Lack of trophic competition among wild and hatchery juvenile chum salmon during early marine residence in Taku Inlet,Southeast Alaska

Early marine trophic interactions of wild and hatchery chum salmon (Oncorhynchus keta) were examined as a potential cause for the decline in harvests of adult wild chum salmon in Taku Inlet, Southeast Alaska. In 2004 and 2005, outmigrating juvenile chum salmon were sampled in nearshore habitats of the inlet (spring) and in epipelagic habitat at Icy Strait (summer) as they approached the Gulf of Alaska. Fish were frozen for energy density determination or preserved for diet analyses, and hatchery stocks were identified from the presence of thermal marks on otoliths. We compared feeding intensity, diets, energy density, and size relationships of wild and hatchery stocks (n = 3123) across locations and weeks. Only hatchery fish feeding intensity was negatively correlated with fish abundance. In both years, hatchery chum salmon were initially larger and had greater energy density than wild fish, but lost condition in early weeks after release as they adapted to feeding on wild prey assemblages. Diets differed between the stocks at all inlet locations, but did not differ for hatchery salmon between littoral and neritic habitats in the outer inlet, where the stocks overlapped most. Both diets and energy density converged by late June. Therefore, if density-dependent interactions affect wild chum salmon, these effects must be very rapid because survivors in Icy Strait showed few differences. Our study also demonstrates that hatchery release strategies used near Taku Inlet successfully promote early spatial segregation and prey partitioning, which reduce the probability of competition between wild and hatchery chum salmon stocks.     

The occurrence and run timing of naturally spawning chum salmon in northern Japan

Since the late 20th century, the biomass of Pacific salmon Oncorhynchus spp. has increased. Hokkaido, northern Japan, is one of the main areas of chum salmon O. keta production in the North Pacific and intensive hatchery programs support the recent high abundance. However, proper management of naturally spawning populations is necessary to conserve healthy stocks of this species. In 2008, we started a program to assess the naturally spawning chum salmon populations in Hokkaido. Of the total of approximately 1,500 rivers in Hokkaido, 238 rivers with lengths of longer than 8 km (excluding those rivers used for hatchery broodstock collection) were surveyed in 2008 and 2009. The number of non-enhanced rivers found to contain naturally reproducing chum salmon was 59 (31.4% of surveyed rivers) and 50 (37.6% of surveyed rivers) rivers in 2008 and 2009, respectively. Including the rivers where hatchery broodstock were collected and rivers shorter than 8 km that contain naturally spawning chum salmon, chum salmon ascended at least 191 and 175 rivers in Hokkaido in 2008 and 2009, respectively. Repeated foot surveys indicated that the run timings of naturally spawning chum salmon may be affected by coastal commercial fisheries. This study showed that naturally spawning chum salmon remain in many rivers in Hokkaido where hatchery programs have been intensively conducted.     

Wild and hatchery reproduction of pink and chum salmon and their catches in the Sakhalin-Kuril region,Russia

In the Sakhalin-Kuril region hatchery culture of pink and chum salmon is of great importance compared to other regions of the Russian Far East. During the last 30 years the number of hatcheries increased two-fold, and significant advances were made in hatchery technologies. As a result, chum salmon capture in regions where hatcheries operate (southwestern and eastern Sakhalin coasts, and Iturup Island) was 9 times as high during 2006–2010 than during 1986–1990, whereas wild chum salmon harvest markedly declined. Recent dynamics in pink salmon catch appear to track trends in natural spawning in monitored index rivers, suggesting natural-origin pink salmon play a dominant role in supporting the commercial fishery. It remains uncertain as to whether hatcheries have substantially supplemented commercial catch of pink salmon in this region, and I recommend continued research (including implementing mass marking and recovery programs) before decisions are made regarding increasing pink salmon hatchery production. Location of hatcheries in spawning river basins poses problems for structuring a management system that treats hatchery and wild populations separately. Debate continues regarding the existence and importance of density-dependent processes operating in the ocean environment and the role hatcheries play in these processes. Loss of critical spawning habitat for chum salmon in the Sakhalin-Kuril region has lead to significant declines in their abundance. I conclude by recommending increases in releases of hatchery chum salmon numbers in the region to help recover depressed wild populations and provide greater commercial fishing benefits in the region.     

Relationship of size at return with environmental variation,hatchery production,and productivity of wild pink salmon in Prince William Sound,Alaska: does size matter?

Pink salmon (Oncorhynchus gorbuscha) returning to Prince William Sound (PWS), Alaska, have increased to historically high levels of abundance in recent years, but average body size at return has declined. We examined how body size at return of PWS pink salmon was related to 10 biophysical factors, including the scale of hatchery production. We also examined the effect of body size at return on productivity of wild pink salmon in PWS. For the 1975–1999 brood years, we found that an index of total abundance of pink salmon in the Gulf of Alaska and sea surface temperature during the year of return best explained the variation in pink salmon body size over time. Body size at return was significantly correlated with productivity of wild pink salmon. We used stepwise-regression to fit a generalized linear version of the Ricker spawner-recruit model to determine if body size would explain significant variation in wild-stock productivity in context with other environmental variation, including hatchery production. The results indicate that variability in wild-stock productivity is primarily driven by density-independent factors in the marine environment, but that body size of wild spawners also significantly affects productivity of wild PWS pink salmon. We conclude that the success of large-scale enhancement increasing the total run in PWS may have contributed to the decline in body size because of density-dependent growth in the Gulf of Alaska. We used a simulation model to estimate the impact of hatchery-induced changes in adult body size on wild-stock production in PWS. We estimated an annual wild-stock yield loss of 1.03 million pink salmon, less than 5% of the annual hatchery return of 24.2 million adult pink salmon for brood years 1990–1999.     

Straying of hatchery salmon in Prince William Sound,Alaska

The straying of hatchery salmon may harm wild salmon populations through a variety of ecological and genetic mechanisms. Surveys of pink (Oncorhynchus gorbuscha), chum (O. keta) and sockeye (O. nerka) salmon in wild salmon spawning locations in Prince William Sound (PWS), Alaska since 1997 show a wide range of hatchery straying. The analysis of thermally marked otoliths collected from carcasses indicate that 0–98% of pink salmon, 0–63% of chum salmon and 0–93% of sockeye salmon in spawning areas are hatchery fish, producing an unknown number of hatchery-wild hybrids. Most spawning locations sampled (77%) had hatchery pink salmon from three or more hatcheries, and 51% had annual escapements consisting of more than 10% hatchery pink salmon during at least one of the years surveyed. An exponential decay model of the percentage of hatchery pink salmon strays with distance from hatcheries indicated that streams throughout PWS contain more than 10% hatchery pink salmon. The prevalence of hatchery pink salmon strays in streams increased throughout the spawning season, while the prevalence of hatchery chum salmon decreased. The level of hatchery salmon strays in many areas of PWS are beyond all proposed thresholds (2–10%), which confounds wild salmon escapement goals and may harm the productivity, genetic diversity and fitness of wild salmon in this region     

Perspectives on wild and hatchery salmon interactions at sea,potential climate effects on Japanese chum salmon,and the need for sustainable salmon fishery management reform in Japan

Pacific salmon (Oncorhynchus spp.) play an important role as a keystone species and provider of ecosystem services in the North Pacific ecosystem. We review our studies on recent production trends, marine carrying capacity, climate effects and biological interactions between wild and hatchery origin populations of Pacific salmon in the open sea, with a particular focus on Japanese chum salmon (O. keta). Salmon catch data indicates that the abundance of Pacific salmon increased since the 1976/77 ocean regime shift. Chum and pink salmon (O. gorbuscha) maintained high abundances with a sharp increase in hatchery-released populations since the late 1980s. Since the 1990s, the biomass contribution of hatchery returns to the total catch amounts to 50% for chum salmon, more than 10% for pink salmon, and less than 10% for sockeye salmon (O. nerka). We show evidence of density-dependence of growth and survival at sea and how it might vary across spatial scales, and we provide some new information on foraging plasticity that may offer new insight into competitive interactions. The marine carrying capacity of these three species is synchronized with long-term patterns in climate change. At the present time, global warming has positively affected growth and survival of Hokkaido populations of chum salmon. In the future, however, global warming may decrease the marine carrying capacity and the area of suitable habitat for chum salmon in the North Pacific Ocean. We outline future challenges for salmon sustainable conservation management in Japan, and recommend fishery management reform to sustain the hatchery-supported salmon fishery while conserving natural spawning populations.     

A global assessment of salmon aquaculture impacts on wild salmonids

Since the late 1980s, wild salmon catch and abundance have declined dramatically in the North Atlantic and in much of the northeastern Pacific south of Alaska. In these areas, there has been a concomitant increase in the production of farmed salmon. Previous studies have shown negative impacts on wild salmonids, but these results have been difficult to translate into predictions of change in wild population survival and abundance. We compared marine survival of salmonids in areas with salmon farming to adjacent areas without farms in Scotland, Ireland, Atlantic Canada, and Pacific Canada to estimate changes in marine survival concurrent with the growth of salmon aquaculture. Through a meta-analysis of existing data, we show a reduction in survival or abundance of Atlantic salmon; sea trout; and pink, chum, and coho salmon in association with increased production of farmed salmon. In many cases, these reductions in survival or abundance are greater than 50%. Meta-analytic estimates of the mean effect are significant and negative, suggesting that salmon farming has reduced survival of wild salmon and trout in many populations and countries.     

Migration of Pacific Rim Chum Salmon on the High Seas: Insights from Genetic Data

Wild stocks of chum salmon, Oncorhynchus keta, have experienced recent declines in some areas of their range. Also, the release of hatchery chum salmon has escalated to nearly three billion fish annually. The decline of wild stocks and the unknown effects of hatchery fish combined with the uncertainty of future production caused by global climate change have renewed interest in the migratory patterns of chum salmon on the high seas. We studied the composition of high-seas mixtures of maturing and immature individuals using baseline data for 20 allozyme loci from 356 populations from throughout the Pacific Rim. Composition estimates were made from three time series. Two of these time series were from important coastal migratory corridors: the Shumagin Islands south of the Alaska Peninsula and the east coast of the Kamchatka Peninsula. The third was from chum salmon captured incidentally in the Bering Sea trawl fishery for walleye pollock. We also analyzed geographically dispersed collections of chum salmon captured in the month of July. The time series show dynamic changes in stock composition. The Shumagin Island corridor was used primarily by Northwest Alaskan and Asian populations in June; by the end of July stocks from the Alaska Peninsula and southern North America dominated the composition. The composition along the Kamchatka coast changed dramatically from primarily Russian stocks in May to primarily Japanese stocks in August; the previously undocumented presence of stocks from the Alaska Peninsula and Gulf of Alaska was also demonstrated. Immature chum salmon from throughout the Pacific Rim, including large proportions of southern North American stocks, contributed to the Bering Sea bycatch during the months of September and October. The migration routes of North American stocks is far more widespread than previously observed, and the Bering Sea is an important rearing area for maturing and immature chum salmon from throughout the species' range.     

Overview of salmon stock enhancement in southeast Alaska and compatibility with maintenance of hatchery and wild stocks

Modern salmon hatcheries in Southeast Alaska were established in the 1970s when wild runs were at record low levels. Enhancement programs were designed to help rehabilitate depressed fisheries and to protect wild salmon stocks through detailed planning and permitting processes that included focused policies on genetics, pathology, and management. Hatcheries were located away from significant wild stocks, local sources were used to develop hatchery broodstocks, and juveniles are marked so management can target fisheries on hatchery fish. Initially conceived as a state-run system, the Southeast Alaska (SEAK) program has evolved into a private, non-profit concept centered around regional aquaculture associations run by fishermen and other stakeholders that pay for hatchery operations through landing fees and sale of fish. Today there are 15 production hatcheries and 2 research hatcheries in SEAK that between 2005 and 2009 released from 474 to 580 million (average 517 million) juvenile salmon per year. During this same period commercial harvest of salmon in the region ranged from 28 to 71 million salmon per year (average 49 million). Contributions of hatchery-origin fish to this harvest respectively averaged 2%, 9%, 19%, 20%, and 78% for pink, sockeye, Chinook, coho, and chum salmon. Both hatchery and wild salmon stocks throughout much of Alaska have experienced high marine survivals since the 1980s and 1990s resulting in record harvests over the past two decades. Although some interactions between hatchery salmon and wild salmon are unavoidable including increasing concerns over straying of hatchery fish into wild salmon streams, obvious adverse impacts from hatcheries on production of wild salmon populations in this region are not readily evident.     

Understanding the adaptive consequences of hatchery-wild interactions in Alaska salmon

About 31% of salmon harvested in Alaska comes from the hatchery production of hundreds of millions of pink and chum salmon and smaller numbers of sockeye, Chinook, and coho salmon. The numbers of hatchery-reared juveniles released in some areas are greater than the numbers of juveniles from wild populations. However, virtually nothing is known about the effects of hatchery fish on wild populations in Alaska. Possible effects of these interactions can be inferred from studies of salmonids in other areas, from studies of other animals, and from theory. Numerous studies show a complex relationship between the genetic architecture of a population and its environment. Adaptive responses to nature and anthropogenic selection can be influenced by variation at a single gene, or more often, by the additive effects of several genes. Studies of salmonids in other areas show that hatchery practices can lead to the loss of genetic diversity, to shifts in adult run timing and earlier maturity, to increases in parasite load, to increases in straying, to altered levels of boldness and dominance, to shifts in juvenile out-migration timing, and to changes in growth. Controlled experiments across generations show, and theory predicts, that the loss of adaptive fitness in hatchery salmon, relative to fitness in wild salmon, can occur on a remarkably short time scale. All of these changes can influence survival and impose selective regimes that influence genetically based adaptive traits. The preservation of adaptive potential in wild populations is an important buffer against diseases and climate variability and, hence, should be considered in planning hatchery production levels and release locations. The protection of wild populations is the foundation for achieving sustained harvests of salmon in Alaska.     

Interregional differentiation of chum salmon from Sakhalin and South Kurils infered from microsatellite markers

Variability at ten microsatellite loci was examined in wild and hatchery populations of chum salmon from the Sakhalin Island and Southern Kuril Islands, Iturup and Kunashir. Substantial genetic differences between Sakhalin and South Kurils chum salmon (the differentiation theta reached 6.0%) were revealed. Statistically significant differences between chum salmon from Iturup and that from Kunashir were demonstrated, as well as between the chum salmon populations from different rivers within the islands. It was shown that in different types of population comparisons, required different marker sets most informative were.     

Mitochondrial DNA variation and genetic population structure of chum salmon Oncorhynchus keta around the Pacific Rim

A survey of mtDNA variation among populations of chum salmon Oncorhynchus keta around the Pacific Rim revealed four large population groups: Rim of the Sea of Japan, the Rim of the Okhotsk Sea and West Bering Sea, North‐west Alaska and Gulf of Alaska. The observed population structure appears to reflect isolation by distance with limited gene flow between regions and larger amounts of gene flow between populations within these four regions.     

Some consequences of Pacific salmon hatchery production in Kamchatka: changes in age structure and contributions to natural spawning populations

Aggregate hatchery production of Pacific salmon in the Kamchatka region of the Russian Federation is very low (< 0.5% of total harvest, with five hatcheries releasing approximately 41 M juvenile salmon annually), but contributions in certain rivers can be substantial. Enhancement programs in these rivers may strongly influence fitness and production of wild salmon. In this paper we document significant divergence in demographic traits in hatchery salmon populations in the Bolshaya River and we estimate the proportion of hatchery chum salmon in the total run in the Paratunka River to demonstrate the magnitude of enhancement in this system. We observed a reduction in the expression of life history types in hatchery populations (ranging from 1 to 9 types) compared to wild populations (17 types) of sockeye salmon in the Bolshaya River. We found similar trends in Chinook salmon in the same river system. This reduced life history diversity may make these fish less resilient to changes in habitat and climate. We estimate hatchery chum salmon currently contribute 17-45% to the natural spawning population in the Paratunka River. As hatchery fish increase in numbers at natural spawning sites, this hatchery production may affect wild salmon production. It is important to investigate the risk of introgression between hatchery and wild salmon that can lead to reduction in salmon fitness in Kamchatka rivers, as well as the potential of ecological interactions that can have consequences on status of wild salmon and overall salmon production in this region.     

Interregional differentiation of chum salmon from Sakhalin and South Kurils inferred from microsatellite markers

Variability at ten microsatellite loci was examined in wild and hatchery populations of chum salmon (Oncorhynchus keta Walbaum) from the Sakhalin Island and Southern Kuril Islands, Iturup and Kunashir. Substantial genetic differences between Sakhalin and South Kurils chum salmon (the differentiation reached 6.0%) were revealed. Statistically significant differences between chum salmon from Iturup and those from Kunashir were demonstrated, as well as between the chum salmon populations from different rivers within the islands. It was shown that in different types of population comparisons, different marker sets were most informative.     

Rapid expansion of an enhanced stock of chum salmon and its impacts on wild population components

A harvested stock of chum salmon homing to Kurilskiy Bay, Iturup Island, consists of two genetically distinct river populations that reproduce in two rivers that drain into the bay and are characterized by limited gene flow. One of these is small and can be regarded as wild, whereas the other is much larger and, until recently, was composed of naturally reproducing components spawning in the river??s mainstem and tributaries, with almost no hatchery reproduction during the past two decades. The only human impact on reproduction of the chum salmon stock was regulation of the escapement, with officially accepted limits to avoid ??over-escapement??. Recently the hatchery began to release a large amount of chum salmon juveniles. As confirmed by data on variation in both age composition and microsatellite DNA, first-generation hatchery-origin fish that returned from the first large releases occupied spawning grounds and presumably competed directly with, and potentially displaced wild fish. The most dramatic example is a genetically distinct beach-spawning form of chum salmon that was swamped by much more numerous hatchery-origin fish of the river-spawning form. In order to restore and support naturally reproduced population components, careful estimation of the carrying capacity of natural spawning grounds is necessary with efforts to increase escapement to these habitats. We also recommend concerted efforts to restore and conserve a unique beach-spawning population of chum salmon. We further recommend development of a marking program for direct estimation of straying and evaluation of ecological and genetic impacts of hatchery fish on neighboring wild and natural populations.     

Juvenile chum salmon consumption of zooplankton in marine waters of southeastern Alaska: a bioenergetics approach to implications of hatchery stock interactions

Bioenergetics modeling was used to estimate zooplankton prey consumption of hatchery and unmarked stocks of juvenile chum salmon (Oncorhynchus keta) migrating seaward in littoral (nearshore) and neritic (epipelagic offshore) marine habitats of southeastern Alaska. A series of model runs were completed using biophysical data collected in Icy Strait, a regional salmon migration corridor, in May, June, July, August, and September of 2001. These data included a temperature (1-m surface versus surface to 20-m average), zooplankton standing crop (surface to 20-m depth versus entire water column), chum salmon diet (percent weight of prey type consumed), energy densities, and weight. Known numbers of hatchery releases were used in a cohort reconstruction model to estimate total abundance of hatchery and wild chum salmon in the northern region of southeastern Alaska, given average survival to adults and for two different (low and high) early marine littoral mortality rate assumptions. Total prey consumption was relatively insensitive to temperature differences associated with the depths potentially utilized by juvenile chum salmon. However, the magnitudes and temporal patterns of total prey consumed differed dramatically between the low and high mortality rate assumptions. Daily consumption rates from the bioenergetics model and CPUE abundance from sampling in Icy Strait were used to estimate amount and percentage of zooplankton standing crop consumed by mixed stocks of chum salmon. We estimated that only a small percentage of the available zooplankton was consumed by juvenile chum salmon, even during peak abundances of marked hatchery and unmarked mixed stocks in July. Total daily consumption of zooplankton by all stock groups of juvenile chum salmon was estimated to be between 330 and 1764 g/km²d¹ from June to September in the neritic habitat of Icy Strait. As with any modeling exercise, model outputs can be misleading if input parameters and underlying assumptions are not valid; therefore, additional studies are warranted, especially to determine physiological input parameters, and to improve abundance and mortality estimates specific to juvenile chum salmon. Future bioenergetics modeling is also needed to evaluate consumption by the highly abundant, vertically migrating planktivorous that co-occurred in our study; we suggest that these fishes have a greater impact on the zooplankton standing crop in Icy Strait than do hatchery stock groups of juvenile chum salmon.     

Genetic stock identification of overwintering chum salmon in the North Pacific Ocean

Understanding stock and age-specific seasonal migrations of Pacific salmon during ocean residence is essential to both the conservation and management of this important resource. Based upon 11 microsatellites assayed on 265 individuals collected aboard international research surveys during winter 2009, we found substantial differences in the age-specific origin of chum salmon (Oncorhynchus keta) in the North Pacific Ocean. Overall, Asian stocks dominated the collections, however, ocean age 1 fish were primarily of Japanese origin and ocean age 2–3+ fish were predominantly of Russian origin. These results suggest that cohorts of chum salmon stocks migrate nonrandomly in the North Pacific Ocean and adjacent seas.     

Interaction between hatchery and wild Pacific salmon in the Far East of Russia: A review

We review studies of interactions between hatchery and wild Pacific salmon in the Russian Far East. This includes the role of hatchery practices that result in premature migration to the sea and increased mortality, and data on feeding and territorial competition between juveniles of hatchery and wild origin. In the course of downstream migration many juvenile hatchery salmon are eliminated by wild salmon predation. During the marine period, Japanese hatchery chum salmon (Oncorhynchus keta) distribution overlaps the distribution of Russian wild salmon. Consequently, replacement of wild populations by hatchery fishes, as a result of abundant juvenile hatchery releases combined with extensive poaching in spawning grounds, is apparent in some Russian rivers. Interactions between the populations occur in all habitats. The importance of conservation of wild salmon populations requires a more detailed study of the consequences of interactions between natural and artificially reared fishes.     

Source-Sink Estimates of Genetic Introgression Show Influence of Hatchery Strays on Wild Chum Salmon Populations in Prince William Sound,Alaska

The extent to which stray, hatchery-reared salmon affect wild populations is much debated. Although experiments show that artificial breeding and culture influence the genetics of hatchery salmon, little is known about the interaction between hatchery and wild salmon in a natural setting. Here, we estimated historical and contemporary genetic population structures of chum salmon (Oncorhynchus keta) in Prince William Sound (PWS), Alaska, with 135 single nucleotide polymorphism (SNP) markers. Historical population structure was inferred from the analysis of DNA from fish scales, which had been archived since the late 1960’s for several populations in PWS. Parallel analyses with microsatellites and a test based on Hardy-Weinberg proportions showed that about 50% of the fish-scale DNA was cross-contaminated with DNA from other fish. These samples were removed from the analysis. We used a novel application of the classical source-sink model to compare SNP allele frequencies in these archived fish-scales (1964–1982) with frequencies in contemporary samples (2008–2010) and found a temporal shift toward hatchery allele frequencies in some wild populations. Other populations showed markedly less introgression, despite moderate amounts of hatchery straying. The extent of introgression may reflect similarities in spawning time and life-history traits between hatchery and wild fish, or the degree that hybrids return to a natal spawning area. The source-sink model is a powerful means of detecting low levels of introgression over several generations.     

An overview of salmon enhancement and the need to manage and monitor natural spawning in Hokkaido, Japan

The chum and pink salmon catches in Hokkaido, Japan have increased dramatically since the 1970s and the 1990s, respectively. In contrast, masu salmon catches have been steadily decreasing. Despite intensive hatchery development in Hokkaido, naturally spawning salmon populations persist based on results from a recent river survey. This paper focuses on the challenges of maintaining hatchery salmon populations while protecting natural chum, pink and masu salmon populations in Hokkaido. Two important initiatives related to meeting this ambitious goal are managing hatcheries in a way that minimizes negative interactions between natural and hatchery salmon populations, and initiating new efforts at restoring and rehabilitating degraded freshwater habitats. In addition, in order to maintain a balance of demand and supply in the domestic market through the exportation of extra salmon, Hokkaido has decided to enter full assessment to gain Marine Stewardship Council (MSC) certification of the Hokkaido chum salmon trap net fishery. This would involve a fundamental shift in fisheries management as practiced in Japan, specifically elevating the importance of managing the fishery in a way that conserves natural salmon populations. A key component of a new salmon management strategy is the establishment of a zone management framework based on the designation of stream units to spatially separate natural salmon from hatchery salmon to minimize negative effects of hatchery fish and to utilize effectively hatchery salmon for commercial fisheries. This effort is allied with similar initiatives in other Pacific Rim countries that are focusing on management reform to restore natural ecosystem function and maintain the coexistence of wild and hatchery salmon.