculture and Fisheries, Occassional Pub-
lication 47: 8–11
Cudmore-Vokey B, Crossman EJ (2000) Checklists of the fish
fauna of the Laurentian Great Lakes and their connecting
channels. Can Manuscr Rep Fish Aquat Sci 2550:39
Davaine P (1991) Sea ranching of coho salmon in the Ker-
guelen Islands (TAAF). ICES, Copenhagen, 21 pp
Davaine P, Beall E (1982) Introductions of salmonids in the
French Austral Territories. Com Natl Fr Rech Antarct
51:289–300
Davaine P, Beall E (1997) Introduction en milieu vierge (Iles
Kerguelen, Subantarctique): ejeux re
´sultats, perspectives.
Bull Fr Peche Piscic 1997(344–345):93–110
Davidson F, Hutchinson SJ (1938) The geographic distribution
and environmental limitations of the Pacific salmon
(genus Oncorhynchus). Bull US Bur Fish 48:667–692
de Groot SJ (1985) Introductions of non-idigenous fish species
for release and culture in the Netherlands. Aquaculture
46:237–257
de Moor IJ, Bruton MN (1988) Atlas of alien and translocated
indigenous aquatic animals in southern Africa. A report of
the Committee for Nature Conservation Research
National Programme for Ecosystem Research. South
African Scientific Programmes Report No. 144, Port
Elizabeth, 310 pp
de Sostoa A, Lobon-Cervia J (1989) Fish and fisheries of the
River Ebro: actual state and recent history. In: Petts GE
(ed) Historical change of large alluvial rivers: western
Europe. John Wiley & Sons, London, 233 pp
Dextrase AJ, Coscarelli MA (1999) Intentional introductions of
nonindigenous freshwater organisms in North America.
In: Claudi R, Leach JH (eds) Nonindigenous freshwater
organisms: vectors, biology and impacts. Lewis Publish-
ers, Boca Raton, p 61
DFO (2003) National code on introductions and transfers of
aquatic organisms. http://www.dfo-mpo.gc.ca/science/
aquaculture/code/prelim_e.htm. Accessed 28 November
2004
Drake DC, Naiman RJ (2000) An evaluation of restoration
efforts in fishless lakes stocked with exotic trout. Conserv
Biol 14:1807–1820
Dunham JB, Adams SB, Schroeter RE et al (2002) Alien
invasions in aquatic ecosystems: toward an understanding
of brook trout invasions and potential impacts on inland
cutthroat trout in western North America. Rev Fish Biol
Fish 12:373–391
Dunham JB, Pilliod DS, Young MK (2004) Assessing the
consequences of nonnative trout in headwater ecosystems
in western North America. Fisheries 29:18–26
Dymond JR, Hart JL, Pritchard AL (1929) The fishes of the
Canadian waters of Lake Ontario. Univ Toronto Stud Biol
37:1–35
Dymond JR (1947) Checklist of the freshwater fishes of Can-
ada east of the Rocky Mountains. R Ont Mus Life Sci
Misc Publ 1:1–36
Dymond JR (1955) The introduction of foreign fishes in Can-
ada. Proc Int Assoc Theor Appl Limnol 12:543–553
EA (2004) Our nations’ fisheries: the migratory and freshwater
fisheries of England and Wales – a snapshot. Environment
Agency, United Kingdom, 97 pp
Eccles DH (1992) FAO species identification sheets for fishery
purposes. Field guide to the freshwater fishes of Tanzania.
FAO, Rome, 145 pp
Economidis PS (1991) Check list of freshwater fishes of Greece
(recent status of threats and protection). Hellenic Society
for the Protection of Nature, Special publication, 48 pp
Economidis PS, Dimitriou E, Pagoni R et al (2000) Introduced
and translocated fish species in inland waters of Greece.
Fish Manag Ecol 7:239–250
Eldredge LG (1994) Freshwater fishes. In: Eldredge LG (ed)
Perspectives in aquatic exotic species management in the
Pacific Islands, vol 1. Introductions of commercially sig-
nificant aquatic organisms to the Pacific Islands. South
Pacific Commission, New Caledonia, p 73
Rev Fish Biol Fisheries
123

Elvira B (1998) Impact of introduced fish on the native
freshwater fish fauna of Spain. In: Cowx IG (ed) Stocking
and introduction of fish. Fishing News Books, United
Kingdom, p 186
Emery L (1985) Review of fish species introduced into the
Great Lakes, 1819–1974. Great Lakes Fishery Commis-
sion Technical Report No. 45, Ann Arbor, 32 pp
Epifanio J (2000) The status of coldwater fishery management
in the United States: an overview of state programs.
Fisheries 25:13–27
Erdman DS (1984) Exotic fishes in Puerto Rico. In: Courtenay
WR Jr, Stauffer JR Jr (eds) Distribution, biology and
management of exotic fishes. Johns Hopkins University
Press, Baltimore, p 162
FAO (1993) Aquaculture production (1985–1991). FAO Fish
Circ 815, Rev 5, 213 pp
FAO (1997) FAO Database on introduced aquatic species.
FAO, Rome http://www.fao.org/fi/website/FIRetrieve
Action.do?dom=collection&xml=dias.xml. Accessed 26
October 2007
Farrington LW, Austin CM, Burridge CP et al (2004) Allo-
zyme diversity in Australian rainbow trout, Oncorhynchus
mykiss. Fish Manag Ecol 11:97–106
Flajshans M, Linhart O, Slechtova
´V et al (1999) Genetic
resources of commercially important fish species in the
Czech Republic: present state and future strategy. Aqua-
culture 173:471–483
Fletcher AR (1986) Effects of introduced fish in Australia. In:
De Dekker P, Williams WD (eds) Limnology in Australia.
CSIRO, Melbourne, p 231
Fricke R (1999) Fishes of the Mascarene Islands (Re
´union,
Mauritius, Rodriguez): an annotated checklist, with
descriptions of new species. Koeltz Scientific Books,
Koenigstein, Theses Zoologicae, vol 31, 759 pp
Frier JO (1994) Danmark. Danske ferskvandsfisk og deres
udbredelsesomra
˚de. In: Frier JO (ed) Truede Ferskv-
andsfiskearter i Norden. TemaNord 1994:625, Nordisk
Ministerra
˚d, København (in Danish), pp 4–6, 83–99
Froese R, Pauly D (2004) FishBase version (10/2004).
http://www.fishbase.org. Accessed 7 December 2004
Frost WE (1974) A survey of the rainbow trout (Salmo
gairdneri) in Britain and Ireland. Salmon and Trout
Asssociation, London, 36 pp
Fuller PL, Nico LG, Williams JD (1999) Nonindigenous fishes
introduced into inland waters of the United States. Special
Publication No. 27. American Fisheries Society, Bethesda
Fulton W (2004) Review of trout stocking in South Australia. S
Aust Fish Manag Ser 41: 1–67
Gabrielyan BK (2001) An annotated checklist of freshwater
fishes of Armenia. Naga ICLARM Q 24:23–29
Gandolfi G, Zerunian S, Torricelli P et al (1991) I pesci delle
acque interne italiane. Ministero dell’Ambiente e Unione
Zoologica Italiana, Instituto Poligrafico e Zecca dello
Stato, Roma, 616 pp
Gerstmeier R, Romig T (1998) Die Su
¨ßwasserfische Europas:
fu
¨r Naturfreunde und Angler. Franckh-Kosmos Verlag,
Stuttgart, 368 pp
Golani D, Mires D (2000) Introduction of fishes to the fresh-
water system of Israel. Isr J Aquacult/Bamidgeh 52:47–60
Golani D, Shefler D (1985) The occurrence of rainbow trout
(Salmo gairdneri) and coho salmon (Oncorhynchus
kisutch) in Lake Kinneret (Lake Tiberias). Bamidgeh
37:27–31
Gopi KC (2000) Freshwater fishes of Kerala State. In: Ponniah
AG, Gopalakrishnan A (eds) Endemic fish diversity of
western Ghats. NBFGR-NATP Publication, National
Bureau of Fish Genetic Resources, Lucknow, 56 pp
Griffiths D (1997) The status of the Irish freshwater fish fauna:
a review. J Appl Ichthyol 13:9–13
Gross M, Patel A, Correa C et al (2004) NSERC strategic grant
research update: breeding ecology of Pacific salmon in
Lake Ontario. Department of Zoology, University of
Toronto, Ontario, 34 pp
Hallock RJ, Fry DH Jr (1967) Five species of salmon Oncor-
phynchus, in the Sacramento River, California. Calif Fish
Game 53:5–22
Ham KD, Pearsons TN (2001) A practical approach for con-
taining ecological risks associated with fish stocking
programs. Fisheries 26:15–23
Hanamura N (1967) Salmon of the north Pacific Ocean, Part IV.
Spawning populations of north Pacific salmon, 1. sockeye
salmon in the Far East. Int Pac Fish Comm 23:1–7
Hardy CJ (1983) Origin of NZ’s sockeye. Freshwater Catch
(NZ) 18:11–13
Hart JL (1973) Pacific fishes of Canada. Bulletin 180, Fisheries
Research Board of Canada, Ottawa, 740 pp
Hasegawa K, Maekawa K (2006) The effects of introduced
salmonids on two native stream-dwelling salmonids
through interspecific competition. J Fish Biol 68:
1123–1132
Hayes JP, Guffey SZ, Kriegler FJ et al (1996) The genetic
diversity of native, stocked, and hybrid populations of
brook trout in the southern Appalachians. Conserv Biol
10:1403–1412
Headland RK (1984) The island of South Georgia. Cambridge
University Press, United Kingdom, 293 pp
Healey MC (1991) Life history of chinook salmon (On-
corhynchus tshawytscha). In: Groot C, Margolis L (eds)
Pacific salmon life histories. UBC Press, Vancouver,
p 311
Hendry AP, Quinn TP (1997) Variation in adult life history and
morphology among Lake Washington sockeye salmon
(Oncorhynchus nerka) populations in relation to habitat
features and ancestral affinities. Can J Fish Aquat Sci
54:75–84
Hendry AP, Quinn TP, Utter FM (1996) Genetic evidence for
the persistence and divergence of native and introduced
sockeye salmon (Oncorhynchus nerka) within Lake
Washington, Washington. Can J Fish Aquat Sci 53:
823–832
Hendry AP, Hensleigh JE, Reisenbichler RR (1998) Incubation
temperature, developmental biology, and the divergence
of sockeye salmon (Oncorhynchus nerka) within Lake
Washington. Can J Fish Aquat Sci 55:1387–1394
Holcı
´k J (1991) Fish introductions in Europe with particular
reference to its central and eastern part. Can J Fish Aquat
Sci 48(Suppl. 1):13–23
Holcı
´k J, Razavi BA (1992) On some new or little known
freshwater fishes from the Iranian coast of the Caspian
Sea. Folia Zool 41:271–280
Holmlund CM (2000) 150 years of Fish Stocking in the
Archipelago of Stockholm: gambling with ecological and
Rev Fish Biol Fisheries
123

social resilience? Unpublished manuscript, Draft version
2000-04-26
Holmlund CM, Hammer M (2004) Effects of fish stocking on
ecosystem services: an overview and case study using the
Stockholm Archipelago. Environ Manage 33:799–820
Horak D (1995) Native and nonnative fish species used in state
fisheries management programs in the United States. Am
Fish Soc Symp 15:61–67
Hubbs CL, Lagler KF (1958) Fishes of the Great Lakes region.
University of Michigan Press, Bloomfield Hills, 213 pp
Hutchinson GE (1957) Concluding remarks. Cold Springs
Harbor Symposium on Quantitative Biology, pp 415–427
Innal D, Erk’akan F (2006) Effects of exotic and translocated
fish species in the inland waters of Turkey. Rev Fish Biol
Fish 16:39–50
ISSG (2007) Global Invasive Species Database.
http://www.issg.org. Accessed 8 Aug 2007
Johnson DM, Behnke RJ, Harpman DA et al (1995) Economic
benefits and costs of stocking catchable rainbow trout: a
synehesis of economic analysis in Colorado. N Am J Fish
Manag 15:26–32
Jonsson G (1992) Islenskir fiskar. Fiolvi, Reykjavik, 568 pp
Kaczynski VW, Alvarado F (2006) Assessment of the southern
range limit of North American coho salmon: difficulties in
establishing natural range boundaries. Fisheries 13:
374–391
Kailola PJ, Williams MJ, Stewart PC et al (1993) Australian
fisheries resources. Bureau of Resource Sciences, Can-
berra, 422 pp
Kamilov G, Urchinov ZU (1995) Fish and fisheries in Uzbe-
kistan under the impact of irrigated agriculture. In: Petr T
(ed) Inland fisheries under the impact of irrigated agri-
culture: central Asia. FAO Fisheries Circular No. 894,
Rome, pp 10–41
Kapoor D, Dayal R, Ponniah AG (2002) Fish biodiversity of
India. National Bureau of Fish Genetic Resources, Luc-
know, India, 775 pp
Keith P, Allardi J (1998) The introduced freshwater fish of
France: status, impacts and management. In: Cowx IG
(ed) Stocking and introduction of fish. Fishing News
Books, MPG Books Ltd., Bodmin, p 153
Keith P, Allardi J, Moutou B (1992) Livre rouge des espe
`ces
menace
´es de poissons d’eau douce de France et bilan des
introductions. Museum National d’Histoire Naturelle,
Secretariat de la Faune et de la Flore, Conseil Supe
´rieur
de la Peche, CEMAGREF and Ministe
`re de l’Environ-
ment, 111 pp
Keith P, Vigneux E, Bosc P (1999) Atlas des poissons et des
crustace
´s d’eau douce de La Re
´union. Patrimoines Natu-
rels (M.N.H.N., Paris/S.P.N.), 39, 136 pp
Keller M, Smith KD, Rybicki RW (1989) Summary of salmon
and trout management in Lake Michigan. Michigan
Department of Natural Resources, Fisheries Division,
Technical Report 89-1, 27 pp
Kerr SJ, Grant RE (2000) Ecological impacts of fish intro-
ductions: evaluating the risk. Ontario Ministry of Natural
Resources, Fish and Wildlife Branch, Peterborough,
473 pp
Kerr SJ, Lasenby TA (2000) Rainbow trout stocking in inland
lakes and streams: an annotated bibliography and litera-
ture review. Ontario Ministry of Natural Resources, Fish
and Wildlife Branch, Fisheries Section, Perterborough,
228 pp
Khurshut EE (2001) Fishes of Uzbekhistan. FishBase printout
checked by Ernest Khurshut
Kido MH, Heacock DE, Asquith A (1999) Alien rainbow trout
(Oncorhynchus mykiss) (Salmoniformes: Salmonidae) diet
in Hawaiian streams. Pac Sci 53:242–251
Kinnison M, Unwin M, Boustead N et al (1998) Population-
specific variation in body dimensions of adult chinook
salmon (Oncorhynchus tshawytscha) from New Zealand
and their source population, 90 years after introduction.
Can J Fish Aquat Sci 55:554–563
Kırkan S, Go
¨ksoy EO, Kaya O (2003) Isolation and antimi-
crobial susceptibility of aeromonas salmonicida in
rainbow trout (Oncorhynchus mykiss) in Turkey hatchery
farms. J Vet Med Ser B 50:339–342
Knapp RA, Matthews KR (2000) Effects of nonnative fishes on
wilderness lake ecosystems in the Sierra Nevada and
recommendations for reducing impacts. USDA Forest
Service Proceedings RMRS-P-15-VOL-5, 312 pp
Knapp RA, Corn PS, Schindler DE (2001) The introduction of
nonnative fish into wilderness lakes: good intentions,
conflicting mandates, and unintended consequences.
Ecosystems 4:275–278
Kocik JF, Jones ML (1999) Pacific salmonines in the Great
Lakes basin. In: Taylor WW, Ferreri CP (eds) Great Lakes
fisheries policy and management: a binational perspective.
Michigan State University Press, East Lansing, p 455
Koli L (1990) Suomen kalat. [Fishes of Finland]. Werner
So
¨derstro
¨m Osakeyhtio
¨, Helsinki, 357 pp
Krueger CC, May B (1991) Ecological and genetic effects of
salmonid introductions in North America. Can J Fish
Aquat Sci 48(Suppl. 1):66–77
Krupp F, Schneider W (1989) The fishes of the Jordan River
drainage basin and Azraq Oasis. Fauna Saudi Arabia 10:
347–416
Kucera PA, Koch DL, Marco GF (1985) Introductions of
Lahontan cutthroat trout into Omak Lake, Washington.
N Am J Fish Manag 5: 296–301
Kullander SO (1999) Swedish fishes: checklist of Swedish fishes.
http://www.nrm.se/ve/pisces/allfish.shtml.en. Accessed 10
June 2007
Kwain W (1987) Biology of pink salmon in the North Amer-
ican Great Lakes. In: Dadswell MJ, Kluada RJ, Moffitt
CM et al (eds) American Fisheries Society Symposium,
vol 1. American Fisheries Society, Bethesda, pp 57–65
Landergren P (1999) Spawning of anadromous rainbow trout,
Oncorhynchus mykiss (Walbaum): a threat to sea trout,
Salmo trutta L., populations? Fish Res 40:55–63
Leach JH, Lewis CA (1991) Fish introductions in Canada:
provincial views and regulations. Can J Fish Aquat Sci
48(Suppl. 1):156–161
Lever C (1996) Naturalized fishes of the world. Academic
Press, California, 408 pp
Liao C-I, Su H-M, Chang EY (2001) Techniques in finfish
larviculture in Taiwan. Aquaculture 200:1–31
Liao IC, Lia HC (1989) Exotic aquatic species in Taiwan. In:
De Silva SS (ed) Exotic aquatic organisms in Asia pro-
ceedings of the workshop on introduction of exotic
aquatic organisms in Asia Spec Publ 3. Asian Fisheries
Society, Manila, Philippines, p 101
Rev Fish Biol Fisheries
123

Lukin AA (1998) The introduction of trout, Parasalmo mykiss,
into Lake Imandra (Kola Peninsula). J Ichthyol 38:
511–516
MacCrimmon H (1965) The beginning of salmon culture in
Canada. Can Geog J 71:96–103
MacCrimmon HR (1971) World distribution of rainbow trout
(Salmo gairdneri). J Fish Res Board Can 28:663–704
MacCrimmon HR (1977) Animal, man and change: alien and
exotic wildlife of Ontario. McClelland and Stewart,
Toronto
MacCrimmon HR, Marshall TL, Gots BL (1970) World dis-
tribution of brown trout, Salmo trutta: further
observations. J Fish Res Board Can 27:811–818
Machidori S, Kato F (1984) Spawning populations and marine
life of masu salmon (Oncorhynchus masou). Int N Pac
Fish Comm Bull 43:1–138
Maciolek JA (1984) Exotic fishes in Hawaii and other islands
of Oceania. In: Courtenay WR Jr, Stauffer JR Jr (eds)
Distribution, biology and management of exotic fishes.
Johns Hopkins University Press, Baltimore, p 131
MacKay HH (1969) Fishes of Ontario. Ontario Department of
Lands and Forests, Toronto, 292 pp
MacKinnon M (1987) Review of inland fisheries – Australia.
FAO Fish Rep 371(Suppl):1–37
Magomedov GM (1978) Some data on the biological basis for
acclimatization of the chum salmon, Oncorhynchus keta,
in the Caspian Sea. J Ichthyol 18:318–323
Marceta B (1999) Check-list of the fishes of the Gulf of Trieste
(North Adriatic Sea). Provided on spreadsheet to FishBase
McDowall RM (1984) Exotic fishes: The New Zealand expe-
rience. In: Courtenay WR Jr, Stauffer JR Jr (eds)
Distribution, biology and management of exotic fishes.
Johns Hopkins University Press, Baltimore, p 200
McDowall RM (1990a) New Zealand freshwater fishes: a
natural history and guide. Heinemann Reed, Auckland
McDowall RM (1990b) Freshwater fishes and fisheries of New
Zealand – the angler’s el dorado. Aquat Sci 2:281–341
McDowall RM (1994a) Gamekeepers for the nation: the story
of New Zealand’s acclimatisation societies, 1861–1990.
Canterbury University Press, Christchurch, 508 pp
McDowall RM (1994b) The origins of New Zealand’s chinook
salmon, Oncorhynchus tshawytscha. Mar Fish Rev 56:1–7
McDowall RM (1999) Different kinds of diadromy: different
conservation problems. ICES J Mar Sci 56:410–413
McDowall RM (2003) Impacts of introduced salmonids on
native galaxiids in New Zealand upland streams: a new
look at an old problem. Trans Am Fish Soc 132:229–238
McDowall RM (2006) Crying wolf, crying foul, or crying
shame: alien salmonids and a biodiversity crisis in the
southern cool-temperate galaxioid fishes? Rev Fish Biol
Fish 16:233–422
McDowall RM, Allibone RM, Chadderton WL (2001) Issues
for the conservation and management of Falkland Islands
freshwater fishes. Aquat Conserv: Mar Freshw Ecosyst
11:473–486
McHugh P, Budy P (2006) Experimental effects of nonnative
brown trout on the individual- and population-level per-
formance of native Bonneville cutthroat trout. Trans Am
Fish Soc 135:1441–1455
McPhail JD, Carveth R (1993) Field key to the freshwater
fishes of British Columbia. Fish Museum, Department of
Zoology, University of British Columbia, Vancouver,
239 pp
Mills EL, Leach JH, Carlton JT, Secor CL (1993) Exotic
species in the Great Lakes: a history of biotic crises and
anthropogenic introductions. J Great Lakes Res 19:1–54
Mirza MR (2003) Checklist of freshwater fishes of Pakistan.
Pakistan J Zool Suppl Ser 3:1–30
Mitrofanov VP, Petr T (1999) Fish and fisheries in the Altai,
Northern Tien Shan and Lake Balkhash (Kazakhstan). In:
Petr T (ed) Fish and fisheries at higher altitudes: Asia.
FAO Fish Tech Pap No 385, Rome, p 149
Molony BW (2001) Environmental requirements and toler-
ances of rainbow trout (Oncorhynchus mykiss) and brown
trout (Salmo trutta) with special reference to Western
Australia: a review. Department of Fisheries, Western
Australia Fisheries Research Report 130, 28 pp
Molony BW, Lenanton R, Jackson G et al (2003) Stock
enhancement as a fisheries management tool. Rev Fish
Biol Fish 13:409–432
Monnerjahn U (1999) Freshwater fishes of Germany. A
checklist compiled from German FischKataster of the
Bundesla
¨nder and from the German ‘Rote Liste’. Pro-
vided as Excel spreadsheet for use by FishBase
Moring JR (2000) The creation of the first public salmon
hatchery in the United States. Fisheries 25:6–12
Morrow JE (1980) The freshwater fishes of Alaska. University
of British Columbia Animal Resources Ecology Library,
Vancouver, 248 pp
Mouslih M (1987) Fish and crayfish introductions in Morocco.
Rev Hydrobiol Trop 20:65–72
Moyle PB (1976) Inland fishes of California. University of
California Press, Berkeley, 405 pp
Moyle PB (1986) Fish introductions into North America: pat-
terns and ecological impact. In: Mooney HA, Drake JA
(eds) Ecology of biological invasions of North America
and Hawaii. Springer, New York, p 27
Moyle PB, Light T (1996) Biological invasions of fresh water:
empirical rules and assembly theory. Biol Conserv
78:149–161
Muus BJ, Dahlstro
¨m P (1968) Su
¨ßwasserfische. BLV Ver-
lagsgesellschaft, Mu
¨nchen, 224 pp
Needham PR, Gard R (1959) Rainbow trout in Mexico and
California with notes on the cutthroat series. University of
California Press, Berkeley, 124 pp
Nico LG, Fuller PL (1999) Spatial and temporal patterns of
nonindigenous fish introductions in the United States.
Fisheries 24:16–27
Nielsen JG, Bertelsen E (1992) Fisk i grønlandske farvande.
Atuakkiorfik, Nuuk, 68 pp
Nijssen H, de Groot SJ (1974) Catalogue of fish species of the
Netherlands. Beaufortia 21:173–207
Okeyo DO (1998) Taxonomy, common names and distribution
of fish in the eastern arm of the Rift Valley Drainage,
Kenya. University of Fort Hare, Eastern Cape, 19 pp
Page LM, Burr BM (1991) A field guide to freshwater fishes of
North America north of Mexico. Houghton Mifflin
Company, Boston, 432 pp
Parsons W (1973) History of salmon in the Great Lakes, 1850–
1970. US Bur Sport Fish Wildl, 80 pp
Pascual MA, Bentzen P, Riva Rossi C et al (2001) First doc-
umented case of anadromy in a population of introduced
Rev Fish Biol Fisheries
123

rainbow trout in Patagonia, Argentina. Trans Am Fish Soc
130:53–67
Pascual MA, Macchi P, Urbanski J et al (2002) Evaluating
potential effects of exotic freshwater fish from incom-
plete species presence-absence data. Biol Invasions
4:101–113
Paulin C, Stewart A, Roberts C et al (1989) New Zealand fish:
a complete guide. National Museum of New Zealand
Miscellaneous Series No. 19, 279 pp
Pavlov PI (1980) Fauna of Ukraine. Fishes. Tunicata (Ascidian,
appendicularian), Acrania (Cephalochordata), Vertebrata
(Cyclostomata; cartilaginous fishes, bony fishes – stur-
geons; clupeids; anchois; salmonides; ombres; brochets;
umbres). Kiev, Naukova Dumka Publishing House, 8(1)
Paxton JR, Hoese DF, Allen GR et al (1989) Pisces. Petro-
myzontidae to Carangidae. Zoological Catalogue of
Australia, vol 7. Australian Government Publishing Ser-
vice, Canberra
Pearsons TN, Hopley CW (1999) A practical approach for
assessing ecological risks associated with fish stocking
programs. Fisheries 24:16–23
Peck JW, Jones TS, MacCallum WR et al (1999) Contribution
of hatchery reared fish to chinook salmon populations and
sport fisheries in Lake Superior. N Am J Fish Manag
19:155–164
Pe
´rez LA, Winkler FM, Dı
´az NF et al (2001) Genetic vari-
ability in four hatchery strains of coho salmon,
Oncorhynchus kisutch (Walbaum), in Chile. Aquac Res
32:41–46
Pethiyagoda R (1991) Freshwater fishes of Sri Lanka. The
Wildlife Heritage Trust of Sri Lanka, Colombo, 362 pp
Pietsch TW, Amaoka K, Stevenson DE et al (2000) Freshwater
fishes of the Kuril Islands and adjacent regions. Interna-
tional Kuril Island Project (IKIP), University of
Washington Fish Collection, Washington, 18 pp
Pister EP (2001) Wilderness fish stocking: history and per-
spective. Ecosystems 4:279–286
Piyakarnchana T (1989) Exotic aquatic species in Thailand. In:
De Silva SS (ed) Exotic aquatic organisms in Asia pro-
ceedings of the workshop on introduction of exotic
aquatic organisms in Asia Spec Publ 3. Asian Fisheries
Society, Manila, p 119
Popov AM (1933) Fishes of Avatcha Bay on the southern coast
of Kamchatka. Copeia 1933:59–67
Quinn TP, Nielsen JL, Gan C et al (1996) Origin and genetic
structure of chinook salmon, Oncorhynchus tshawytscha,
transplanted from California to New Zealand: allozyme
and mtDNA evidence. Fish Bull 94:506–521
Raat AJP (2003) Stocking of sea trout, Salmo trutta, in Lake
Veere, south-west Netherlands. Fish Manag Ecol 10:
61–71
Rahel FJ (2000) Homogenization of fish faunas across the
United States. Science 288:854–856
Reshetnikov YS, Bogutskaya NG, Vasil’eva ED et al (1997)
An annotated check-list of the freshwater fishes of Russia.
J Ichthyol 37:687–736
Ricker WE, Loftus KH (1968) Pacific salmon move east. Fish
Council Can Annu Rev 23:37–39
Rinne JH, Janisch J (1995) Coldwater fish stocking and native
fishes in Arizona: past, present, and future. Am Fish Soc
Symp 15:397–406
Riva Rossi CM, Lessa EP, Pascual MA (2004) The origin of
introduced rainbow trout (Oncorhynchus mykiss) in the
Santa Cruz River, Patagonia, Argentina, as inferred from
mitochondrial DNA. Can J Fish Aquat Sci 61:1095–1101
Robins CR, Bailey RM, Bond CE et al (1991) Common and
scientific names of fishes from the United States and
Canada. American Fisheries Society Special Publication
20, Bethesda, 183 pp
Roughley TC (1961) Fish and Fisheries of Australia. Angus
and Robertson, Sydney, 343 pp
Sartorio A (2003) Tropical adaptation of rainbow trout (On-
corhynchus mykiss). Aliens Newsletter of Invasive
Species Specialist Group, 17, 30 pp
Savvaitova KA, Petr T (1999) Fish and fisheries in Lake Issyk-
kul (Tien Shan), River Chu and Pamir lakes. In: Petr T
(ed) Fish and fisheries at higher altitudes: Asia. FAO Fish
Tech Pap No. 385, Rome, p 168
Scott D (1984) Origin of the New Zealand sockeye salmon,
Oncorhynchus nerka (Walbaum). J R Soc N Z 14:245–247
Scott WB, Crossman EJ (1973) Freshwater fishes of Canada.
Bulletin 184, Fisheries Research Board of Canada,
Ottawa, 966 pp
Scott D, Hewitson SJ, Fraser JS (1978) The origin of rainbow
trout, Salmo gairdneri Richardson, in New Zealand. Calif
Fish Game 64:200–209
Shaji CP, Easa PS, Gopalakrishnan A (2000) Freshwater fish
diversity of Western Ghats. In: Ponniah AG, Gopala-
krishnan A (eds) Endemic fish diversity of Western Ghats.
NBFGR-NATP Publication. National Bureau of Fish
Genetic Resources, Lucknow, p 33
Shetty HPC, Nandeesha MC, Jhingran AG (1989) Impact of
exotic aquatic species in Indian waters. In: De Silva SS
(ed) Exotic aquatic organisms in Asia proceedings of the
workshop on introduction of exotic aquatic organisms in
Asia Spec Publ 3. Asian Fisheries Society, Manila, p 45
Shrestha J (1994) Fishes, fishing implements and methods of
Nepal. Smt. M.D. Gupta, Lalitpur Colony, Lashkar
(Gwalior), India, 150 pp
Skelton PH (1987) South African red data book – fishes. South
African National Scientific Programmes Report No. 137,
CSIR Pretoria, 199 pp
Skelton PH (1993) A complete guide to the freshwater fishes of
southern Africa. Southern Book Publishers (Pty) Ltd.,
388 pp
Sklower A (1951) Carp breeding in Israel. Arch Fis-
chereiweeiss 11:42–54
Smedley HH (1938) Rainbow, Salmo irrideus. In: Trout in
Michigan, Muskegon, p 25
SNH (2003) River Spey: catchment management plan. Scottish
Natural Heritage, Achantoul, 84 pp
Soto D, Jara F, Moreno C (2001) Escaped salmon in the inner
seas, southern Chile: facing ecological and social con-
flicts. Ecol Appl 11:1750–1762
Stephenson SA (2005) The distribution of Pacific salmon
(Oncorhynchus spp.) in the Canadian western Arctic.
Canadian Manuscript Report of Fisheries and Aquatic
Sciences 2737, Fisheries and Oceans Canada, Winnepeg,
Manitoba, 29 pp
Stewart DJ, Ibarra M (1991) Predation and production by
salmonine fishes in Lake Michigan, 1978–88. Can J Fish
Aquat Sci 48:909–922
Rev Fish Biol Fisheries
123

Stiassny MLJ, Raminosoa N (1994) The fishes of the inland
waters of Madagascar, Biological diversity of African
fresh- and brackish-water fishes. Geographical overviews
presented at the PARADI Symposium, Senegal, 15–20
November 1993, Ann Mus R Afr Centr, Sci Zool,
pp 133–148
Svetovidov AN (1984) Salmonidae. In: Whitehead PJP, Bau-
chot M-L, Hureau J-C et al (eds) Fishes of the North-
Eastern Atlantic and the Mediterranean. UNESCO, Paris,
pp 373–385
Swaby SE, Potts GW (1990) Rare British marine fishes –
identification and conservation. J Fish Biol 37(Suppl.
A):133–143
Swales S (1990) Conservation status of large galaxiids (Pisces:
Galaxiidae) in tributaries of the Waikato River, New
Zealand. J Fish Biol 37(Suppl. A):217–218
Taft AC (1937) A red salmon (Oncorhynchus nerka) taken in
the Klamath River. Calif Fish Game 23:178
Takami T, Aoyama T (1999) Distribution of rainbow trout and
brown trout in Hokkaido, northern Japan. Wildl Conserv
Japan 4:41–48
Talwar PK, Jhingran AG (1991) Inland fishes of India and
adjacent countries, vol 2. A.A. Balkema, Rotterdam
Tan Y, Tong HE (1989) The status of the exotic aquatic
organisms in China. Exotic aquatic organisms in Asia
proceedings of the workshop on introduction of exotic
aquatic organisms in Asia Spec Publ 3, Manila, Philip-
pines, Asian Fisheries Society, p 35
Tedla S, Meskel FH (1981) Introduction and transplantation of
freshwater fish species in Ethiopia. Ethiop J Sci 4:69–72
Thurow RF, Lee DC, Rieman BE (1997) Distribution and
status of seven native salmonids in the interior Columbia
River basin and portions of the Klamath River and Great
Basins. N Am J Fish Manag 17:1094–1110
Thys van den Audenaerde DFE (1994) Introduction of aquatic
species into Zambian waters, and their importance for
aquaculture and fisheries. Aquaculture for Local Com-
munity Development Programme, ALCOM Field
Document No. 24, 29 pp
Trautman MB (1981) The fishes of Ohio. Ohio State University
Press, Columbus, 782 pp
UN (2007) United Nations Member States. http://www.un.org.
Accessed 8 Aug 2007
Unwin MJ (1991) Hatchery releases and the salmon sports
fishery. Salmon towards 2000 Addington Raceway,
Christchurch, 27–28 October 1990, New Zealand Ministry
of Agriculture and Fisheries, pp 18–25
Unwin MJ, Glova GJ (1997) Changes in life history parameters
in a naturally spawning population of chinook salmon
(Oncorhynchus tshawytscha) associated with releases of
hatchery-reared fish. Can J Fish Aquat Sci 54:1235–1245
Unwin MJ, James GD (1998) Occurrence and distribution of
adult chinook salmon in the New Zealand commercial
fishery. Trans Am Fish Soc 127:560–575
USGS (2004) Nonindigenous Aquatic Species (NAS) infor-
mation resource. http://www.nas.er.usgs.gov/. Accessed
24 November 2004
van Zyll de Jong MC, Gibson RJ, Cowx IG (2004) Impacts of
stocking and introductions on freshwater fisheries of
Newfoundland and Labrador, Canada. Fish Manag Ecol
11:183–193
Waddington DG, Laughland A (1996) Trout fishing in the U.S.
U.S. Fish and Wildlife Service, Report 91-5, Arlington
Walczak P (1972) A brief review of Salmonidae in Iran.
Manuscript of the Fisheries Research Institute, Bandhar,
7pp
Walker KF, Yang HZ (1999) Fish and fisheries in western
China. In: Petr T (ed) Fish and fisheries at higher altitudes:
Asia. FAO Fish Tech Pap No. 385, Rome, p 237
Walker BW, Whitney RR, Barlow GW (1961) The fishes of the
Salton Sea. In: Walker BW (ed) The ecology of the salton
sea, California, in relation to the sport fishery of Cali-
fornia. California Department of Fish and Game, Fish
Bulletin 113, Sacramento, p 77
Watkins BP, Cooper J (1986) Introduction, present status and
control of alien species at the Prince Edward Islands, sub-
Antarctic. S Afr J Antarct Res 16:86–94
Weigel DE, Peterson JT, Spruell P (2003) Introgressive
hybridization between native cutthroat trout and intro-
duced rainbow trout. Ecol Appl 13:38–50
Weiss S, Kummer H (1999) Movement of wild and experi-
mentally stocked brown trout, Salmo trutta L., before and
after a flood in a small Austrian stream. Fish Manag Ecol
6:35–45
Welcomme RL (1988) International introductions of inland
aquatic species. FAO, Rome, Fish Tech Pap 294, 328 pp
Went AEJ (1957) List of Irish fishes. Department of Lands,
Fisheries Division, Dublin, 31 pp
West GJ, Glucksman J (1976) Introduction and distribution of
exotic fish in Papua New Guinea. Papua New Guinea
Agric J 27:19–48
Westerman FA (1930) Fish division. Mich Dep Conserv 5th
Bienn Rep., 1927–28, pp 191–233
Winkler HM, Skora K, Repecka R et al (2000) Checklist and
status of fish species in the Baltic Sea. ICES CM 2000/
Mini:11, 15 pp
Worth SG (1895) Report on the propagation and distribution of
food-fishes. U.S. Comm. Fish, Fish. Rep. 1893. Part
19:73–138
WRI (2003) Water Resources eAtlas: watersheds of the world.
http://www.iucn.org. Accessed 8 Aug
Xie Y, Lin Z, Gregg WP et al (2001) Invasive species in
China – an overview. Biodivers Conserv 10:1317–1341
Yamamoto MN (1992) Occurrence, distribution and abundance
of accidentally introduced freshwater aquatic organisms in
Hawaii. State of Hawaii, Federal Aid in Sportfish Resto-
ration, Dingell-Johnson JOR. Freshwater Fisheries
Research and Surveys, Project No. F-14-R-16
Yoshiyama RW, Fisher FW (2001) Long time past: Baird
Station and the McCloud Wintu. Fisheries 26:6–22
Zamorano RM (1991) Salmon farming in Chile. Proceedings of
the special session on salmonid aquaculture, World
Aquaculture Society, Los Angeles, 16 February 1989.
Canadian Technical Report of Fisheries and Aquatic
Sciences 1831:51–63
Rev Fish Biol Fisheries
123