Interactions of Atlantic Salmon in the Pacific Northwest IV. Impacts on The
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Fisheries Research 62 (2003) 307–328 Interactions of Atlantic salmon in the Pacific Northwest IV. Impacts on the local ecosystems F. William Waknitz∗, Robert N. Iwamoto, Mark S. Strom Northwest Fisheries Science Center, National Marine Fisheries Service, 2725 Montlake Boulevard East, Seattle, WA 98112, USA Abstract The paper begins with the introduction of Atlantic salmon into the local ecosystems of the Pacific Northwest (Puget Sound), and describes potential interactions with native salmonids. Specific sections review possible hybridization between Atlantic and Pacific salmon, genetic dilution and alteration of the gene pool, the colonization of the aquatic environment by Atlantic salmon, and finally the interactions of wild salmon and genetically altered transgenics. This is followed by possible epidemics and transmission of waterborne disease, and reviews the potential for cultured Atlantic salmon, a non-native species, to introduce new diseases into the local ecosystem. There are nine specific items, from the diseases which might be involved, to potential interactions, and current policies for disease control. After a review of the potential ecological impacts in the Pacific Northwest, specifically the interaction with Pacific salmon and predation, there are three parts on the effects of artificial propagation practices in the region in general, the impacts of the introduction of various non-indigenous salmonid species, and a comparison of escapes or releases of propagated Atlantic and Pacific salmon. The last part examines the potential effects of Atlantic salmon vis-à-vis the Biological Status Reviews of west coast Pacific salmon stocks carried out by the National Marine Fisheries Service. The review ends by summarizing the varying degrees of risk carried by these issues. Published by Elsevier Science B.V. Keywords: Atlantic salmon; Salmonids; Pacific Northwest ecosystems; Colonization; Predation; Genetic interactions; Waterborne diseases; Risks 1. The artificial propagation of salmonids in the the extremely large complex of federal, state, tribal, Pacific Northwest and cooperative hatcheries in Alaska and the western states, concerns about the potential adverse impacts of 1.1. Background private trout and salmon culture in Washington have also been described by Ellis (1996), and Alverson and Artificial propagation of salmon and trout in the Ruggerone (1997). Pacific Northwest has come under increasing scrutiny Concerns about genetic interactions, the transmis- in recent years. This is due to the recognition that sion of disease, and ecological interactions are most hatchery-cultured salmon and trout may have the po- commonly voiced. In 1997, e.g., the Pollution Control tential to impact natural populations adversely. Al- Hearing Board of the State of Washington (PCHB) though the weight of attention has been focused on heard testimony that Atlantic salmon had the poten- tial to hybridize with Pacific salmon, based on un- Corresponding author. Tel.: 1-360-871-8322; published Canadian laboratory studies, and that it was ∗ + fax: 1-206-842-8364. possible the 369,000 Atlantic salmon which escaped + E-mail address: [email protected] (F.W. Waknitz). into Puget Sound in 1996 would produce 10 million 0165-7836/03/$ – see front matter. Published by Elsevier Science B.V. doi:10.1016/S0165-7836(03)00066-3 308 F.W. Waknitz et al. / Fisheries Research 62 (2003) 307–328 healthy smolts in local rivers. Subsequently, in 1999, salmon in particular, is the potential genetic effects the Alaska Department of Fish and Game (ADF&G) of deliberate releases and inadvertent escapees on the published a press release that escaped Atlantic salmon native salmonids. For the salmon farming industry from west coast salmon farms would compete with in British Columbia, Canada, where both Pacific and wild salmon, and spread diseases and parasites for Atlantic salmon are extensively farmed, the British which Pacific salmon had little resistance. Columbia Salmon Aquaculture Review published by the Environment Assessment Office (EAO, 1997) 1.2. The origin and disease status of the Atlantic listed four major areas of concern: salmon stocks in Puget Sound Hybridization between Pacific and Atlantic salmon; • Genetic dilution and alteration of the wild salmonid In 1971 scientists from the National Marine Fish- • gene pool; eries Service (NMFS) at the Manchester Research Colonization by Atlantic salmon; Station began testing the feasibility of rearing New • Interactions between wild salmon and genetically England stocks of Atlantic salmon in seawater • altered transgenics. net-pens in Puget Sound to provide 3.5 million eyed eggs annually for restoring depleted runs in southern These concerns are both geographically and species New England. Between 1971 and 1983, the station specific. Across the border in Puget Sound, the con- received eggs from many North American stocks, cern is only with farmed Atlantic salmon, as Pacific including the Grand Cascapedia River in Quebec (via salmon, with rare exception, are not cultured by pri- Oregon State), and the Penobscot, Union, St. John, vate enterprises. and Connecticut Rivers in the United States. Each delivery of eggs was examined in accordance 2.1. Hybridization with federal regulations and certified by federal pathol- ogists to be free of bacterial and viral pathogens prior No genetic interactions between Atlantic and wild to transfer from New England to Washington. How- Pacific salmon have been reported in the Pacific ever, few eggs were ever sent back as a panel of New Northwest. Similarly, under controlled and protected England state and federal fisheries officials determined laboratory conditions where survival of hybrid off- in 1984 that raising Atlantic salmon in Puget Sound spring should be optimized, viable hybrids between rendered the eggs unfit for transfer back to the east Atlantic and Pacific salmonid species are difficult to coast because of the risk of introducing Pacific salmon produce. Refstie and Gjedrem (1975), Sutterlin et al. diseases to New England Atlantic salmon populations. (1977), and Blanc and Chevassus (1982) found that As a result of this decision, millions of Atlantic crosses between Atlantic salmon and rainbow trout salmon eggs originally meant for New England failed to produce any viable progeny. A similar lack restoration programs were available for distribution of vitality was observed in pairings of Atlantic salmon to salmon farmers in Washington. These eggs proved and coho salmon (Chevassus, 1979), and Atlantic to be beneficial to the local industry as, by this time, salmon and pink salmon (Loginova and Krasnoperova, it was clear from work in Norway and Scotland, that 1982). Gray et al. (1993) attempted to produce diploid Atlantic salmon were superior to Pacific salmon in and triploid hybrids by crossing Atlantic salmon with all aspects of culture, including survival to hatching, chum and coho salmon, and rainbow trout. All em- growth rate in fresh and sea water, and resistance to bryos died in early developmental stages, leading to infectious diseases. the conclusion that hybridization of Atlantic salmon with Pacific salmon species was unlikely to happen. Recently, two pilot studies from British Columbia 2. Genetic interactions of artificially propagated have provided more data regarding the lack of genetic Pacific and Atlantic salmon compatibility between Atlantic and Pacific salmon (R. Devlin, Department of Fisheries and Oceans (DFO), Amajorconcernwithartificialpropagationingen- Canada, reported in Alverson and Ruggerone, 1997). eral, and farming of Pacific salmonids and Atlantic In the first study, using a small number of eggs, crosses F.W. Waknitz et al. / Fisheries Research 62 (2003) 307–328 309 with Atlantic salmon produced a few hybrids with pink of producing hybrids, might produce genetic distur- salmon, but no hybrids with coho, chum, chinook, bances by interfering with wild salmonid breeding be- sockeye salmon, and rainbow trout. In the same exper- havior. For example, by beating Pacific salmon males iment, by contrast, the interspecific crosses between to a redd, the resulting non-viable eggs would reduce Oncorhynchus species produced hybrids to hatch the number of juvenile salmonids available for recruit- ranging from 10 to 90% in 15 of the 42 crosses, with ment in depressed populations. each species of Pacific salmon readily produced hy- Although this scenario could possibly occur in brids with between two and five other Pacific salmon some locations, it is improbable in the tributaries of species, confirming previous observations in this Puget Sound for a number of reasons. First, salmon genus (Foerster, 1935; Seeb et al., 1988). Moreover, farmers in Puget Sound use Atlantic salmon de- because of dissimilar natural spawning times between rived from stocks provided to them by NMFS in Atlantic salmon in the fall, and steelhead in the spring, the mid-1980s, primarily Penobscot River and Grand this particular cross was performed using cryopre- Cascapedia River strains. The Penobscot River hatch- served Atlantic salmon sperm. In the second study, ery strain is know to have a remarkably low incidence using a larger number of eggs and involving crosses of early maturity, either after 1 or 2 years in freshwa- between Atlantic salmon and rainbow and steel- ter (precocious male parr), or at 2 or 3 years of age (1 head trout, coho, chum, chinook, and pink salmon, a year at sea), known as grilse (Ritter et al., 1986). As few hybrids were produced. Cryopreserved Atlantic age at maturity is a genetically inherited trait,