A Historical Perspective on Salmonid Production from Pacific Rim Hatcheries

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A Historical Perspective on Salmonid Production
                                 from Pacific Rim Hatcheries
                Conrad Mahnkenl, Gregory Ruggerone2 , William Waknitzl, and Thomas Flagg l
                      lNorthwest Fisheries Science Center, 2725 Montlake Boulevard East
                                   Seattle Washington, 98112-2097, U.S.A.

                               2Natural Resource Consultants, 4055 21st Avenue West
                                         Seattle, Washington, 98199, U.S.A.

         Mahnken, C., G. Ruggerone, W. Waknitz, and T. Flagg. 1998. A historical perspective on salmonid
            production from Pacific rim hatcheries. N. Pac. Anadr. Fish Comm. Bull. No.1: 38-53

              Annual hatchery production rates of chinook (Oncorhynchus tshawytscha), coho (0. kisutch) ,
         sockeye (0. nerka) , pink (0. gorbuscha) , chum (0. keta), masu (0. masu} salmon, and steelhead trout
         (0. mykiss} were obtained from published and unpublished sources and compiled as a computer
         database. Pacific Rim hatchery production trends for the 40-year period from 1950-1992 were
         analyzed for all species from four geographic areas: Pacific Northwest (Washington, Oregon, Idaho,
         and California), Canada (British Columbia), Alaska, and Japan (Honshu and Hokkaido). Production
         of chum, sockeye, and pink salmon has increased dramatically in Japan, Canada, and Alaska in the
         past 20 years. Chinook, coho, steelhead, and masu have also experienced moderate increases in the
         same time period; however, production of coho, chinook and steelhead has declined since 1985.
         Trends in survival of hatchery fish over the period 1970-90 are demonstrated where data were
         available. We noted that survival of coho salmon was greatest for releases into large estuaries, such
         as southeast Alaska, Georgia Strait, and Puget Sound, than into drainages discharging directly into the
         Eastern Pacific Ocean. A negative cline in survival of coho salmon was observed moving both north
         and south from the center of the distribution of the species in British Columbia. Survival trends for fall
         chinook released north of Puget Sound tended to be the opposite of those released to the south.
         Survival of Japanese chum salmon released into the North Pacific Ocean has increased steadily from
         the mid-1960's to the present.

                  INTRODUCTION                                    Alaska, declining fish size and altered return timing
                                                                  and age at maturity, have raised concerns over limits
     Development of the North Pacific sahnonid                    on ocean carrying capacity.
hatchery system began in the late 19th century and has                  In the Pacific Northwest, recent Endangered
played a prominent role in enhancement of the                     Species Act listings of Redfish Lake sockeye in the
salmonid resource in Pacific Rim nations since the                Stanley Basin of Idaho, Sacramento River winter
1950s.     Until recently, the artificial propagation             chinook in California, and fall, summer, and spring
approach to enhancement of fisheries has not been                 runs of chinook in the Snake River Basin of Idaho and
seriously questioned, but the recent alarming declines            Oregon have focused attention not only on habitat loss
in wild spawning stocks have forced a re-evaluation of            and fishery-related impacts on the wild stock" but on
industrial-scale hatchery production of north Pacific             the genetic and demographic consequences of
sahnonids. Declines have been observed in chinook,                uncontrolled expansion of hatcheries as well (Meffe
coho, and sockeye sahnon stocks in the Pacific                    1992; Nehlsen et al. 1991). Proposed recovery plans
Northwest; high harvest rates of wild fish in fisheries           for listed Snake River chinooks and sockeye call for a
targeted on the more abundant hatchery stock., have               limit on annual releases from Columbia Basin
continued, and high production of hatchery chum                   hatcheries to 1994 levels (Schmitten et al. 1995).
sahnon in Japan and both pinks and chum sahnon in                       In Alaska, the hatchery successes that were hailed

                                                            38
in the mid 1980s are being assailed in the 1990s. The          and Oceans, 555 West Hastings Street, Vancouver,
return of record numbers of hatchery pink and chum             B.C., Pers. commun., Oct. 1996). Japanese release
salmon and high abundance of natural fish in the               information on chum and pink salmon was supplied by
North Pacific has led to record high catches and               the Hokkaido Salmon Hatchery (Masahide Kaeriyama
record low revenues to fishermen, and has brought              and Shigihiko Urawa, Hokkaido Salmon Hatchery, 2-
forward new criticism of hatchery management                   2 Nakano shima , Toyohira-ku, Sapporo 062, Japan,
strategies. Concerns over decreasing fish size in the          Pers. commun., Oct., 1996).
hatchery-based fishery for chum salmon in northern                  Until recently, a consolidated data set for hatchery
Japan has led to a decision by the Japanese                    production has been unavailable for the western
government to reduce hatchery releases. Similar                United States (Wahle and Smith 1979; Malmken et al.
concerns for declining size and increasing age at              1983; Isaksson 1988). We compiled a comprehensive
maturity observed in North Pacific stocks of five              historical data set from previously unreported raw data
salmon species suggests that large-scale hatchery              forms archived by fishery agencies, annual reports of
production is resulting in density-dependant growth            hatcheries, and electronic databases. We obtained
reduction (Kaeriyama and Urawa 1992; Bigler et al.             earlier salmon release records from Oregon and Idaho
1996).                                                         and steelhead from Washington which were previously
     This report provides managers with an historical          not available.
data set of hatchery releases from Pacific Rim nations,
excluding Russia, for the six Pacific salmon species           Survival Data
and steelhead through 1992. In addition, an analysis
of survival trends for hatchery coho and fall chinook               Survival information for coho and chinook salmon
from the eastern Pacific and chum from Japan are               was obtained from coded wire tag (CWT) data bases
presented.                                                     maintained by the Pacific States Marine Fisheries
                                                               Commission. The historical CWT data set was
                     METHODS                                   assembled by Maria Claribel Coronado-Hernandez
                                                               (1995) and is contained in her doctoral dissertation.
Release Data                                                   This seminal work synthesizes and analyzes, for the
                                                               first time, spatial and temporal factors affecting
     Annual production rates from Pacific Rim                  survival of hatchery-reared chinook and coho salmon
hatcheries were obtained for six species of Pacific            and steelhead in western North America over the time
salmon: chinook (Oncorhynchus tshawytscha), coho               period for which coded wire tag recoveries are
(0. ldsutch) , sockeye, (0. nerka) , pink (0.                  available (1971-1989). Release-recovery information
gorbuscha) , chum (0. keta) , masu (0. masu), and              is presented in the form of expanded CWT recoveries
steelhead trout (0. myldss).     We compiled this              using only non-experimental production groups.
information from published and unpublished sources             Recoveries were standardized to the most common age
and organized it in a computer database. Production            at return using virtual population analysis (Coronado-
trends for the 40-year period from 1950-1990 were              Hernandez 1995).
analyzed for all species from four geographic areas:                We computed mean survival rates from this data
Pacific Northwest (Washington, Oregon, Idaho, and              set and used two-way analysis of variance to make
California), Canada (British Columbia), Alaska, and            statistical comparisons of mean survival values over
Japan (Honshu and Hokkaido). Data from Russia and              time. We chose coho and fall chinook salmon for
Korea were unavailable as a continuous historical              analysis of temporal and geographic variatIOn because
series, however discontinuous data were available              of the large data sets available for these species.
from three sources and were used to estimate total             Pairwise comparisons between sampling dates were
present Pacific Rim output (Heard 1995; Malmken et             made using the Fishers PLSD test. All statements
al. 1983; Konovalov 1980).                                     about statistical comparisons are based on the P
     Release data from Alaska for chum, pink, coho,             < 0.05 significance level.
sockeye, and chinook salmon were obtained through                   Survival data for Japanese chum salmon was
Alaska Department of Fish and Game annual reports              provided by the Hokkaido Salmon Hatchery
(McNair 1996). Alaska steelhead releases were                  (Masahide Kaeriyama and Shigihiko Urawa, Hokkaido
supplied by Alaska Department of Fish and Game                 Salmon Hatchery, 2-2 Nakanoshima, Toyohira-ku,
(Marianne McNair, Alaska Department of Fish and                Sapporo 062, Japan, Pers. commun., Oct., 1996).
Game, P.O. Box 25526 Juneau Alaska, Pers.                      These data were compiled through records of fishery
commun., Oct., 1996). Release data for all species             catch and hatchery escapement to the Japanese coastal
from British Columbia were provided by Canadian                net fishery and to Hokkaido and Honshu hatcheries,
Fisheries and Oceans (Ted Perry, Canadian Fisheries            respectively.

                                                          39
NPAFC Bulletin No.1                                                                                   Mahnken et al. (1998)

    RESULTS - HISTORICAL PRODUCTION OF                            decline can be attributed to various factors: Survival
       PACIFIC SALMON AT PACIFIC RIM                              of adult hatchery fish declined following the oceanic
                HATCHERIES                                        regime shift in 1976 and a series of EI Nmos events in
                                                                  the late 1970s and early 1980s. These conditions
 Coho Salmon                                                      resulted in reduced escapement, and hatcheries were
                                                                  unable to meet egg needs for full production. More
       Coho salmon are among the most successful of               restrictive inter-basin egg transfer policies were
  hatchery-cultivated species in the Pacific Northwest            introduced to protect remaining wild populations, and
  and Canada. Coho salmon hatcheries were highly                  reduced operating budgets at some hatcheries further
  successful in returning adults to the fisheries in the          reduced coho salmon production.
  1970s, when record smolt-to-adult survivals were
  recorded in the British Columbia and Puget Sound                Chinook salmon
  regions.
       Coho salmon production in western North                           Like coho salmon, chinook salmon are released
 America grew slowly from its inception at the tum of              either as fry or as yearling smolts. Chinook salmon
 the century, and before 1940 the output from                      were the first salmon species to be artificially
 hatcheries never exceeded about 25 million fish                   propagated in western north America and are
 annually (Fig. lA). The slow rate of growth can be                artificially propagated in hatcheries along the eastern
 attributed to failure of hatcheries to contribute to              Pacific seaboard from California to Alaska. More
 expansion of the fishery (McNeil and Bailey 1975).                chinook salmon have been produced from hatcheries
 Following a period of reduced production during                   than any other species in the Pacific Northwest. The
 World War II, coho salmon hatcheries entered an                   first effort to artificially propagate chinook salmon in
 industrial phase of expansion that lasted through the             North America was at the Baird Fish Hatchery on the
 1970s.                                                           McCloud River in California in 1872. This hatchery
      In the 1950s and 1960s, advances in the                     was established to obtain chinook salmon eggs for
 knowledge of feeds, diseases, and the early life history         transport to Atlantic Ocean tributaries to replace
 culture requirements of coho salmon led to improved              depleted Atlantic salmon (Salmo salar) runs. Today,
post-release survival of hatchery fish (Lichatowich and           the center of hatchery production is the Columbia
McIntyre 1987). Coho salmon are released either as                River Basin where approximately 27 % of world
 fry or as yearling smolts (25-30 g) and size at release          chinook salmon is produced.
had also been increased over the years and contributed                  Fall chinook are the most commonly cultured life
to improved adult survival (Wahle and Smith 1979).                history type in both British Columbia and the Pacific
These achievements were even more important in that               Northwest. Also known as "ocean-type" (Healey
they coincided with a period of rapid deterioration of            1991), fall chinook salmon most frequently inhabit
freshwater habitat and blockage of major migratory                coastal rivers, although ocean-type fish are cultured in
pathways by hydroelectric dams.            Given these            the Columbia River as far upstream as the Methow
successes, fishery managers came to believe that                  River. Fall chinook salmon are sometimes released as
hatcheries were a means by which the Pacific                      fry but are more commonly reared for three months
Northwest could continue to develop its water                    and released at hatcheries in the spring at
resources for power, irrigation, and industrial or               approximately 7-10 g. To improve survival, some
domestic use and at the same time, maintain fisheries            hatchery populations of fall chinook are reared to
at historic levels (Lichatowich and McIntyre 1987).              yearling size and generally released in the spring at
Increased reliance on hatchery coho salmon led to the            25-30 g as age-l smolts (Wahle and Smith 1979).
rapid expansion of production through the 1970s. In                     Spring and summer chinook salmon, or "stream-
the late 1970s and early 1980s, private sea ranches              type" life history stocks (Healey 1991), are produced
added 9.3 million smolts per year to Oregon coastal              at hatcheries located primarily on large river systems
production (Fig. lA) and helped lead to a record                 of the Pacific Northwest, British Columbia, and
production of 198 million hatchery coho salmon in                Alaska. Spring chinook salmon are the predominant
1981. In the years that followed however, coho                   stocks produced in Alaskan hatcheries. Spring and
salmon production in the Pacific Northwest stabilized            summer stocks of chinook salmon are seldom released
and began to decline (Fig. lA).                                  as underyearlings and are grown in hatcheries to the
      Begmning in 1989, a period of declining                    largest size, often over 100 g average, of any of the
production began in most sectors of the coho salmon              Pacific salmon species (NRC, 1995). Underyearling
hatchery system. The decline in overall production               releases have been attempted at hatcheries where high
from the contiguous western United States was                    rearing water temperatures result in accelerated
partially offset by the added production of 40 million           growth, but survival to adult from releases of these
hatchery fish from Alaska and British Columbia. This             intermediate sized fish is generally lower than those of

                                                            40
larger yearling fish (Zaugg et al. 1985,                         Coulee dam and other mid-Columbia hydroelectric
1986).                                                           projects. Between 1960 and 1976, 30 hatcheries and
     Hatchery production of chinook sahnon began in              12 rearing ponds raised anadromous sahnonids in this
Washington State in 1895 at the Kalama hatchery on               region (Wahle and Smith 1979). Another surge in
the Columbia River, and production grew slowly to                production occurred in Puget Sound hatcheries in
around 50 million fish released until the late 1930's            Washington State during the 1960s. New lower
(Fig. lB). Production dropped slightly during World              Columbia River hatcheries and growing production in
War II, then accelerated following improvements in               other sectors of the Pacific Northwest drove annual
culture technology and construction of new hatcheries.           releases to more than 300 million chinook salmon
The decade of 1950-1960 began the industrial phase of            smolts by the early 1980s. Production increases from
chinook sahnon hatchery production, as development               British Columbia and Alaska hatcheries in the 1980s
in the Pacific Northwest resulted in loss of freshwater          added another 100 million fish annually. By 1988,
habitat. Growth of the fisheries also added pressure             when production peaked, more than 420 million fry,
to the hatchery system to increase production. New               fingerling, and smolts were being released from
hatcheries on the middle Columbia River were                     eastern Pacific hatcheries, a seven-fold increase from
constructed to mitigate for lost habitat above Grand             the base level of 59 million fish released in 1949.

 Fig. 1 Hatchery production of (A) coho salmon and (B) chinook salmon juveniles from Pacific Northwest, British
        Columbia, and Alaska hatcheries, 1990-1992.

                       200~--------------------------------~--~

                                A                              COHO SALMON

                       150

                               .. PACIFIC NORTHWEST
                               G
               U 100
NPAFe Bulletin No.1                                                                                 Mahnken et al. (1998)

Chinook salmon hatchery production began to decline             Japan. Another interesting feature of the Japanese
in the late 1980s for the same reasons that coho                hatchery system is that it has reduced reliance on high-
salmon production is now falling.                               seas fisheries. The imposition of foreign EEZs has
                                                                restricted the Japanese high-seas fishery on chum
Chum salmon                                                     salmon destined for Russia and North America, but
                                                                the loss to the national fishery has been more than
      From the late 1880s until the recent expansion of         replaced by hatcheries.
 chum salmon ranching in Japan, hatcheries released                  In North America, enhancement efforts for chum
 fry from rearing ponds to the stream soon after yolk           salmon also accelerated in the 1970s, primarily in
 sac absorption.      Egg and fry development was               Alaska and British Columbia (Fig. 2A), adding an
 accelerated in Hokkaido through the use of constant,           additional 850 million fry to the already impressive
 8°C ground water. These condition favored releases             releases by Japan for a total Pacific Rim production of
 from early to mid-February when fry were sometimes             nearly three billion chum salmon fry (exclusive of
 subjected to severe conditions in the streams and              Russian hatchery releases).
 coastal marine waters. With temperatures as low as 0-
SoC during February and March, survival was                     Pink salmon
 minimal.       In 1962, production-scale hatchery
 experiments were undertaken to delay release to a                    Pink salmon are second to chum salmon in the
 time of more favorable sea temperatures. These                 numbers of juveniles released into the North Pacific
 experiments involved feeding up to 300 million fry for         Ocean; accounting for 29% of the total reported in
short periods (Mayama 1985) and demonstrated that                1992 (Heard 1995). Pink salmon are released from
dry diets could increase body weight from 0.6 to 1.0            most hatcheries at 0.5-2.0 g size (Isaksson 1988).
 g in about one month. Fry fed on these diets could be          Alaska is the largest producer of pink salmon and
released from Hokkaido hatcheries in May, when                  released more than 800 million juveniles in 1992
coastal water temperatures exceeded 10°C. These fry             (Heard 1995; McNair 1996). Heard (1995) also notes
are usually released 50 d before average sea surface            that Russia is the second largest producer of hatchery
temperature reaches 15°C. Larger fish of 2-3 g                  pinks; with 584 million juveniles released in 1992.
average weight released .to coastal waters during                    Pink salmon releases remained low (less than 100
spring periods of high primary and secondary                    million) until the early 1980s, when an
productivity survived at a much higher rate than                industrialization period began in Alaska, and numbers
smaller ones. Adult returns from unfed fry released             of released fish from North America increased tenfold
in 1950-60 averaged 1.2%; returns improved to 2.3%              to a total of 1.006 billion in 1992 (Fig. 2B). When
after 1966 as the percentage of fed fish increased              added to Russian production, total world production
(Isaksson 1988).                                                of pink salmon was 1.590 billion produced in 1992.
      Japanese chum salmon catch was high prior to              Pink salmon is the most recent of the Pacific salmon
World War II, but dropped during the war years.                 species to be industrially produced.
Following the war, catch increased for a few years,             Sockeye salmon
then decreased as stocks were overexploited.
Beginning in the 1970s, catch of Asian chum salmon                   Sockeye salmon were propagated in Alaska before
rose again, primarily as a result of massive Japanese           the tum of the century to enhance existing runs, and
hatchery releases. From 1950 until 1970 Japanese                millions of eggs were sent to the Atlantic coast in an
hatchery production rose from 260 million fish                  attempt to establish runs there (Roppel 1982).
released to around 580 million (Fig. 2A). Following             However, initial culture efforts in Alaska were
improvements in adult contribution through the release          unsuccessful, and sockeye salmon programs were
of fed fry, and the loss of foreign fishing grounds to          discontinued early in the 20th century (Allee 1990).
exclusive economic zones (EEZs), the Japanese chum              Similarly, 11 sockeye salmon hatcheries built in
salmon hatchery system entered a phase of rapid                 British Columbia before 1917 produced no consistent
industrialization. Production rose from 260 million             benefits, and production ceased soon thereafter
fry released in 1970 to 2 billion released in 1981.             (Foerster 1968). In Washington, artificial propagation
After the early 1980s, Japanese production of chum              of sockeye salmon began in 1896 at the Baker Lake
salmon leveled off.                                             Station in the Skagit River basin and continued until
      The Japanese hatchery system is the largest in the        this facility was closed in 1933. In addition to
world in terms of fish released and returned 78 million         supplementing the run of sockeye salmon to Baker
adult fish to the Japanese coastal fisheries in 1995            Lake, this facility was the source for the sockeye
(H. Urawa, pers. commun.). The Japanese land-                   salmon introduced into Lake Washington where a
based fishery is almost entirely hatchery-dependant;            strong run was eventually established (Kemmerich
wild stocks are virtually non-existent in northern              1945).

                                                           42
Fig. 2   Hatchery production of (A) chum salmon and (B) pink salmon juveniles from Pacific Northwest, British
           Columbia, Alaska, and Japanese hatcheries, 1950-1993.

                      3000~------------------------------------~

                      2500        A                     CHUM SALMON

                      2000
                                            PACIFIC NORTHWEST
              v(!)    1500         D        BRITISH COLUMBIA
               V')                          ALASKA
               o(!)   1000         I2l      JAPAN

               (!)    500
              0::::
              ..c
              .-
               V')

              UL      1250~--------------------------------------~
              "+-
               o
              C 1000              B                       PINK SALMON

              o
                      750

                      500         o      PACIFIC NORTHWEST
                                  o      BRITISH COLUMBIA
                                  [] ALASKA
                      250         fa     JAPAN

                                     IJ')
                                     IJ')
                                                                                   o        IJ')
                                                                                   CO       CO
                                     0-.                                           0-.      0-.

                                                      Release Year
      Sockeye salmon culture began in the Columbia                programs for any species in the PacIfic Northwest.
River in the 1940s at the Leavenworth Hatchery in                 Recent attempts to reinvigorate sockeye salmon
eastern Washington state. This production effort                  culture program in the Pacific Northwest include a
attempted to mitigate for losses of sockeye due to                combined hatchery and net-pen culture system
construction of hydroelectric dams on the middle                  operated by the State of Washington in Lake
Columbia River (Mullan 1986).            Smolts were              Wenatchee and a recovery program for the
           for a period of about 20 years, but disease            endangered Redfish Lake sockeye salmon in Idaho's
and low returns forced abandonment of the program                 Stanley Basin (Flagg et al. 1991, 1995).
     the 1960s. Small smolt and fry-release programs                   British Columbia is now by far the largest
still exist for sockeye in Puget Sound and on the                 producer of artificially propagated sockeye salmon,
Washington coast, but by and large, sockeye salmon                producing more than 290 million fry in 1993 (Fig.
culture in the Pacific Northwest is insignificant.                3A).      Fry are produced ill spawning channels
Along with pink salmon, sockeye salmon production                 containing gravel substrates, where returning adults
constitutes one of the smallest artificial propagation            spawn naturally.       Fry are allowed to migrate

                                                            43
NPAFCBulletinNo.l                                                                                    Mahnken et al. (1998)

volitionally out of the channels upon swimup, usually           75 million smolts in 1994, or 21 % of total North
into a lake. Spawning-channel culture of sockeye                American production (Fig. 3A). Alaska also releases
salmon in British Columbia constitutes the least                age-O sockeye after brief holding in sea-pens.
invasive culture technique used to mass culture Pacific
salmon.      These extensive artificial propagation             Steelhead and masu salmon
techniques require no feed, use natural spawning
substrates, and require no handling of either juveniles              Hatchery steelhead are produced entirely in North
or adults during rearing. Although spawning-channel             America (Fig. 3B), while masu salmon are only
culture of sockeye was initiated prior to the 1950s, the        produced in significant numbers in Japan (Fig. 4). In
pro gram did not accelerate until the 1960s when                terms of total production, steelhead and masu salmon
output increased from 2 million fry in 1960 to 258              are minor species, but in terms of their contribution to
million in 1980 (Fig. 3A).                                      regional fisheries, steelhead programs are important
     The Alaskan hatchery system is now the major               and produced an estimated 738,000 adults annually
producer of sockeye smolts in North America, and                from 1978 to 1987 (Light 1989). Like coho salmon
although a recent entrant (1974) into large-scale               and chinook salmon, steelhead coho salmon and
sockeye production, has grown rapidly and produced              chinook salmon, steelhead coho salmon and chinook

  Fig. 3   Hatchery production of (A) sockeye salmon and (B) steelhead salmon juveniles from Pacific Northwest, British
           Columbia, and Alaska hatcheries and spawning channels, 1900·1992.

                   400

                   300
                             A                    SOCKEYE SALMON

                              [] PACIFIC NORTHWEST
                   200        fIl BRITISH COLUMBIA
                              •     ALASKA

                   100

                     O+---~-'~~-=F=~~~~--~
                   40~----------------------------------------~

             V')
             C                B                    STEElHEAD TROUT
             o     30

                               [J PACIFIC NORTHWEST
                   20          1m   BRITISH COLUMBIA
                               •    ALASKA

                    10

                    O~~~~~~~~~~~~~~~~~~~~
                                o                                o
                         8
                         ~      ~
                                                                 ll')
                                                                 ~

                                                 Release Year
                                                           44
Fig. 4   Production of masu salmon juveniles from Japanese hatcheries, 1960-1992.

                      20

              "'0                                          MASU SALMON
Fig.5 Total hatchery production of coho, steelhead, masu, sockeye, chinook, pink, and chum salmon juveniles from the
        Pacific Northwest, British Columbia, Alaska, and Japan, 1950-1992.

                  5000.-------------------------------------~
                                                      ALL SPECIES
           "'0
Fig. 6   Mean survival (coded-wire tags) of coho salmon released from Pacific coast hatcheries 1970-1990. (A) Georgia
          Strait estuary, coastal regions of north British Columbia/outer Vancouver IslandlWashington, and coastal regions
          of Oregon/California. (B) Puget Sound and the Columbia River. Error bars are 1 standard error. Asterisks denote
          differences (P
                .-
                 >
                 ~

                 ~
                (/')
                -+-      0
                 C      15
                 (l)
                                --0-
                                __......_-   PUGET SOUND
                 U
                 ~
                 (l)
                                .
                                             COLUMBIA RIVER
                                                                                          B
                0-
                        10

                                                           Year
stock survivals fluctuate widely and exhibit a severe               Strait of Georgia, Puget Sound, and southeast Alaska.
decline in the 1980s, they remained more than twice                 These areas are known to be excellent areas for
as productive (approximately 5 %) as the coastal stocks             juvenile rearing and for survival of sahnonids (Healey,
(approximately 2%) until 1987. After 1987, all stocks                1980; Simenstead et aI., 1982), with relatively high
declined to survivals around 2 %.                                   survivals of 5.5-7.5% during this period.
     A comparison of bi-decadal mean survival by
region revealed another interesting feature of hatchery             Fall chinook salmon
coho sahnon releases from 1970-90 (Fig.            When
arranged in order of declining                                           Fan chinook sahnon exhibit               different
latItudinal clines appeared among the coastal stocks,               survival characteristics from coho sahnon. Because of
both north and south from the center of the geographic              smaller size at release, surVIval of hatchery fall
distributIon of the species, at about the latitude of               chinook sahnon is less than that of coho sahnon.
Vancouver Island. This region had the highest                       Furthermore, survivals and trends in survival are
survival (4%), with coastal California and coastal                  distinctly different between fall chinook sahnon
Alaska (at either end of the geographic           having            ._...._.J stocks north of        Sound and those to the
the lowest survivals, at 1-1. 5 %. Latitudinal anomolies            south         8). SurVIvals           in Puger Sound,
were observed in large coastal estuaries such as the                Strait of Georgia and outer Vancouver Island ill the

                                                              47
NPAFC Bulletin No.1                                                                                                 Mahnken et al. (1998)

 Fig. 7 Mean survival of hatchery coho salmon by region, 1970-1990.

                      9,--------------,-------------------------,
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                      7-
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                                                                                  Coastal

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mid-1970s at between 3-4%, then declined sharply to                       satiation of predators at higher levels of juvenile
less than 2 % in the 1980s following the EI Nifio of                      production. However, Kaeriyama (1996) presented
1977 (Fig. 8A). A further decline to about 0.5%                           evidence that other life history characteristics of
survival followed the 1983 EI Nifio and survival has                      enhanced chum salmon, namely decreased body size
continued downward since. Stocks south of Puget                           and an increase of age-at-maturity may indicate the
Sound showed the opposite trend, with survival rates                      beginning of a density-dependant effect of continued
of less than 1 % through 1982 rising to around 2%                         large-scale releases from Japanese hatcheries.
following the 1983 El Nifio (Fig. 8B). It appears that
the 1983 El Nifio acted to enhance survival of                                                      DISCUSSION
southern fall chinook salmon in the mid-1980s rather
than to decrease survival. However, mean survival of                           Researchers from North America and Japan have
fall chinook salmon, aggregated by region, failed to                      noted the dramatic decline in salmon stock abundance
show the same geographic cline as that of coho                            and body size in the southern portion of the species
salmon.                                                                   range in North America. These declines have been
                                                                          especially apparent over the past two decades (NeWsen
Chum salmon                                                               et al. 1991; Bigler et al. 1996; Ricker 1981).
                                                                          However, while the abundance of stocks in the Pacific
      Survival of chum salmon released from Hokkaido                      Northwest has declined, the abundance of populations
hatcheries has followed a trend that appears to be less                   to the north, both in Asia and North America, remain
influenced by ocean conditions and more by                                healthy and some have reached historical highs (Heard
improvements in hatchery technology (Figure 9).                           1995; Kaeriyama and Urawa 1992; Burger and
Survival of Hokkaido chum salmon has risen                                Wertheimer 1995; Zorpette 1995). Attempts have
uniformly from 2.5 to 4% from 1965-1988, a period                         been made to assess the cause of these declines based
in which production increased from 550 to 970 million                     on changes in freshwater conditions, fishing, or on
fry released. The number of juveniles released and                        variations in the marine environment (Beamish and
their survival both increased with time. This apparent                    Bouillon 1993; Cooper and Johnson 1992; Johnson
inverse density-dependant survival has been noted by                      1984; Lawson 1993; Lichatowich 1993; Nickelson
McNeil (1991) who suggests that the relationship may                      1986; Northcote and Atagi 1994; Pearcy 1992;
be an artifact of improved hatchery technology, or                        Richards and Olsen 1993; Olsen and Richards 1994;

                                                                  48
Fig.8 Mean survival (coded-wire tags) of fall chinook salmon released from Pacific coast hatcheries 1970-1990. (A) Puget
      Sound, Strait of Georgia, and outer Vancouver Island regions. (B) upper and lower Columbia River, and Washington
      and Oregon coastal regions. Error bars are omitted for simplicity. Asterisks denote differences (P
            .-
             >
             I.-
             ::)
            V)         0
            -+-        3
             C
             Q)
             U 2.5
             I.-
                                  B
            Q)                    ----.---   UPPER COLUMBIA RIVER
            CL         2
                                  --0--      OREGON COASTAL
                                  -a-        WASHINGTON COASTAL
                     1.5          --+---     LOWER COLUMBIA R.

                     0.5

                                                           o                l.{')             o
                                                           CO               CO                0-..
                                                           0-..             0-..              0-..

                                                    Year
Fig.9 Mean survival of chum salmon released from Japanese hatcheries in Hokkaido, 1963-1988.
                       5
                                             HOKKAIDO CHUM SALMON
               0       4
              .2:
               2:
               ::J     3

              -
              C/)

               C
               ())
               U
                      2
               ~
               ())
              D-

                      O
                           (")                                                              CO
                           '0                                                               CO
                           0-..                                                             0-..

                                                   Brood Year
                                                          49
NPAFe Bulletin No.1                                                                                  Mahnken et al. (1998)

 Francis and Sibley 1991; Kaeriyama 1996). Most                   Nino events is most evident in regions where survival
 researchers have concluded that, whatever the major              has been historically high (southeast Alaska and
 factor(s) affecting survival of Pacific salmonids, they         Georgia Strait following the 1982-83 EI Nino, Puget
 are most likely to occur in the ocean environment.               Sound following the 1976-77 EI Nino). In the past 40
       Cooper and Johnson (1992), compared trends in              years, nine EI Ninos have affected the coastal regions
 abundance of Washington, Oregon, and British                    of the eastern Pacific. In the 1970s and 1980s, the
 Columbia steelhead and concluded that there were                coastal regions of the Pacific Northwest and Canada
similarities in trends over the entire geographic range          have been beset by a series of four moderate-to-strong
 that indicated common factors were responsible for the          El Nino events, most notably the 1982-83 EI Nino,
 observed changes in survival. Because freshwater,                which by many measures was the strongest this
estuarine, and nearshore conditions differ considerably          century. Since 1970, El Ninos have occurred in 1972-
 from year to year within this region, they concluded            73, 1976-77, 1982-83, and 1987-88.
that these factors alone could not explain the                         A strong negative cline in survival of coho salmon
similarities in steelhead survivals. They suggested              is observed moving both north and south from the
 that similarities in steelhead abundance trends in              center of the species distribution in British Columbia,
widely separated geographical regions indicated that             with stocks in western Alaska and California
common factors were responsible for the observed                 exhibiting the lowest survival. Fall chinook salmon,
declines, and that oceanic conditions were responsible.          on the other hand, do not show the same strong
Olsen and Richards (1994) came to a similar                      latitudinal cline.
conclusion while working with aggregated coastwide                     Fall chinook salmon survival, although apparently
chinook salmon production data, namely that similar              also affected by EI Nino events, seems directed by
chinook salmon run-size trends can be observed                   other external factors. Stock survival north of the
between several west coast river basins, and that the            Columbia River peaked in the mid 1970s, while
data support the hypothesis that ocean conditions have           survival for regions south of the river peaked in the
had a marked and uniform impact on chinook salmon                mid-1980s. Furthermore, mean survival of fall
production in the Pacific Northwest. Lichatowich                 chinook salmon aggregated by region failed to show
(1993) has pointed out that the magnitude of oceanic             the same geographic cline as coho salmon. The Strait
environmental changes and their impacts on salmon               of Georgia estuary produced highest survivals, but
survival may be so large as to mask changes that                Puget Sound fall chinook salmon did not produce
occur in the freshwater habitat. He cautioned that this         higher survivals than Coastal Oregon, and produced
may cause managers to falsely attribute increased               only slightly better survival than outer Vancouver
ocean survival to restoration effects in freshwater.            Island or coastal California. This is surprising, given
Hilborn et al. (1993) further emphasizes the same               the well-documented importance of estuaries for
point by stating that attempts to understand the impact         growth and survival of juvenile chinook salmon
of in-river (Columbia River) actions on survival will           (Healey 1991; McCabe et al. 1986). Nevertheless,
be confounded by changes in ocean conditions.                   there is some indication that hatchery fall chinook
      Coded-wire tag data shows that for the period             salmon juveniles spend less time in estuaries than wild
1970-1990, coho salmon adult survival was highest for           juveniles which may reduce the benefits of such areas
stocks released into large coastal estuaries. Survival          to artificially propagated fish (Levings et al. 1986). It
in these estuaries is typified by widely fluctuating            may be that the overall lower survival of fall chinook
mean survivals. Conversely, survival of hatchery                salmon masks regional geographic differences so
coho salmon released into coastal regions that lack             evident with coho salmon.
protective coastal estuaries is typified by lower, more               It is tempting to postulate a cause and effect
constant survival. However, differences in survival             relationship between the occurrence of EI Nino events
between estuarine and coastal releases of fall chinook          and declines in survival of hatchery fish in the eastern
salmon are not as dramatic, with regions like outer             Pacific, but no convincing ecological relationship
Vancouver Island and coastal Oregon performing as               exists. Climate conditions are known to have changed
well or better than Puget Sound and the Strait of               recently in the Pacific Northwest. Most Pacific
Georgia. It is possible that such factors as size and           salmonid stocks south of British Columbia have been
orne of entry to seawater, location and length of time          affected by changes in ocean production that occurred
in estuaries prior to outmigration, and predation may           during the 1970s. Pearcy (1992) and Lawson (1993)
influence differences in absolute survival and temporal         attribute this decline largely to ocean factors, but do
trends between the species.                                     not identify specific effects. However, given the
       Coho salmon survivals were depressed following           increased frequency of EI Nino events in the past two
the unusually strong El Nino events of the past two             decades, and large-scale secular warming of the region
decades (Fig. 6) and continued to decline throughout            (Freeland 1990), it is certainly plausible that there is
the 1980s. Depressed survival associated with El                at least some response to EI Nino events in the form

                                                           50
of reduced survival of the species.                                                REFERENCES
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                                                         53
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