Range-Wide Selection of Catchments for Pacific Salmon Conservation

Abstract:  Freshwater ecosystems are declining in quality globally, but a lack of data inhibits identification of areas valuable for conservation across national borders. We developed a biological measure of conservation value for six species of Pacific salmon ( Oncorhynchus spp.) in catchments of the northern Pacific across Canada, China, Japan, Russia, and the United States. We based the measure on abundance and life-history richness and a model-based method that filled data gaps. Catchments with high conservation value ranged from California to northern Russia and included catchments in regions that are strongly affected by human development (e.g., Puget Sound). Catchments with high conservation value were less affected by agriculture and dams than other catchments, although only 1% were within biodiversity reserves. Our set of high-value areas was largely insensitive to simulated error, although classification remained uncertain for 3% of catchments. Although salmon face many threats, we propose they will be most likely to exhibit resilience into the future if a complementary mosaic of conservation strategies can be proactively adopted in catchments with healthy salmon populations. Our analysis provides an initial map of where these catchments are likely to be located.     

Population interconnectivity and implications for recovery of a species of concern, the Pacific hake of Georgia Basin

To aid the recovery of a species, understanding the extent to which populations are connected is useful for targeting conservation efforts. Pacific hake within waters of Puget Sound, Washington State, USA, and Georgia Strait, British Columbia, Canada are listed as a species of concern under the U.S. Endangered Species Act due to dramatic declines in the Puget Sound population. To assess the role of dispersal in the recovery of Pacific hake, we sought to quantify patterns of connectivity between populations in Puget Sound and Georgia Strait. Using natural chemical markers from otoliths of fish sampled from these two populations, we linked natal signatures of fish to signatures of individuals from known spawning grounds. Results indicated that 82 % of individuals collected from Puget Sound (n = 78) were estimated to have originated there, while 40 and 92 % of the individuals collected from two cohorts within Georgia Strait (n = 9 and 24, respectively) had originated from Puget Sound. A trend of “population abandonment” of fish from Puget Sound suggests that recovery of this Pacific hake population will depend on local management practices.     

Extinction-effective population index: incorporating life-history variations in population viability analysis

Viability status of populations is a commonly used measure for decision-making in the management of populations. One of the challenges faced by managers is the need to consistently allocate management effort among populations. This allocation should in part be based on comparison of extinction risks among populations. Unfortunately, common criteria that use minimum viable population size or count-based population viability analysis (PVA) often do not provide results that are comparable among populations, primarily because they lack consistency in determining population size measures and threshold levels of population size (e.g., minimum viable population size and quasi-extinction threshold). Here I introduce a new index called the "extinction-effective population index," which accounts for differential effects of demographic stochasticity among organisms with different life-history strategies and among individuals in different life stages. This index is expected to become a new way of determining minimum viable population size criteria and also complement the count-based PVA. The index accounts for the difference in life-history strategies of organisms, which are modeled using matrix population models. The extinction-effective population index, sensitivity, and elasticity are demonstrated in three species of Pacific salmonids. The interpretation of the index is also provided by comparing them with existing demographic indices. Finally, a measure of life-history-specific effect of demographic stochasticity is derived.     

Colonization of the Southern Patagonia Ocean by Exotic Chinook Salmon

Abstract:  Anadromous salmonids have been particularly successful at establishing wild populations in southern Patagonia, in contrast to their limited success elsewhere outside their native ranges. The most recent such discovery is a spawning population of Chinook salmon in the Santa Cruz River, which flows into the Atlantic Ocean from Argentina. We used mitochondrial DNA analysis to discriminate between alternative potential sources of this population and were able to discard in situ introductions of fish imported directly from California in the early twentieth century. Our results showed that the fish most likely came from Puget Sound, Washington, imported into southern Chile for salmon-ranching experiments in the 1980s. This finding provides concrete evidence of colonization of Atlantic rivers from Pacific locations. The southern Pacific and Atlantic oceans provide a favorable marine environment for the success of invading salmon. In particular, the waters associated with fjords, southern channels, and the inshore portion of the Patagonian shelf provide a rather bounded, continuous waterway for exotic anadromous salmonids, rich in diverse forage species.     

Use of trace element analysis of feathers as a tool to track fine-scale dispersal in birds

Tracking dispersal and migratory movements of animals over small and large spatial scales is a challenge. In birds, a promising yet underutilized tool is the trace elemental composition of feathers. The elemental profile of a feather may reveal information about the geographic origin of a bird provided that molting occurs on the breeding grounds and that elemental differences exist between breeding areas. Here, we explore the use of trace elemental composition in body feathers of the Puget Sound white-crowned sparrow Zonotrichia leucophrys pugetensis as a tool (1) to discriminate among birds collected in four different song dialect populations along a 400-km stretch of the Pacific Northwest coast and (2) to assign males singing nonlocal dialects in one population to potential natal populations. Inductively coupled plasma mass spectrometry detected 34 trace elements in sampled feathers and in a discriminant function analysis seven of these elements differed among the four source populations. Half of the six nonlocal dialect singers, who were likely to have immigrated into the focal population, were assigned to a population that matched their song dialect. Our study suggests that feather microchemistry is a promising tool for identifying geographic origins of dispersing birds over small geographic scales and in combination with other markers, such as song, may give insight into ecological and evolutionary processes.     

Public Willingness to Pay for Recovering and Downlisting Threatened and Endangered Marine Species

Abstract: Nonmarket valuation research has produced economic value estimates for a variety of threatened, endangered, and rare species around the world. Although over 40 value estimates exist, it is often difficult to compare values from different studies due to variations in study design, implementation, and modeling specifications. We conducted a stated‐preference choice experiment to estimate the value of recovering or downlisting 8 threatened and endangered marine species in the United States: loggerhead sea turtle (Caretta caretta), leatherback sea turtle (Dermochelys coriacea), North Atlantic right whale (Eubalaena glacialis), North Pacific right whale (Eubalaena japonica), upper Willamette River Chinook salmon (Oncorhynchus tshawytscha), Puget Sound Chinook salmon (Oncorhynchus tshawytscha), Hawaiian monk seals (Monachus schauinslandi), and smalltooth sawfish (Pristis pectinata). In May 2009, we surveyed a random sample of U.S. households. We collected data from 8476 households and estimated willingness to pay for recovering and downlisting the 8 species from these data. Respondents were willing to pay for recovering and downlisting threatened and endangered marine taxa. Willingness‐to‐pay values ranged from $40/household for recovering Puget Sound Chinook salmon to $73/household for recovering the North Pacific right whale. Statistical comparisons among willingness‐to‐pay values suggest that some taxa are more economically valuable than others, which suggests that the U.S. public's willingness to pay for recovery may vary by species.     

Use of Linkage Disequilibrium Data to Estimate Effective Size of Hatchery and Natural Fish Populations

A primary parameter in the assessment of the viability of a population is its effective population size ( Ne ). Allozyme analysis of four groups of fishes provided data on linkage disequilibrium, which were then used to estimate Ne . The groups included hatchery samples of juvenile white seabass, Atractoscion nobilis , juvenile rainbow trout, Oncorhynchus mykiss , from the Shasta Hatchery, and juvenile chinook salmon, O. tshawytscha , from the Coleman National Fish Hatchery. The fourth sample consisted of juvenile chinook salmon from the threatened winter run in the upper Sacramento River. The groups of fish were chosen to represent different applications of the methodology to conservation of fishes. For a variety of reasons. Ne may be considerably lower than census counts of fish present in the parental populations. The Ne of the hatchery broodstock that produced the sample of juvenile white seabass was estimated to be approximately 10, although 25 adult white seabass were present in a mass spawning tank. Ne estimates for the parental populations of the Shasta and Coleman Hatchery samples were 35.8 and 132.5, respectively. The actual number of fish spawned at the Shasta Hatchery was approximately 40, whereas nearly 10,000 salmon were spawned at the Coleman Hatchery. The threatened winter run of chinook salmon had an estimated Ne of 85.5 and an approximate run size of 2000 salmon. The method of estimating effective population size from linkage disequilibrium data appears to result in realistic estimates of effective population size when adequate sample size and a sufficient number of polymorphic loci are available.     

Reconstruction of Pacific salmon abundance from riparian tree-ring growth.

We use relationships between modern Pacific salmon (Oncorhynchus spp.) escapement (migrating adults counted at weirs or dams) and riparian tree-ring growth to reconstruct the abundance of stream-spawning salmon over 150-350 years. After examining nine sites, we produced reconstructions for five mid-order rivers and four salmon species over a large geographic range in the Pacific Northwest: chinook (O. tschwatcha) in the Umpqua River, Oregon, USA; sockeye (O. nerka) in Drinkwater Creek, British Columbia, Canada; pink (O. gorbuscha) in Sashin Creek, southeastern Alaska, USA; chum (O. keta) in Disappearance Creek, southeastern Alaska, USA; and pink and chum in the Kadashan River, southeastern Alaska, USA. We first derived stand-level, non-climatic growth chronologies from riparian trees using standard dendroecology methods and differencing. When the chronologies were compared to 18-55 years of adult salmon escapement we detected positive, significant correlations at five of the nine sites. Regression models relating escapement to tree-ring growth at the five sites were applied to the differenced chronologies to reconstruct salmon abundance. Each reconstruction contains unique patterns characteristic of the site and salmon species. Reconstructions were validated by comparison to local histories (e.g., construction of dams and salmon canneries) and regional fisheries data such as salmon landings and aerial surveys and the Pacific Decadal Oscillation climate index. The reconstructions capture lower-frequency cycles better than extremes and are most useful for determination and comparison of relative abundance, cycles, and the effects of interventions. Reconstructions show lower population cycle maxima in both Umpqua River chinook and Sashin Creek pink salmon in recent decades. The Drinkwater Creek reconstruction suggests that sockeye abundance since the mid-1990s has been 15-25% higher than at any time since 1850, while no long-term deviations from natural cycles are detected for salmon in the Kadashan River or in Disappearance Creek. Decadal-scale cycles in salmon abundance with periods of 25-68 years were detected in all of the reconstructions. This novel approach provides river-specific, long-term perspectives on salmon abundance and cycles. Additionally, it provides a new frame of reference for maintaining and rebuilding individual stocks and for striking a balance between societal demands and the limited, always-changing salmon resource.     

Two populations of the marine fish Porichthys notatus,one lacking in luciferin essential for bioluminescence

Porichthys notatus is a common benthic fish found along the Pacific coast of North America. It possesses more than 700 small dermal photophores on its head and trunk. When P. notatus collected in Monterey Bay, California, is injected subcutaneously with norepinephrine, the photophores emit a long-lasting luminescence that is readily visible to the dark-adapted eye. The light emission is due to the oxidation of luciferin (substrate) by molecular oxygen, catalyzed by luciferase. In contrast, P. notatus collected in Puget Sound, Washington, is nonluminous, even though the photophores do not differ ultrastructurally from those of the California fish. The inability of the Puget Sound fish to luminesce is due to lack of luciferin in its photophores. Luminescence capability, however, may be induced in the Puget Sound fish by the oral or intraperitoneal administration of a small amount of luciferin from the tiny luminescent marine ostracod crustacean, Vargula hilgendorfii, suggesting that the origin of luciferin in P. notatus may be from the diet. In this study, specimens of P. notatus were collected over the entire known range of the species, between November 1981 and September 1987, and the presence of luciferin in the photophores determined. The results indicate that there are two populations of P. notatus: a luciferin-deficient, nonluminescent population located north of northern California and a luciferin-containing, luminescent population extending south of Cape Mendocino, California, to Baja California, Mexico. At the northern end of the southern population, a mixture of luminescent and nonluminescent P. notatus was found.     

Zinc and cadmium in the Pacific hake Merluccius productus off the western U.S. coast

Concentrations of zinc and cadmium in muscle tissue or whole Pacific hake, Merluccius productus (Ayres), increase with increasing total wet weight, especially for smaller fish. These trends appear to be related to the euphausiid diet of hake. Except for the isolated population in Puget Sound, no relationship between zinc or cadmium concentration and collection site is apparent.     

Latitudinal trends in size-dependence of bioluminescence in the midshipman fish Porichthys notatus

The bioluminescent fish Porichthys notatus (plainfin midshipman), has a discontinuous distribution along the Pacific coast of North America. The fish is present from Cape Mendocino southward to Baja California, Mexico, absent off the coast of Oregon, USA, and abundant, northward, in Puget Sound, Washington. Interestingly, the population in Puget Sound lacks the substrate (luciferin) necessary for the luminescence reaction and, despite possessing an otherwise fully functional photophore system, is nonluminescent. The California population of P. notatus is uniformly luminescent south of Monterey Bay, but 15% of the speciments tested from San Francisco Bay and the Gulf of the Farallons have been reported to be nonluminescent. Explanations for nonluminescent midshipman in both Puget Sound and the San Francisco Bay area have focussed on a dietary requirement for luciferin. To gain further insight into reasons for nonluminescence in the San Francisco Bay region, the distribution of bioluminescence in P. notatus was studied from Monterey Bay to Cape Mendocino during 1985. A complex pattern of bioluminescence was found, in which nonluminescent individuals reflected neither a local anomaly in the San Francisco Bay region nor a simple gradient of decreasing luminescence towards the northern end of the range of the California population. Instead, a distinct size-dependent component in luminescence capability of the fish was observed. Aspects of the life history of P. notatus and related factors which might influence the bioluminescence characteristics of this population are discussed.Please address all correspondence to Dr. F.I. Tsuji at the Osaka Bioscience Institute     

Identifying interactions among salmon populations from observed dynamics

A simple direct correlation analysis of individual counts between different populations often fails to characterize the true nature of population interactions; however, the most common data type available for population studies is count data, and one of the most important objectives in population and community ecology is to identify interactions among populations. Here, I examine the dynamics of the spawning abundance of fall-run chinook salmon spawning within the California Central Valley and the Klamath Basin, California, and the Columbia River Basin, Oregon. I analyzed multiple time series from each watershed using a multivariate time-series technique called maximum autocorrelation factor analysis. This technique was used for finding common underlying trends in escapement abundance within each watershed. These trends were further investigated to identify potential resource-mediated interactions among the three groups of salmon. Each group is affected by multiple trends that are likely to be affected by environmental factors. In addition, some of the trends are coherent with each other, and the differences in population dynamics originate from variations in the relative importance of these trends among the three watershed groups.     

High mercury bioaccumulation in Pacific salmons from the Sea of Okhotsk and the Bering Sea

Mercury (Hg) accumulation in fishes is a serious threat for food security because mercury induces dieseases. Here we studied the total mercury content in four species of Pacific salmon (pink, chum, sockeye and chinook), caught in the Sea of Okhotsk, the Bering Sea, and the Kuril oceanic waters. Results show that the lowest concentration of mercury was found in the pink salmon, and the highest was 5–6 times higher in sockeye. Hg concentrations exceed several times those in other species of Pacific salmon from the North Pacific     

Temperature-associated population diversity in salmon confers benefits to mobile consumers

Habitat heterogeneity can generate intraspecific diversity through local adaptation of populations. While it is becoming increasingly clear that population diversity can increase stability in species abundance, less is known about how population diversity can benefit consumers that can integrate across population diversity in their prey. Here we demonstrate cascading effects of thermal heterogeneity on trout-salmon interactions in streams where rainbow trout rely heavily on the seasonal availability of anadromous salmon eggs. Water temperature in an Alaskan stream varied spatially from 5 degrees C to 17.5 degrees C, and spawning sockeye salmon showed population differentiation associated with this thermal heterogeneity. Individuals that spawned early in cool regions of the 5 km long stream were genetically differentiated from those spawning in warmer regions later in the season. Sockeye salmon spawning generates a pulsed resource subsidy that supports the majority of seasonal growth in stream-dwelling rainbow trout. The spatial and temporal structuring of sockeye salmon spawn timing in our focal stream extended the duration of the pulsed subsidy compared to a thermally homogeneous stream with a single population of salmon. Further, rainbow trout adopted movement strategies that exploited the multiple pulses of egg subsidies in the thermally heterogeneous stream. Fish that moved to track the resource pulse grew at rates about 2.5 times higher than those that remained stationary or trout in the reference stream with a single seasonal pulse of eggs. Our results demonstrate that habitat heterogeneity can have important effects on the population diversity of dominant species, and in turn, influence their value to species that prey upon them. Therefore, habitat homogenization may have farther-reaching ecological effects than previously considered.     

Incorporating covariates into fisheries stock assessment models with application to Pacific herring.

We present a framework for evaluating the cause of fishery declines by integrating covariates into a fisheries stock assessment model. This allows the evaluation of fisheries' effects vs. natural and other human impacts. The analyses presented are based on integrating ecological science and statistics and form the basis for environmental decision-making advice. Hypothesis tests are described to rank hypotheses and determine the size of a multiple covariate model. We extend recent developments in integrated analysis and use novel methods to produce effect size estimates that are relevant to policy makers and include estimates of uncertainty. Results can be directly applied to evaluate trade-offs among alternative management decisions. The methods and results are also broadly applicable outside fisheries stock assessment. We show that multiple factors influence populations and that analysis of factors in isolation can be misleading. We illustrate the framework by applying it to Pacific herring of Prince William Sound, Alaska (USA). The Pacific herring stock that spawns in Prince William Sound is a stock that has collapsed, but there are several competing or alternative hypotheses to account for the initial collapse and subsequent lack of recovery. Factors failing the initial screening tests for statistical significance included indicators of the 1989 Exxon Valdez oil spill, coho salmon predation, sea lion predation, Pacific Decadal Oscillation, Northern Oscillation Index, and effects of containment in the herring egg-on-kelp pound fishery. The overall results indicate that the most statistically significant factors related to the lack of recovery of the herring stock involve competition or predation by juvenile hatchery pink salmon on herring juveniles. Secondary factors identified in the analysis were poor nutrition in the winter, ocean (Gulf of Alaska) temperature in the winter, the viral hemorrhagic septicemia virus, and the pathogen Ichthyophonus hoferi. The implication of this result to fisheries management in Prince William Sound is that it may well be difficult to simultaneously increase the production of pink salmon and maintain a viable Pacific herring fishery. The impact can be extended to other commercially important fisheries, and a whole ecosystem approach may be needed to evaluate the costs and benefits of salmon hatcheries.     

Population viability analysis for several populations using multivariate state-space models

The International Union for the Conservation of Nature and Natural Resources (IUCN), the world's largest and most important global conservation network, has listed approximately 16,000 species worldwide as threatened. The most important tool for recognizing and listing species as threatened is population viability analysis (PVA), which estimates the probability of extinction of a population or species over a specified time horizon. The most common PVA approach is to apply it to single time series of population abundance. This approach to population viability analysis ignores covariability of local populations. Covariability can be important because high synchrony of local populations reduces the effective number of local populations and leads to greater extinction risk. Needed is a way of extending PVA to model correlation structure among multiple local populations. Multivariate state-space modeling is applied to this problem and alternative estimation methods are compared. The multivariate state-space technique is applied to endangered populations of pacific salmon, USA. Simulations demonstrated that the correlation structure can strongly influence population viability and is best estimated using restricted maximum likelihood instead of maximum likelihood.     

Population structure of the black tiger prawn, Penaeus monodon, among western Indian Ocean and western Pacific populations

We examined the population structure of the black tiger prawn, Penaeus monodon Fabricius, 1798, in the Indo-West Pacific by analyzing the geographic distribution of elongation factor 1-alpha intron sequences from specimens collected during the winter and spring of 1997. Both the molecular phylogeny of alleles and F-statistics indicated very strong differentiation between populations from the western Indian Ocean and western Pacific. This pattern is concordant with other recent studies of marine species in this region, implying that the Indo-Australian Archipelago represents a biogeographic break between populations in the Indo-West Pacific. F _ST-values among populations in the western Indian Ocean also indicate structure within this region, whereas no structure was found among western Pacific populations. Nucleotide diversity was significantly lower in the western Indian Ocean populations than in the western Pacific, implying that the populations have regional differences in demographic history. Received: 16 November 1998 / Accepted: 26 May 1999     

Long-term demographic and genetic effects of releasing captive-born individuals into the wild

Because of continued habitat destruction and species extirpations, the need to use captive breeding for conservation purposes has been increasing steadily. However, the long-term demographic and genetic effects associated with releasing captive-born individuals with varied life histories into the wild remain largely unknown. To address this question, we developed forward-time, agent-based models for 4 species with long-running captive-breeding and release programs: coho salmon (Oncorhynchus kisutch), golden lion tamarin (Leontopithecus rosalia), western toad (Anaxyrus boreas), and Whooping Crane (Grus americana). We measured the effects of supplementation by comparing population size and neutral genetic diversity in supplemented populations to the same characteristics in unaltered populations 100 years after supplementation ended. Releasing even slightly less fit captive-born individuals to supplement wild populations typically resulted in reductions in population sizes and genetic diversity over the long term when the fitness reductions were heritable (i.e., due to genetic adaptation to captivity) and populations continued to be regulated by density-dependent mechanisms over time. Negative effects for species with longer life spans and lower rates of population replacement were smaller than for species with shorter life spans and higher rates of population replacement. Programs that released captive-born individuals over fewer years or that avoided breeding individuals with captive ancestry had smaller reductions in population size and genetic diversity over the long term. Relying on selection in the wild to remove individuals with reduced fitness mitigated some negative demographic effects, but at a substantial cost to neutral genetic diversity. Our results suggest that conservation-focused captive-breeding programs should take measures to prevent even small amounts of genetic adaptation to captivity, quantitatively determine the minimum number of captive-born individuals to release each year, and fully account for the interactions among genetic adaptation to captivity, population regulation, and life-history variation.     

Co-existing populations of Pacific ocean perch, Sebastes alutus, in Queen Charlotte Sound, British Columbia

Variation at five microsatellite loci (Sal1, Sal2, Sal3, Sal4 and Sal5) was examined in approximately 1300 Pacific ocean perch (Sebastes alutus) sampled from 14 coastal sites in British Columbia, Canada. Mean observed heterozygosities by locus ranged from 71% to 88%, and by sample ranged from 75% to 84%. Theta values ranged from 0 to 0.04 over the five loci, and averaged 0.015. Among Pacific ocean perch samples, Š ranged from 0.001 to 0.056. Canonical discriminate analysis of multilocus genotypes and neighbour-joining analysis of pairwise genetic distances between samples both indicated the presence of three populations, one off the west coast of Vancouver Island (the Vancouver Island population) and two co-existing populations in Queen Charlotte Sound, Dixon Entrance and along the west coast of the Queen Charlotte Islands (the eastern and western QCI populations). Pacific ocean perch of the eastern and western QCI populations were caught in close proximity to each other, but individual samples showed little evidence of admixture. Fall and spring samples collected within geographic areas were genetically similar, indicating seasonally stable population structure. Restricted gene flow between the Vancouver Island and the two more northerly populations may result from limited adult dispersal and larval retention within the California Current and Alaska Gyre, respectively, but the presence of two populations within Queen Charlotte Sound cannot be explained entirely by larval retention hypotheses. The presence of two Pacific ocean perch populations in central British Columbia has implications for fisheries management.     

Evidence for multiple recruitment-cohorts of Puget Sound Dungeness crab,Cancer magister

While sampling intertidally in Puget Sound, Washington, USA, for juvenile Dungeness crab (Cancer magister) in 1984, we found evidence of two distinct cohorts of the same year-class based on sizes of first-instar juveniles (J1) and the spatial/temporal patterns of settlement. In 1988, three distinct cohorts were observed to settle in Puget Sound and its approaches. Settlement of one cohort occurred during May in the Strait of Juan de Fuca and in those areas of Puget Sound closest to the Strait. J1 individuals of this cohort were large (x=7.4 mm carapace width, CW) and comparable in both size and timing of settlement to populations along the Washington coast (e.g. Grays Harbor and Willapa Bay). Initial settlement density of the May cohort was as high as 215 crabs/m² in intertidal eelgrass beds along the Strait of Juan de Fuca and decreased to <2 crabs/m² within Puget Sound and the Strait of Georgia. A second cohort apparently originated in Hood Canal (a deep inland fjord), its size upon settlement in June was significantly smaller (J1 x= 5.3 mm CW) than the May cohort, and it was limited to Hood Canal and areas of Puget Sound close to the mouth of Hood Canal. A third cohort, which settled in late July and August, was the smallest of the three cohorts (J1 x= 4.8 mm CW), and was widely distributed around Puget Sound from Seattle in the south to the USA/Canadian border in the north. We hypothesize that most juvenile recruitment in Hood Canal and Puget Sound originates from parental stocks endemic to their respective basins (Hood Canal and Puget Sound cohorts), but that, on occasion, oceanographic conditions allow substantial influx of Pacific Ocean Dungeness crab larvae (oceanic cohort) through the Strait of Juan de Fuca into Puget Sound. Tracking of spatial/temporal settlement patterns and comparison of J1 sizes proved useful for estimating the probable sources and dispersion of Dungeness crab larvae. Differences in size and time of settlement between various larval cohorts of C. magister may prove useful as biomarkers for tracing circulation patterns within and between inland waters of Washington and the Pacific Ocean. Causes of smaller size and later settlement of the Puget Sound cohort relative to oceanic conspecifics of the same year-class are discussed.Contribution No. 856, School of Fisheries, University of Washington, Seattle