The School Aquatic & Fishery Sciences' Alaska Salmon Program is unique in that it represents the world's longest-running effort to monitor salmon and their ecosystems. Several accounts of various aspects of the history of this program and the industry and resources it has supported have been written over the years, including the account below by Tom Quinn. In addition, we anticipate further contributions in the future, including a biography by Robert L. "Bud" Burgner who, as director of the school's Fisheries Research Institute, played a major role in the development and expansion of the Alaska Salmon Program. Links to Burgner's memoirs and two books detailing the history of fisheries at UW and studies of Alaska sockeye salmon are provided below:

Studies of Alaska Red Salmon (ed. Koo 1962)

Flagship: A History of Fisheries at the University of Washington (Stickney 1989)

The Memoirs of Robert L. (Bud) Burgner (Burgner 2009)

Introduction

Commercial catches of Pacific salmon (Oncorhynchus spp.) in Bristol Bay and the Alaska Peninsula began in the late 1800s and, by the early 1900s, these were the largest salmon fisheries in the world. Management of the fisheries was based on the simple premise that salmon should not be overfished. Initially, restrictions on fishing gear and location were used to limit fishing efficiency; then in 1924 Congress passed the White Act, which required that 50% of the returning salmon be allowed to spawn. There was no biological basis for this figure and it bore no relationship to principles of population dynamics. Beginning in 1919, sockeye (O. nerka) salmon catches declined from approximately 22 to 16 million fish per year, followed by a second significant decline to approximately 10 million per year in the mid-1940s. With the hope of bringing the salmon runs back to their original level, the fishing industry commissioned Dr. W.F. Thompson, Director of the School of Fisheries at the University of Washington, to investigate factors influencing the dramatic decline in salmon abundance in southwestern Alaska.

The University of Washington's salmon research program in Alaska began in 1946, and during 1947 a permanent field station was constructed at Aleknagik in the Wood River system, a series of five interconnected lakes stretching over 200 kilometers (km) and flowing into Bristol Bay. A year later, the Fisheries Research Institute (FRI) was established on the University of Washington campus as a base for salmon research in Alaska. In 1955 and 1960, additional research stations were built at Chignik and Iliamna lakes, and the suite of facilities now includes six camps: Iliamna village and Porcupine Island (both on Iliamna Lake), Aleknagik and Lake Nerka (Wood River system), and Chignik. There is also a cabin at the outlet of Lake Kulik in the upper Wood River system.

The research objectives of this program were to determine physical and biological factors influencing sockeye salmon production. These objectives were developed at a time when the fundamental biology of salmon was poorly known and there were no long-term studies integrating salmon and their ecosystems in a holistic manner. Many basic techniques for counting salmon and understanding their life-history patterns were developed at these camps, and the data were used for the management of the salmon runs. The early period of research preceded Alaska’s statehood (in 1959), but the fisheries are now managed by the Alaska Department of Fish and Game (ADF&G). Some of the sampling programs that were initiated by FRI (e.g., counting the salmon smolts leaving Iliamna Lake) have been taken over as routine operations by ADF&G. However, we have continued a great deal of biological sampling that provides basic information on life-history patterns and population dynamics, and that is directly used in the conservation of the populations (e.g., abundance, size at age, and age structure of adult salmon). We also have many sampling programs with no direct connection to fisheries management but that document the within-season and interannual patterns of the physical environment (e.g., temperature, solar radiation and lake level) and biotic communities (e.g., emerging insects, chlorophyll, zooplankton, non-salmonid fishes, juvenile salmon growth).

Most research at the FRI field stations since 1946 has addressed applied aspects of salmon and aquatic ecology that were directly relevant to the salmon processing industry, which has supported the research program. We continue to maintain this close relationship with industry and will also take advantage of the long-term datasets on climate change and biotic responses (limnology, fish growth, community structure, etc.) to pursue new areas of research. The field stations represent a unique and invaluable opportunity for aquatic and terrestrial research because of their near-pristine environment and the absence of introduced populations or exotic species. The sites are located on the edge of Beringia (the great Pleistocene refuge from glaciation) and contain animals and plants found nowhere else in North America.

For a list of the physical and biological attributes under long-term study, click here.

Research Foci

Wood River Lakes

Initial research at the Wood River Lakes and throughout Bristol Bay was aimed at determining the carrying capacity of spawning and rearing environments (Burgner et al. 1969). To achieve this goal, basic techniques were devised for estimating spawning densities of salmon, determining the age of fish by scale or otolith patterns, conducting stream surveys, estimating the abundance of emigrating juvenile salmon (smolts), and quantitative sampling of salmon catches. Additional studies described the life-history characteristics of sockeye salmon and associated species (predators, competitors, prey, and parasites). Much of the earlier work by individual investigators was published in two books (Koo 1962, Burgner 1968) devoted entirely to research in the Wood River lakes, Iliamna Lake, and the Chignik lakes. More recently, Burgner (1991) summarized the life history and ecology of sockeye salmon, including extensive data from southwestern Alaska.

Research at the Wood River Lakes during the 1970s and 1980s was focused on factors influencing the survival of juvenile and adult sockeye salmon. Studies on freshwater ecology included seasonal and interannual variations in juvenile fish abundance and distribution (Rogers 1973, Newcome 1976); effects of lake enrichment on the phytoplankton, zooplankton, and limnetic fishes in small, oligotrophic Little Togiak Lake (Rogers 1979, Hardy 1979); density-dependent growth of juvenile (Burgner 1987) and maturing sockeye salmon (Rogers 1980); functional response and size-biased predation by Arctic char, Salvelinus alpinus, on juvenile sockeye salmon (Ruggerone and Rogers 1984); and parasites of juvenile sockeye salmon (Burke 1978). But research was not restricted to sockeye salmon: studies were also conducted on the life history of Arctic char (Moriarity 1977, McBride 1979) and on natural selection and phenotypic variation in the threespine stickleback, Gasterosteus aculeatus (Hagen and Gilbertson 1972, 1973a,b; Gilbertson 1980).

During the last decade we have continued many of the same themes initiated by earlier researchers but have taken advantage of the long periods of record now available. We have investigated interrelationships between abundance and growth of planktivorous fishes (especially sticklebacks and juvenile sockeye salmon) and the seasonal and interannual trends in phytoplankton and zooplankton (e.g., Work 1992, Reischauer 1996). Our measurements of salmon growth in freshwater and the sea, as determined from scales, extend back to 1907 and can be related to salmon abundance and long-term trends in climate and the ocean environment (Zimmerman 1991, Rogers and Ruggerone 1993). We have also had several students interested in the effects of adult salmon on the stream ecosystem. For example, Eastman (1996) documented the shift in diet from insects to sockeye salmon eggs in rainbow trout and Arctic char, and Peterson (1998) showed that digging by female salmon disturbs a great number of insects that are consumed by rainbow trout (Oncorhynchus mykiss), Arctic char, and grayling (Thymallus arcticus) before eggs become available as food.

In addition to the work on limnology and fish growth, research at the Wood River system during the 1990s was focused on the evolutionary factors driving population-specific adaptations among different spawning habitats. We have documented differences in egg size that correspond to the sizes of the spawning substrates (Wetzel, Quinn et al. 1995). There are also differences in size, age structure, and morphology (especially the sexually dimorphic dorsal hump of male sockeye salmon) that correspond to stream size (Bishop 1990, Wetzel 1993) and vulnerability to bear predation (Hanson 1992, Quinn in prep.). In addition to these forms of natural selection, we have also examined the selective effects of the gillnet fishery on size and shape of adult sockeye salmon (Hamon 1995). These studies of selection are being combined with detailed work at selected streams (notably Pick and Hansen creeks) on the relative influences of size and spawning date on behavior, stream life, and reproductive success of adult salmon (McPhee and Quinn 1998, Quinn and McPhee 1998, Hendry et al. in press, Steen and Quinn in press).

Iliamna Lake

Much of the early research at Iliamna Lake involved investigating the limnology of the system (e.g., Mathisen 1966). Lenarz (1966), Gunnerod (1971), and Carlson (1974) examined the life histories and population dynamics of key zooplankters (Cyclops scutifer and Bosmina coregoni). Other studies involved fish/zooplankton interactions (Hoag 1972), limnological effects of volcanic ash (Mathisen and Poe 1978), and density-dependent growth of juvenile sockeye salmon (Mathisen 1969). Margolis (1967) and Pennell et al. (1973) described the parasite fauna of sockeye salmon, and more recently, researchers examinedthe possible relationship between parasites and sexual dimorphism of adult male salmon (Berg et al. 1995) and fluctuating asymmetry (Berg et al. 1997).

The Iliamna Lake system was noteworthy for the cyclic abundance of adult sockeye salmon (Mathisen and Poe 1981), and the exceptional runs in some years (up to 20 million salmon escaped the fishery) stimulated early investigations of the role of salmon carcasses in nutrient cycling. Donaldson (1967) developed a phosphorus budget for the lake and evaluated the effect on salmon growth and survival of nutrients from the millions of salmon carcasses deposited annually. Subsequent workers used stable isotope ratios of nitrogen to demonstrate that the marine-derived contributions from salmon carcasses represent the major source of limiting nutrients to these oligotrophic lakes and streams (Mathisen et al. 1988, Kline et al. 1993).

The biological basis for the cyclic behavior of Iliamna Lake populations has not been determined, but modeling by Eggers and Rogers (1987) indicated that it might have been generated by random fluctuations in abundance and depensatory fishing mortality. This research resulted in a dramatic change in the management of the system’s populations. Fishing pressure was reduced in "off-peak" years to increase sockeye salmon abundance. Over the next few years, we will be able to better understand the success or failure of this approach to conserving the overall abundance and diversity of salmon populations. The lake system has about 100 identified spawning populations, which differ dramatically in abundance. Their conservation is complicated by the fact that they all pass through the fishery and enter the lake at about the same time (Jensen and Mathisen 1987) and have unequal tendencies to cycle.

The Iliamna Lake populations occupy diverse physical habitats and show a corresponding diversity of life-history patterns. Early investigations on fecundity by Mathisen and Gunnerod (1969) led to more detailed studies on the variation in egg size and fecundity among populations (Blair et al. 1993, Quinn et al. 1995). Studies on male salmon indicated that, like the Wood River populations, those in the Iliamna Lake system also display great variation in secondary sexual traits. In particular, males that spawn in beach environments show extreme development of the dorsal hump. Kerns and Donaldson (1968) described the behavior of these beach-spawning populations, and subsequent research showed that sexual selection favors not only large males but those with deep bodies for their length (Quinn and Foote 1994). These breeding populations spawn at high densities in very clear water near shore and are ideal for testing hypotheses in behavioral ecology related to homing and spawning-site selection (Blair and Quinn 1991, Hendry et al. 1995), effects of skewed operational sex ratio on male behavior (Quinn et al. 1996), sexual dimorphism (Quinn and Blair 1992), and other features of reproductive biology. The beaches are an unusual physical environment for spawning, characterized by very large substrate and wind-driven water circulation (Leonetti 1995). The large substrate makes the salmon eggs vulnerable to predation by sculpins (Cottus spp.), initially studied by Roger (1971) and more recently by Foote and Brown (1998) and Dittman et al. (1998).

Chignik Lakes

Early FRI work in the Chignik Lakes formed the basis for current fisheries management in this region. Investigators examined the carrying capacity of the lakes for juvenile sockeye (Narver 1966), competition between sockeye and other planktivores (Narver 1966, Parr 1972), food habits of fishes and predation on juvenile sockeye (Roos 1959, 1960; Narver and Dahlberg 1965), sockeye spawner/return relationships for the purpose of maximizing harvests (Dahlberg 1968, 1973), scale patterns of juvenile sockeye as a means of identifying lake of origin (Narver 1963, Marshall 1977, Conrad 1983, Marshall et al. 1987) and run timing and forecasting of adult sockeye (Parker 1986). Other studies have examined the distribution, seasonal movements and mortality of brown bears (Glenn and Miller 1980, Miller and Sellers 1989), phenotypic variation in threespine stickleback (Narver 1969), distribution of birds (Narver 1970) and the life history of an isopod (Narver 1968). The FRI has maintained a long-term data set (1922 to present) of adult sockeye production (returns per spawner) and fish scales. These scales provide a historical record of fish age and growth in freshwater and the ocean (Bumgarner 1993). Data on phytoplankton abundance, zooplankton species composition and abundance, fish species composition, size, and abundance have been collected periodically since 1955 and are maintained in FRI's archives.

One recent focal area of research in the Chignik lakes has been the interactions between piscivorous coho salmon and their primary prey, sockeye salmon. Ruggerone (1989a, b) quantified the effect of water temperature and meal size on the gastric evacuation rate of coho that consumed sockeye fry, then developed and tested a model for predicting evacuation rates of coho in the lake. By comparing gastric evacuation and bioenergetic approaches for estimating fish food consumption and two independent estimates of coho abundance, Ruggerone (1989c) and Ruggerone and Rogers (1992) estimated that about 55 million juvenile sockeye, or 59% of the sockeye population, were consumed each year during 1985—87. Inverse trends in adult sockeye and coho run size since 1971 also suggested the strong effect of predation on sockeye and the need for multispecies management of the commercial fishery. Ruggerone (1989c) provided evidence for evolutionary effects of size-biased predation on sockeye size at emergence and size-related shifts in habitat use by juvenile sockeye. Juvenile sockeye experienced less risk of predation when they aggregated with threespine sticklebacks than when they form single-species aggregations (Ruggerone 1992).

Marine Research

In addition to various research conducted in lakes and streams at our field stations, the anadromous life cycle of Pacific salmon gives our research program a connection to marine processes. Donald Rogers was involved in several studies addressing the effects of the marine phase on salmon life history:

  1. density-dependent influences of sockeye salmon growth at sea (Rogers 1980, Rogers and Ruggerone 1993);
  2. temporal trends in the production of salmon and the marine environment (Rogers 1984);
  3. genetic and environmental effects on the age of sockeye maturity (Rogers 1987), and;
  4. the effects of salmon smolt size and migration timing on the distribution and survival at sea (Rogers 1988).

Since the late 1980s, we have been operating a test fishery in the outer region of Bristol Bay (from Port Moller) to obtain samples and to estimate the abundance and timing of the run. We combine these and other samples collected by FRI with data collected by ADF&G and other agencies to examine the changes in size at age and age structure of the populations in response to changing ocean temperature regimes, abundance of prey organisms, and salmon density.

Bibliography

A list of thesis and dissertation citations resulting from research conducted under the Alaska Salmon Program (1987–present) is available. A more comprehensive list of publications pertaining to or derived from Alaska Salmon Program research also is available.

References

Berg, O.K., M.D. Adkison, and T.P. Quinn. 1997. Bilateral asymmetry, sexual dimorphism, and nematode parasites in mature male sockeye salmon Oncorhynchus nerka. Northw. Sci. 71:305-312.

Berg, O.K., C.J. Foote, and T.P. Quinn. 1995. Fish age, nematode, Philonema oncorhynchi, infection and development of sexual dimorphism by the adult male sockeye salmon, Oncorhynchus nerka, in western Alaska. Can. J. Zool. 73:1999-2004.

Bishop, S. 1990. Morphometric characteristics of Wood River Lakes sockeye spawning populations. M.S. thesis, Univ. Washington, Seattle.

Blair, G.R. and T.P. Quinn. 1991. Homing and spawning site selection by sockeye salmon (Oncorhynchus nerka) in Iliamna Lake, Alaska. Can. J. Zool. 69:176-181.

Blair, G.R., D.E. Rogers, and T.P. Quinn. 1993. Variation in life history characteristics and morphology of sockeye salmon (Oncorhynchus nerka) in the Kvichak River system, Bristol Bay, Alaska. Trans. Am. Fish. Soc. 122:550-559.

Bumgarner, J.D. 1993. Long-term variation in growth and abundance of Chignik sockeye salmon. M.S. thesis, Univ. Washington. Seattle.

Burgner, R.L. (ed.). 1968. Further Studies of Alaska Sockeye Salmon. Univ. Washington Publ. Fisheries, New Ser. 3:199-267

Burgner, R.L. 1987. Factors influencing age and growth of juvenile sockeye salmon (Oncorhynchus nerka) in lakes. Can. Spec. Publ. Fish. Aquat. Sci. 96:129-142.

Burgner, R.L. 1991. The life history of sockeye salmon (Oncorhynchus nerka). Pages 3-117 in C. Groot and L. Margolis (eds), Pacific Salmon Life Histories. Univ. British Columbia Press.

Burgner, R.L., C.J. DiCostanzo, R.J. Ellis, G.Y. Harry, Jr., W.L. Hartman, O.E. Kerns, Jr., O.A. Mathisen, and W.F. Royce. 1969. Biological studies and estimates of optimum escapements of sockeye salmon in the major river systems in southwestern Alaska. Fish. Bull. U.S. 67:405-459.

Burke, J.M. 1978. Occurrence and effects of the parasite Triaenophorus crassus in sockeye salmon, Oncorhynchus nerka, of the Wood River lake system, Bristol Bay, Alaska. M.S. thesis, Univ. Washington. Seattle.

Carlson, T.J. 1974. The life history and population dynamics of Bosmina coregoni in Pedro Bay, Iliamna Lake, Alaska. M.S. thesis, Univ. Washington. Seattle.

Conrad, R.H. 1983. Management applications of scale pattern analysis methods for the sockeye salmon runs to Chignik, Alaska. M.S. thesis, Univ. Washington. Seattle.

Dahlberg, M. L. 1968. Analysis of the dynamics of sockeye salmon returns to the Chignik Lakes, Alaska. Ph.D. dissertation, Univ. Washington. Seattle.

Dahlberg, M.L. 1973. Stock-and-recruitment relationships and optimum escapements of sockeye salmon stocks of the Chignik Lakes, Alaska. Cons. int. Explor. Mer, Rapp. et Proc.-Verb. 164:98-105.

Dittman, A.H., G.S. Brown, and C.J. Foote. 1998. The role of chemoreception in salmon egg predation by coastrange (Cottus aleuticus) and slimy (C. cognatus) sculpins in Iliamna Lake, Alaska. Can. J. Zool. 76:406-413.

Donaldson, J.R. 1967. The phosphorus budget of Iliamna Lake, Alaska, as related to the cyclic abundance of sockeye salmon. Ph.D. dissertation, Univ. Washington. Seattle.

Eastman, D. 1996. Response of freshwater fish communities to spawning sockeye salmon. M.S. thesis, Univ. Washington. Seattle.

Eggers, D.M. and D.E. Rogers. 1987. The cycle of runs of sockeye salmon (Oncorhynchus nerka Walbaum) to the Kvichak River, Bristol Bay, Alaska: Cyclic dominance or depensatory fishing? Can. Spec. Publ. Fish. Aquat. Sci. No. 96:343-366.

Foote, C.J. and G.S. Brown. 1998. Ecological relationships between freshwater sculpins (genus Cottus) and beach-spawning sockeye salmon (Oncorhynchus nerka) in Iliamna Lake, Alaska. Can. J. Fish. Aquat. Sci. 55:1524-1533.

Glenn, L.P. and L.H. Miller. 1980. Seasonal movements of an Alaska Peninsula brown bear population. Pages 307-312 in Bears--Their Biology and Management. Intl. Assoc. Bear Biol. Conf. Ser. 4.

Gilbertson, L.G. 1980. Variation and natural selection in an Alaskan population of the threespine stickleback (Gasterosteus aculeatus L.). Ph.D. dissertation, Univ. Washington. Seattle.

Gunnerod, T. 1971. The life history and population dynamics of Cyclops scutifer Sars in Pedro Bay, Iliamna Lake, Alaska. Ph.D. dissertation, Univ. Washington. Seattle.

Hagen, D.W. and L.G. Gilbertson. 1972. Geographic variation and environmental selection in Gasterosteus aculeatus L. in the Pacific Northwest, America. Evolution 26:32-51.

Hagen, D.W. and L.G. Gilbertson. 1973a. Selective predation and the intensity of selection acting upon the lateral plates of threespine sticklebacks. Heredity 30:273-287.

Hagen, D.W. and L.G. Gilbertson. 1973b. The genetics of plate morphs in freshwater threespine sticklebacks. Heredity 31:75-84.

Hamon, T. 1995. Artificial selection in the wild: selection on spawner morphology of wild sockeye salmon (Oncorhynchus nerka) by a gillnet fishery. M.S. thesis, Univ. Washington.

Hanson, R. 1992. Brown bear (Ursus arctos) predation on sockeye salmon (Oncorhynchus nerka) spawners in two tributaries of the Wood River Lake system, Bristol Bay, Alaska. M.S. thesis, Univ. Washington. Seattle.

Hardy, F.J. 1979. Effects of inorganic fertilization on phytoplankton in Little Togiak Lake, Alaska. M.S. thesis, Univ. Washington. Seattle.

Hendry, A.P., F.E. Leonetti, and T.P. Quinn. 1995. Spatial and temporal isolating mechanisms: the formation of discrete breeding aggregations of sockeye salmon (Oncorhynchus nerka). Can. J. Zool. 73:339-352.

Hendry, A.P., O.K. Berg, and T.P. Quinn. In press. Breeding date, life history, and energy allocation in a population of sockeye salmon (Oncorhynchus nerka). Oikos.

Hoag, S.H. 1972. The relationship between the summer food of juvenile sockeye salmon, Oncorhynchus nerka, and the standing stock of zooplankton in Iliamna Lake, Alaska. Fish. Bull. 70:355-362.

Jensen, K.A. and O.A. Mathisen. 1987. Migratory structure of the Kvichak sockeye salmon (Oncorhynchus nerka) escapement, 1983. Can. Spec. Publ. Fish. Aquat. Sci. 96:101-109.

Kerns, O.E. and J.R. Donaldson. 1967. Behavior and distribution of spawning sockeye salmon on island beaches in Iliamna Lake, Alaska, 1965. J. Fish. Res. Board Can. 25:485-494.

Kline, T.C., J.J. Goering, O.A. Mathisen, P. Poe, P.L. Parker, and R.S. Scalan. 1993. Recycling of elements transported upstream by runs of Pacific salmon: II. d 13C and d 15N evidence in the Kvichak River watershed, Bristol Bay, Southern Alaska. Can. J. Fish. Aquat. Sci. 50:2350-2365.

Koo, T.S.Y. 1962. Studies of Alaska Red Salmon. Univ. Washington Publications in Fisheries, New Ser., Vol. 1. Univ. Washington Press, Seattle.

Leonetti, F.E. 1997. Estimation of surface and intragravel water flow at sockeye salmon spawning beaches in Iliamna Lake, Alaska. N. Amer. J. Fish. Manag. 17:194-201.

Lenarz, W.H. 1966. Population dynamics of Cyclops scutifer and an evaluation of a sampling program for estimating the standing crop of zooplankton in Iliamna Lake. M.S. thesis, Univ. Washington. Seattle.

Margolis, L. 1967. Triaenophorus crassus plerocercoids in sockeye salmon smolts from the Kvichak River system, Alaska. J. Fish. Res. Board Can. 24:893-894.

Marshall, S.L. 1977. Lacustrine scale patterns of sockeye salmon of the Chignik Lakes, Alaska. M.S. thesis, Univ. Washington. Seattle.

Marshall, S.L. and S.S. Parker. 1982. Pattern identification in the microstructure of sockeye salmon (Oncorhynchus nerka) otoliths. Can. J. Fish. Aquat. Sci. 39:542-547.

Marshall, S.L., D. Bernard, R. Conrad, B. Cross, O. McBride, A. McGregor, S. McPherson, G. Oliver, S. Sharr, and B. Van Alen. 1987. Application of scale patterns analysis to management of Alaska's sockeye salmon (Oncorhynchus nerka) fisheries. Can. Spec. Publ. Fish. Aquat. Sci. 96:307-326.

Mathisen, O.A. 1966. Some adaptations of sockeye salmon races to limnological features of Iliamna Lake, Alaska. Verh. Internat. Verein. Limnol. 16:1025-1035.

Mathisen, O.A. 1969. Growth of sockeye salmon in relation to abundance in the Kvichak District, Bristol Bay, Alaska. FiskDir. Skr. Ser. HavUnders. 15:172-185.

Mathisen, O.A. and T. Gunnerod. 1969. Variance components in the estimation of potential egg deposition of sockeye salmon escapements. J. Fish. Res. Board Can. 26:655-670.

Mathisen, O.A. and P.H. Poe. 1978. Effect of volcanic ash deposits on sockeye salmon lakes. Verh. Intl. Ver. Limnol. 20:165-172.

Mathisen, O.A. and P.H. Poe. 1981. Sockeye salmon cycles in the Kvichak River, Bristol Bay, Alaska. Verh. Internat. Verein. Limnol. 21:1206-1213.

Mathisen, O.A., P.L. Parker, J.J. Goering, T.C. Kline, P.H. Poe, and R.S. Scalan. 1988. Recycling of marine elements transported into freshwater by anadromous salmon. Verh. Int. Ver. Limnol. 23:2249-2258.

McBride, D.N. 1979. Homing of arctic char, Salvelinus alpinus (Linnaeus), to feeding and spawning sites in the Wood River lake system, Alaska. M.S. thesis, Univ. Alaska.

McPhee, M.V. and T.P. Quinn. 1998. Factors affecting the duration of nest defense and reproductive lifespan of female sockeye salmon, Oncorhynchus nerka. Envir. Biol. Fish. 51:369-375.

Miller, S.D. and R.A. Sellers. 1989. Density and structure of a hunted population of brown bears at Black Lake, Alaska. Progr. Rept. Alaska Dept. Fish Game.

Moriarity, D.S. 1977. Arctic char in the Wood River lakes. M.S. thesis, Univ. Washington. Seattle.

Narver, D.W. 1963. Identification of adult red salmon groups by lacustrine scale measurement, time of entry, and spawning characteristics. M.S. thesis, Univ. Washington. Seattle.

Narver, D.W. 1966. Pelagial ecology and carrying capacity of sockeye salmon in the Chignik Lakes, Alaska. Ph.D. dissertation, Univ. Washington. Seattle.

Narver, D.W. 1968. Notes on the isopod Mesidota entomon in Chignik lakes, Alaska. J. Fish. Res. Board Canada 25:157-167.

Narver, D.\W. 1969. Phenotypic variation in threespine stickleback (Gasterosteus aculeatus) of the Chignik River system, Alaska. J. Fish. Res. Board Canada 26:405-412.

Narver, D. W. 1970. Birds of the Chignik River drainage, Alaska. The Condor 72:102-105.

Narver, D.W. and M. L. Dahlberg. 1965. Estuarine food of Dolly Varden at Chignik, Alaska. Trans. Amer. Fish. Soc. 94:405-408.

Newcome, N. 1976. Relative distribution, abundance, size, and food habits of juvenile sockeye salmon and associated species in the Igushik Lakes, Alaska. M.S. thesis, Univ. Washington. Seattle.

Parker, S.S. 1986. Forecasting the abundance and timing of sockeye salmon migrations to Chignik, Alaska. M.S. thesis, Univ. Washington. Seattle.

Parr, W.H. 1972. Interactions between sockeye salmon and lake resident fish in the Chignik Lakes, Alaska. M.S. thesis, Univ. Washington. Seattle.

Pennell, D.A., C.D. Becker, and N.R. Scofield. 1973. Helminths of sockeye salmon (Oncorhynchus nerka) from the Kvichak River system, Bristol Bay, Alaska. Fish. Bull. 71:267-277.

Peterson, D. 1996.The effects of spawning sockeye salmon, Oncorhynchus nerka, on small streams in the Wood River Lakes, Alaska: evidence from the benthos and invertebrate drift. M.S. thesis, Univ. Washington. Seattle.

Quinn, T.P. and G.R. Blair. 1992. Morphological changes in senescing adult male sockeye salmon (Oncorhynchus nerka Walbaum). J. Fish Biol. 41:1045-1047.

Quinn, T.P. and C.J. Foote. 1994. The effects of body size and sexual dimorphism on the reproductive behaviour of sockeye salmon (Oncorhynchus nerka). Anim. Behav. 48:751-761.

Quinn, T.P., A.P. Hendry, and L.A. Wetzel. 1995. The influence of life history trade-offs and the size of incubation gravels on egg size variation in sockeye salmon (Oncorhynchus nerka). Oikos 74:425-438.

Quinn, T.P., M.D. Adkison, and M.B. Ward. 1996. Behavioral tactics of male sockeye salmon (Oncorhynchus nerka) under varying operational sex ratios. Ethology 102:304-322..

Quinn, T.P. and M.V. McPhee. 1998. Effects of senescence and density on the aggression of adult female sockeye salmon. J. Fish Biol. 52:1295-1300.

Quinn, T.P., E.C. Volk, and A.P. Hendry. In press. Natural otolith microstructure patterns reveal precise homing to natal incubation sites by sockeye salmon (Oncorhynchus nerka). Can. J. Zool.

Reeves, J. 1969. A comparative analysis of Arctic char populations in sockeye salmon lakes of the Nushagak district, Alaska. Ph.D. dissertation, Univ. Washington. Seattle.

Reischauer, A. 1996. A comparison of the zooplankton in sockeye and barren lakes in the Nushagak district of Bristol Bay, Alaska. M.S. thesis, Univ. Washington. Seattle.

Roger, P.B. 1971. The ecology of two species of cottids in Iliamna Lake, Alaska, and their relation to sockeye salmon. M.S. thesis, Univ. Washington. Seattle.

Rogers, B.J. 1979. Responses of juvenile sockeye salmon and their food supply to inorganic fertilization of Little Togiak Lake, Alaska. M.S. thesis, Univ. Washington. Seattle.

Rogers, D.E. 1973. Abundance and size of juvenile sockeye salmon, and associated species in Lake Aleknagik, Alaska, in relation to their environment. Fish. Bull. 71:1061-1075.

Rogers, D.E. 1980. Density-dependent growth of Bristol Bay sockeye salmon. Pages 267-283 in W. McNeil, W. and D. Himsworth (eds.), Salmonid Ecosystems of the North Pacific. Oregon State Univ. Press.

Rogers, D.E. 1984. Trends in abundance of northeastern Pacific stocks of salmon. Pp. 100-127 in W.G. Pearcy (ed.), The Influence of Ocean Conditions on the Production of Salmonids in the North Pacific. Oregon State Univ. Press, Corvallis.

Rogers, D.E. 1987. The regulation of age at maturity in Wood River sockeye salmon. Can. Spec. Publ. Fish. Aquat. Sci. 96:78-89.

Rogers, D.E. 1988. Bristol Bay smolt migrations: timing and size composition and the effects on distribution and survival at sea. Pages 87-101 in W. J. McNeil (ed.), Salmon Production Management and Allocation. Oregon State Univ. Press, Corvallis.

Rogers, D.E. and G.T. Ruggerone. 1993. Factors affecting marine growth of Bristol Bay sockeye salmon. Fish. Res. 18:89-103.

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