Effects of the North Pacific Current on the productivity of 163 Pacific salmon stocks

Horizontal ocean transport can influence the dynamics of higher‐trophic‐level species in coastal ecosystems by altering either physical oceanographic conditions or the advection of food resources into coastal areas. In this study, we investigated whether variability in two North Pacific Current (NPC) indices was associated with changes in productivity of North American Pacific salmon stocks. Specifically, we used Bayesian hierarchical models to estimate the effects of the north‐south location of the NPC bifurcation (BI) and the NPC strength, indexed by the North Pacific Gyre Oscillation (NPGO), on the productivity of 163 pink, chum, and sockeye salmon stocks. We found that for salmon stocks located in Washington (WA) and British Columbia (BC), both the BI and NPGO had significant positive effects on productivity, indicating that a northward‐shifted bifurcation and a stronger NPC are associated with increased salmon productivity. For the WA and BC regions, the estimated NPGO effect was over two times larger than the BI effect for pink and chum salmon, whereas for sockeye salmon the BI effect was 2.4 times higher than the NPGO. In contrast to WA and BC stocks, we found weak effects of both horizontal ocean transport processes on the productivity of salmon stocks in Alaska. Our results indicated that horizontal transport pathways might strongly influence population dynamics of Pacific salmon in the southern part of their North American ranges, but not the northern part, suggesting that different environmental pathways may underlie changes in salmon productivity in northern and southern areas for the species under consideration.     

Linking Northeast Pacific recruitment synchrony to environmental variability

We investigated the hypothesis that synchronous recruitment is due to a shared susceptibility to environmental processes using stock–recruitment residuals for 52 marine fish stocks within three Northeast Pacific large marine ecosystems: the Eastern Bering Sea and Aleutian Islands, Gulf of Alaska, and California Current. There was moderate coherence in exceptionally strong and weak year‐classes and correlations across stocks. Based on evidence of synchrony from these analyses, we used Bayesian hierarchical models to relate recruitment to environmental covariates for groups of stocks that may be similarly influenced by environmental processes based on their life histories. There were consistent relationships among stocks to the covariates, especially within the Gulf of Alaska and California Current. The best Gulf of Alaska model included Northeast Pacific sea surface height as a predictor of recruitment, and was particularly strong for stocks dependent on cross‐shelf transport during the larval phase for recruitment. In the California Current the best‐fit model included San Francisco coastal sea level height as a predictor, with higher recruitment for many stocks corresponding to anomalously high sea level the year before spawning and low sea level the year of spawning. The best Eastern Bering Sea and Aleutian Islands model included several environmental variables as covariates and there was some consistent response across stocks to these variables. Future research may be able to utilize these across‐stock environmental influences, in conjunction with an understanding of ecological processes important across early life history stages, to improve identification of environmental drivers of recruitment.     

Interannual and spatial variation in the population genetic composition of young‐of‐the‐year Pacific ocean perch (Sebastes alutus) in the Gulf of Alaska

Little is known about the population structure of Alaskan rockfishes, including Pacific ocean perch (POP, Sebastes alutus), and how persistent and variable oceanographic features may influence their structures. Moreover, early life history information is sparse for many species. We used data from 14 microsatellite loci to characterize the genetic structure of young‐of‐the‐year Pacific ocean perch collected during 1998–2003 from the Gulf of Alaska and Bering Sea. Broad‐scale geographic variation in genetic structure of the young‐of‐the‐year (F_ST = 0.005, P < 10^?4) had similarities to that observed in a previous adult study. The overall correlation between genetic and geographic distance (isolation by distance) was nearly identical to that observed in the adults. Fine‐scale geographic divergence was also observed and may be the result of oceanographic circulation features within the Gulf of Alaska. Interannual variation (between cohorts) at locations sampled in more than one year is consistent with variable oceanography and fine‐scale population structure rather than the influence of a sweepstakes effect. The similarities of the young‐of‐the‐year with the adults and the pattern of genetic divergence confirm that dispersal of Pacific ocean perch is limited in all life stages.     

Establishing climate–growth relationships for yelloweye rockfish (Sebastes ruberrimus) in the northeast Pacific using a dendrochronological approach

We applied dendrochronology (tree‐ring) methods to develop multidecadal growth chronologies from the increment widths of yelloweye rockfish (Sebastes ruberrimus) otoliths. Chronologies were developed for the central California coast, a site just north of Vancouver Island, British Columbia, and at Bowie Seamount west of the Queen Charlotte Islands, British Columbia. At each site, synchronous growth patterns were matched among otoliths via the process of cross‐dating, ensuring that the correct calendar year was assigned to all increments. Each time series of growth‐increment measurements was divided by the values predicted by a best‐fit negative exponential function, thereby removing age‐related trends. These detrended time series were averaged into a master chronology for each site, and chronologies were correlated with monthly averages of sea surface temperatures, upwelling, the Northern Oscillation Index, and the Pacific Decadal Oscillation. The two northern growth chronologies positively correlated with indices of warm ocean conditions, especially from the prior summer through the spring of the current year. During the same period, the California chronology positively correlated with indices of cool ocean conditions, indicating an opposing productivity regime for yelloweye rockfish between the California Current and the Gulf of Alaska. Overall, this study demonstrates how tree‐ring techniques can be applied to quickly develop annually resolved chronologies and establish climate–growth relationships across various temporal and spatial scales.     

Mid-latitude wind stress: the energy source for climatic shifts in the North Pacific Ocean

Analyses of atmospheric observations in the North Pacific demonstrate extensive decadal-scale variations in the mid-latitude winter surface wind stress. In the decade after 1976 winter, eastward wind stress doubled over a broad area in the central North Pacific and the winter zero wind stress curl line was displaced about 6° southward. This resulted in increased southward Ekman transport, increased oceanic upwelling, and increased turbulent mixing as well as a southward expansion of the area of surface divergence. All these factors contributed to a decadal winter cold anomaly along the subtropical side of the North Pacific Current. In summer the cold anomaly extended eastward, almost reaching the coast of Oregon. The increased gradient in wind stress curl and southward displacement of the zero curl line also resulted in an increase in total North Pacific Current transport, primarily on the Equator side of this Current. Thus, surface water entering the California Current was of more subtropical origin in the post-1976 decade. Southward (upwelling favourable) wind stress and sea surface temperature (SST) in the area off San Francisco exhibit at least three different types of decadal departures from mean conditions. In association with the 1976 climatic shift, marine fishery production in the Oyashio, California and Alaska Currents altered dramatically, suggesting that these natural environmental variations significantly alter the long-term yields of major North Pacific fisheries.     

Biogeography of pelagic food webs in the North Pacific

The tufted puffin (Fratercula cirrhata) is a generalist seabird that breeds throughout the North Pacific and eats more than 75 different prey species. Using puffins as samplers, we characterized the geographic variability in pelagic food webs across the subarctic North Pacific from the composition of ~10,000 tufted puffin meals (~56,000 prey items) collected at 35 colonies in the Gulf of Alaska (GoA) and Aleutian Archipelago. Cluster analysis of diet species composition suggested three distinct forage fish communities: (i) in the northern GoA, multiple age‐classes of coastal and shelf residents such as capelin, sand lance and herring dominated the food web, (ii) in the western GoA to eastern Aleutians, the shelf community was dominated by transient age‐0 walleye pollock, and (iii) in the western Aleutians, shelf‐edge and mesopelagic forage species such as squid, lanternfish, and Atka mackerel were prevalent. Geographic patterns of abundance of capelin and sand lance in tufted puffin diets were corroborated by independent research fisheries and diets of piscivorous fish, indicating that puffin diets reflect the local abundance of forage species, not just selection of favored species. Generalized additive models showed that habitat characteristics predict, in a non‐linear fashion, forage species distribution and abundance across two large marine ecosystems. We conclude that major biogeographic patterns in forage fish distribution follow gradients in key habitat features, and puffin diets reflect those patterns.     

Distribution and community structure of ichthyoplankton from the northern and central California Current in May 2004–06

The distributions, concentrations, and community structure of pelagic larval fishes collected from the central and northern California Current in the northeast Pacific Ocean during May 2004, 2005, and 2006 were analyzed to investigate inter-annual, latitudinal, cross-shelf, and depth-stratified variability. The inter-annual climate-induced variability during the sampling period provided a unique opportunity to observe how larval fish communities adjust to rapidly changing environmental conditions. The 170 depth-stratified samples collected from three cruises yielded 14 819 fish larvae from 56 taxa representing 23 families. Dominant larval taxa were Engraulis mordax , Citharichthys spp., Sebastes spp., and Stenobrachius leucopsarus . Larval concentrations decreased significantly in 2006 from 2004 and 2005 levels following the anomalous oceanic conditions observed in 2005 and decreased water temperature in 2006. Larvae were generally found in higher concentrations at northern (>43°N) versus southern (<43°N) stations, with larval E. mordax and Citharichthys spp. found almost exclusively in the north during all sampled years. Inter-annual variability related to dynamic upwelling intensity was observed in cross-shelf larval distributions, although concentrations of S. leucopsarus larvae consistently increased in the offshore direction, while larval Sebastes spp. were generally found in highest concentrations at intermediate stations along the shelf. Multivariate analyses revealed that latitude, station depth, and sea-surface temperature were the most important factors explaining variability in larval concentrations. The present study shows that the ichthyoplankton community of the central and northern California Current changed dramatically in response to the variable environmental conditions of 2004–06.     

Potential trophodynamic and environmental drivers of steelhead (Oncorhynchus mykiss) productivity in the North Pacific Ocean

Information on prey availability, diets, and trophic levels of fish predators and their prey provides a link between physical and biological changes in the ecosystem and subsequent productivity (growth and survival) of fish populations. In this study two long‐term data sets on summer diets of steelhead (Oncorhynchus mykiss) in international waters of the central North Pacific Ocean (CNP; 1991–2009) and Gulf of Alaska (GOA; 1993–2002) were evaluated to identify potential drivers of steelhead productivity in the North Pacific. Stable isotopes of steelhead muscle tissue were assessed to corroborate the results of stomach content analysis. We found the composition of steelhead diets varied by ocean age group, region, and year. In both the GOA and CNP, gonatid squid (Berryteuthis anonychus) were the most influential component of steelhead diets, leading to higher prey energy densities and stomach fullness. Stomach contents during an exceptionally warm year in the GOA and CNP (1997) were characterized by high diversity of prey with low energy density, few squid, and a large amount of potentially toxic debris (e.g., plastic). Indicators of good diets (high proportions of squid and high prey energy density) were negatively correlated with abundance of wild populations of eastern Kamchatka pink salmon (O. gorbuscha) in the CNP. In conclusion, interannual variations in climate, abundance of squid, and density‐dependent interactions with highly‐abundant stocks of pink salmon were identified as potential key drivers of steelhead productivity in these ecosystems. Additional research in genetic stock identification is needed to link these potential drivers of productivity to individual populations.     

Time‐varying relationships between ocean conditions and sockeye salmon productivity

Environmental change is occurring at unprecedented rates in many marine ecosystems. Yet, environmental effects on fish populations are commonly assumed to be constant across time. In this study, I tested whether relationships between ocean conditions and productivity of North American sockeye salmon (Oncorhynchus nerka) stocks have changed over the past six decades. Specifically, I evaluated the evidence for non‐stationary relationships between three widely used ocean indices and productivity of 45 sockeye salmon stocks using hierarchical Bayesian models. The ocean indices investigated were the Pacific Decadal Oscillation (PDO), North Pacific Gyre Oscillation (NPGO), and sea surface temperature (SST). I found partial support for time‐varying salmon–ocean relationships. Non‐stationary relationships were strongest for the NPGO and weaker for the SST and PDO indices. Productivity–NPGO correlations tended to shift gradually over time with opposite trends for stocks in British Columbia (B.C.) and western Alaska; for B.C. stocks, the NPGO correlations shifted from significantly negative prior to 1980 to significantly positive after 1990, whereas for western Alaska stocks, the correlations shifted from positive to negative. Productivity–SST correlations showed declining trends for B.C. and Gulf of Alaska stocks, that is, correlations became more negative (B.C.) or less positive (Gulf of Alaska) over time. For the PDO, correlations weakened during the 1980s for western Alaska and B.C. stocks. Overall, these results provide evidence for time‐varying relationships between salmon productivity and environmental conditions over six decades, highlighting the need to recognize that historical responses of salmon populations to environmental change may not be indicative of future responses.     

Change in feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in the central Gulf of Alaska in relation to climate events

The effects of climate events on the feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in offshore waters of the central Gulf of Alaska were investigated during early summers (1994–2000), based on analyses of stomach contents, and carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N). Gonatid squids (mainly Berryteuthis anonychus) were the dominant prey of all salmon species except for chum salmon (O. keta). During the 1997 El Niño event and the 1999 La Niña event, squids decreased sharply in the diets of all Pacific salmon except coho salmon (O. kisutch) in the Subarctic Current, and chum salmon diets changed from gelatinous zooplankton (1995–97) to a more diverse array of zooplankton species. A δ¹³C and δ¹⁵N analysis indicated that all salmon species occupied the same branch of the food web in 1999–2000. We hypothesize that high‐seas salmon adapt to climate‐induced changes in their prey resources by switching their diets either within or between trophic levels. To understand the effects of climate change on Pacific salmon in the Gulf of Alaska, biological oceanographic research on B. anonychus and other important prey resources is needed.     

How “The Blob” affected groundfish distributions in the Gulf of Alaska

We investigated the distributional shifts of groundfish in response to anomalous ocean conditions, particularly the recent anomalously warm period (2014–2016; “The Blob”), based on data from ten Gulf of Alaska bottom trawl surveys conducted by the Alaska Fisheries Science Center during 1996–2015. Six groundfish species were considered: Pacific cod (Gadus macrocephalus), arrowtooth flounder (Atheresthes stomias), walleye pollock (Gadus chalcogrammus), Pacific ocean perch (Sebastes alutus), northern rock sole (Lepidopsetta polyxystra), and southern rock sole (Lepidopsetta bilineata). Ontogenetic differences were examined by dividing data for each fish species into size classes. Our study demonstrated that after accounting for size‐specific depth preferences, the spatial responses of groundfish to anomalous ocean conditions differed by species and foraging guild in the central Gulf of Alaska. Pacific cod and arrowtooth flounder showed similar responses to ocean warming, but different responses to cooling. In general, Pacific cod moved to deeper depths in warmer years and moved to shallower depths in colder years. Arrowtooth flounder also moved deeper in warmer years. However, in colder years, large arrowtooth flounder (>40 cm) shifted toward shallower depths while smaller‐sized fish shifted toward deeper depths. In warmer years, large pollock (>30 cm) moved to deeper waters while smaller pollock (10–20 cm) moved to shallower waters. Pacific ocean perch exhibited an opposite response to thermal changes in habitat compared with Pacific cod and arrowtooth flounder. They moved deeper in colder years, but there was no clear change in depth as a function of size in response to warmer habitat.     

Bottom-up forcing and the decline of Steller sea lions (Eumetopias jubatus) in Alaska: assessing the ocean climate hypothesis

Declines of Steller sea lion (Eumetopias jubatus) populations in the Aleutian Islands and Gulf of Alaska could be a consequence of physical oceanographic changes associated with the 1976–77 climate regime shift. Changes in ocean climate are hypothesized to have affected the quantity, quality, and accessibility of prey, which in turn may have affected the rates of birth and death of sea lions. Recent studies of the spatial and temporal variations in the ocean climate system of the North Pacific support this hypothesis. Ocean climate changes appear to have created adaptive opportunities for various species that are preyed upon by Steller sea lions at mid‐trophic levels. The east–west asymmetry of the oceanic response to climate forcing after 1976–77 is consistent with both the temporal aspect (populations decreased after the late 1970s) and the spatial aspect of the decline (western, but not eastern, sea lion populations decreased). These broad‐scale climate variations appear to be modulated by regionally sensitive biogeographic structures along the Aleutian Islands and Gulf of Alaska, which include a transition point from coastal to open‐ocean conditions at Samalga Pass westward along the Aleutian Islands. These transition points delineate distinct clusterings of different combinations of prey species, which are in turn correlated with differential population sizes and trajectories of Steller sea lions. Archaeological records spanning 4000 yr further indicate that sea lion populations have experienced major shifts in abundance in the past. Shifts in ocean climate are the most parsimonious underlying explanation for the broad suite of ecosystem changes that have been observed in the North Pacific Ocean in recent decades.     

Climate‐induced nonlinearity in pelagic communities and non‐stationary relationships with physical drivers in the Kuroshio ecosystem

Climate‐induced nonlinearity in biological variability and non‐stationary relationships with physical drivers are crucial to understand responses of marine organisms to climate variability. These phenomena have raised concerns in the northeastern North Pacific, but are out of the spotlight in the northwestern North Pacific in spite of potential implications for this productive system under increased climate variability. Pelagic communities in the Kuroshio ecosystem have both ecological and economic importance. However, patterns of climate‐induced nonlinearity in pelagic communities are not well understood, and existence of non‐stationarity in their relationships with physical drivers remains obscure. Here, we compile large numbers of climatic, oceanic and biological long‐term time‐series data and employ diverse statistical techniques to reveal such climate‐induced nonlinearity and non‐stationarity. Results show that pelagic communities in the Tsushima and Pacific areas (major areas in the Kuroshio ecosystem) had regime shifts in the late 1990s and late 1980s, respectively. Winter sea surface temperatures in the Kuroshio Current path and in the eastern part of East China Sea, which are respectively affected by the Kuroshio Current and Siberian High, correlate with dominant variability patterns in their pelagic communities. Furthermore, non‐stationarity was identified with threshold years in the 1990s in the Tsushima area and in the 1980s in the Pacific area as a possible result of the declined variances in the Siberian High and Aleutian Low, respectively. Our findings provide insights on spatial differentiation of climate‐induced nonlinearity and non‐stationarity, which are valuable for the management of pelagic communities in the northwestern North Pacific under changing climatic conditions.     

Influence of mesoscale eddies on ichthyoplankton assemblages in the Gulf of Alaska

Mesoscale eddies (100–200 km in diameter) propagating along the shelf‐break in the Gulf of Alaska are ubiquitous and have been shown to influence the ecosystem, but their influence on ichthyoplankton species composition and diversity has not been described. Evidence for larval fish entrainment in these eddies was examined using data from a cruise in 2005 that sampled three eastern Gulf of Alaska mesoscale eddies, and sampling that compared shelf to slope ichthyoplankton assemblages in the northern Gulf of Alaska (2002–2004). Hierarchical cluster analysis of oceanographic data showed that stations grouped according to location within an eddy. Species hierarchical cluster analysis revealed a latitudinal turnover in species composition, and an abundant species group. Species richness was correlated with distance from eddy center (P = 0.00025), and assemblages within eddies were significantly different (P < 0.05) from those in surrounding basin and shelf waters. These results suggest that mesoscale eddies propagating along the continental shelf‐break influence larval fish assemblages over the shelf and slope, which has implications for the timing and extent of larval fish distribution in the Gulf of Alaska.     

Covariation between the average lengths of mature coho (Oncorhynchus kisutch) and Chinook salmon (O. tshawytscha) and the ocean environment

We used the average fork length of age‐3 returning coho (Oncorhynchus kisutch) and age‐3 ocean‐type and age‐4 stream‐type Chinook (Oncorhynchus tshawytscha) salmon along the northeast Pacific coast to assess the covariability between established oceanic environmental indices and growth. These indices included the Multivariate El Niño‐Southern Oscillation Index (MEI), Pacific Decadal Oscillation (PDO), Northern Oscillation Index, and Aleutian Low Pressure Index. Washington, Oregon, and California (WOC) salmon sizes were negatively correlated with the MEI values indicating that ultimate fish size was affected negatively by El Niño‐like events. Further, we show that the growth trajectory of WOC salmon was set following the first ocean winter. Returning ocean‐type British Columbia‐Puget Sound Chinook salmon average fork length was positively correlated with the MEI values during the summer and autumn of return year, which was possibly a result of a shallower mixed layer and improved food‐web productivity of subarctic Pacific waters. Size variation of coho salmon stocks south of Alaska was synchronous and negatively correlated with warm conditions (positive PDO) and weak North Pacific high pressure during ocean residence.     

The parasite Ichthyophonus sp. in Pacific herring from the coastal NE Pacific

The protistan parasite Ichthyophonus occurred in populations of Pacific herring Clupea pallasii Valenciennes throughout coastal areas of the NE Pacific, ranging from Puget Sound, WA north to the Gulf of Alaska, AK. Infection prevalence in local Pacific herring stocks varied seasonally and annually, and a general pattern of increasing prevalence with host size and/or age persisted throughout the NE Pacific. An exception to this zoographic pattern occurred among a group of juvenile, age 1+ year Pacific herring from Cordova Harbor, AK in June 2010, which demonstrated an unusually high infection prevalence of 35%. Reasons for this anomaly were hypothesized to involve anthropogenic influences that resulted in locally elevated infection pressures. Interannual declines in infection prevalence from some populations (e.g. Lower Cook Inlet, AK; from 20–32% in 2007 to 0–3% during 2009–13) or from the largest size cohorts of other populations (e.g. Sitka Sound, AK; from 62.5% in 2007 to 19.6% in 2013) were likely a reflection of selective mortality among the infected cohorts. All available information for Ichthyophonus in the NE Pacific, including broad geographic range, low host specificity and presence in archived Pacific herring tissue samples dating to the 1980s, indicate a long‐standing host–pathogen relationship.     

Competition between Asian pink salmon (Oncorhynchus gorbuscha) and Alaskan sockeye salmon (O. nerka) in the North Pacific Ocean

The importance of interspecific competition as a mechanism regulating population abundance in offshore marine communities is largely unknown. We evaluated offshore competition between Asian pink salmon and Bristol Bay (Alaska) sockeye salmon, which intermingle in the North Pacific Ocean and Bering Sea, using the unique biennial abundance cycle of Asian pink salmon from 1955 to 2000. Sockeye salmon growth during the second and third growing seasons at sea, as determined by scale measurements, declined significantly in odd‐numbered years, corresponding to years when Asian pink salmon are most abundant. Bristol Bay sockeye salmon do not interact with Asian pink salmon during their first summer and fall seasons and no difference in first year scale growth was detected. The interaction with odd‐year pink salmon led to significantly smaller size at age of adult sockeye salmon, especially among younger female salmon. Examination of sockeye salmon smolt to adult survival rates during 1977–97 indicated that smolts entering the ocean during even‐numbered years and interacting with abundant odd‐year pink salmon during the following year experienced 26% (age‐2 smolt) to 45% (age‐1 smolt) lower survival compared with smolts migrating during odd‐numbered years. Adult sockeye salmon returning to Bristol Bay from even‐year smolt migrations were 22% less abundant (reduced by 5.9 million fish per year) compared with returns from odd‐year migrations. The greatest reduction in adult returns occurred among adults spending 2 compared with 3 years at sea. Our new evidence for interspecific competition highlights the need for multispecies, international management of salmon production, including salmon released from hatcheries into the ocean.     

Climate effects and bottom‐up controls on growth and size‐at‐age of Pacific halibut (Hippoglossus stenolepis) in Alaska (USA)

Pacific halibut (Hippoglossus stenolepis) are an ecologically, commercially, and culturally important Alaskan groundfish species. Commercial harvest of halibut dates back to the late 19th century and has been managed by the International Pacific Halibut Commission (IPHC) since 1921. IPHC surveys have revealed declining trends in survey biomass in multiple regions and region‐specific declines in mean size‐at‐age (size‐at‐age) over the past two decades (>50% in some areas). Changes in size‐at‐age can arise from a variety of physical, ecological, sampling, and fishery effects, including size‐dependent fishery or predation mortality, alteration in growth from variability in prey quality or quantity, and changes in temperature‐dependent metabolic demands. Here, we develop and apply a bioenergetics model for halibut using survey‐based diet and temperature data for Alaska to evaluate potential environmental drivers of size‐at‐age. In general, juvenile (<40 cm fork length) foraging rates were highest in the Gulf of Alaska concomitant with higher potential growth and elevated basal metabolic demands during warm summer conditions. In contrast, adult (40–120 cm FL) potential growth was highest in the Eastern Bering Sea, potentially reflecting lower metabolic costs and higher rates of prey consumption in that region. We additionally find evidence for interannual variation in potential growth, with a higher frequency of reduced growth potential in the last decade, particularly in the Eastern Bering Sea in 2015 and 2016 for both juvenile and adult halibut. These results suggest the potential for patterns in size‐at‐age to arise from trophic and environmental constraints that collectively limit growth in some regions and years.     

Plankton dynamics: observed and modelled responses to physical conditions in Prince William Sound,Alaska

Plankton populations in Prince William Sound, Alaska, exhibited pronounced seasonal, annual and longer‐period variability in composition and standing stock in response to physically influenced differences in nutrient availability, and possibly currents that modify local biomass by exchanges with water from the bordering Gulf of Alaska. During springs in which early, strong physical stratification developed, intense, short‐lived phytoplankton blooms occurred. These blooms had relatively short residence times in the water column. In contrast, during springs in which slower, weaker stratification developed, phytoplankton blooms were prolonged and took longer to peak. These slower blooms prolonged the period of phytoplankton production, prolonged interaction with the springtime grazing community and led to the incorporation of more organic matter into pelagic food webs. A coupled biological‐physical simulation of plankton production was used to examine the implications of seasonally varying air and mixed‐layer temperatures, surface winds and incident light on the timing, duration, annual production and standing stock of plankton. Our modelling results reproduced the observed characteristics of the springtime production cycle, and the magnitude of zooplankton stocks for the period 1992–97 but not for 1981–91. These results suggest that for most of the 1990s, bottom‐up influences on nutrient supplies controlled levels of primary consumers, whereas for the 11 years before that, other unknown factors dominated this process. We present the results of a comprehensive, multiyear study of relationships between plankton and physical limitations, and a retrospective analysis of earlier conditions to explore the possible causes for these differences.     

Horizontal and vertical movements of juvenile bluefin tuna (Thunnus orientalis) in relation to seasons and oceanographic conditions in the eastern Pacific Ocean

Electronically tagged juvenile Pacific bluefin, Thunnus orientalis, were released off Baja California in the summer of 2002. Time‐series data were analyzed for 18 fish that provided a record of 380 ± 120 days (mean ± SD) of ambient water and peritoneal cavity temperatures at 120 s intervals. Geolocations of tagged fish were estimated based on light‐based longitude and sea surface temperature‐based latitude algorithms. The horizontal and vertical movement patterns of Pacific bluefin were examined in relation to oceanographic conditions and the occurrence of feeding events inferred from thermal fluctuations in the peritoneal cavity. In summer, fish were located primarily in the Southern California Bight and over the continental shelf of Baja California, where juvenile Pacific bluefin use the top of the water column, undertaking occasional, brief forays to depths below the thermocline. In autumn, bluefin migrated north to the waters off the Central California coast when thermal fronts form as the result of weakened equatorward wind stress. An examination of ambient and peritoneal temperatures revealed that bluefin tuna fed during this period along the frontal boundaries. In mid‐winter, the bluefin returned to the Southern California Bight possibly because of strong downwelling and depletion of prey species off the Central California waters. The elevation of the mean peritoneal cavity temperature above the mean ambient water temperature increased as ambient water temperature decreased. The ability of juvenile bluefin tuna to maintain a thermal excess of 10°C occurred at ambient temperatures of 11–14°C when the fish were off the Central California coast. This suggests that the bluefin maintain peritoneal temperature by increasing heat conservation and possibly by increasing internal heat production when in cooler waters. For all of the Pacific bluefin tuna, there was a significant correlation between their mean nighttime depth and the visible disk area of the moon.