Nonlocal wind-driven fjord–coast advection and its potential effect on plankton and fish recruitment

The Bergen Ocean Model (BOM), a three-dimensional physical coastal ocean model, was used for a numerical simulation experiment to investigate short-term effects of wind-generated coastal upwelling and downwelling on the dynamics of adjacent large outer and smaller inner fjords. The effect of the real alongshore wind regime on advection for an idealized fjord topography, resembling Masfjorden, western Norway, is used as an example. This modelling exercise is a supplement to, and its predictions support, the various hypotheses investigated in ecosystem simulation studies of the Masfjorden. The model predicts that coastal winds from the north cause upwelling and transport the upper water layer out from the fjords. Winds from the south cause downwelling and transport the upper water layer into the fjords. The transport is rapid and ≈50% of the upper water layer may be replaced within 1–2 days. Implications of these physical processes for the dispersal and retention of planktonic organisms and the early life stages of fish are discussed. If strong southerly winds occur frequently, this will transport planktonic organisms into the fjord and may increase the carrying capacity for planktivorous fish. In contrast, frequent strong northerly winds may reduce the abundance of planktonic organisms, including the early life stages of marine fish, and thus possibly reduce recruitment to fjord fish populations. Frequent shifts between southerly and northerly winds would cause an exchange of early life stages between neighbouring fjords and thus enhance genetic exchange.     

The influence of diel vertical migration on zooplankton transport and recruitment in an upwelling region: estimates from a coupled behavioral-physical model

Diel vertical migration (DVM) is a common zooplankton behavior in which organisms reside in surface or near‐surface waters at night and at deeper depths during the day. In many upwelling regions, DVM reduces the transport of organisms away from the region. It is unclear, however, what role DVM plays in recruitment (the arrival of larvae or juveniles to locations where they will become reproducing adults) to upwelling regions. In this study, we estimate the influence of DVM on zooplankton transport, the level of recruitment of locally produced propagules (self‐recruitment), and sources of recruits in the upwelling region near Monterey Bay, California, by simulating the trajectories of fixed‐depth and vertically migrating organisms with a drifter‐tracking algorithm driven by climatological velocity fields from a three‐dimensional hydrodynamic model. Our simulations suggest that DVM into subsurface poleward and onshore currents during the day does not fully compensate for equatorward and offshore transport in the surface Ekman layer at night and does not retain zooplankton in the Monterey Bay region. Our simulations also suggest that DVM decreases the ability of zooplankton to return to the region after being transported away and shifts source regions for recruits closer to the bay. While DVM does not appear to substantially increase the potential for self‐recruitment to the region, this study indicates that other mechanisms, such as transport during non‐upwelling periods, continuous transport below the surface, increases in mean transport depth over time, or seasonal changes in hydrography, may still enable relatively high levels of self‐recruitment to this highly advective region.     

Fish larvae distribution off Mejillones Peninsula (northern Chile) during a coastal upwelling event in Spring 1999: interactions with the cold upwelling plume

We examined the interaction between vertical and horizontal distribution of fish larvae off Mejillones Peninsula (23°S), northern Chile, under conditions of active coastal upwelling. An oceanographic survey covered spatial variability in temperature, chlorophyll- a (chl a ), dissolved oxygen, salinity and water density. Fish larvae were sampled during daytime and nighttime periods through two consecutive days in four stations: two inside and two outside of a well-developed upwelling plume, and at three depth strata: 0–20, 20–80 and 80–200 m. Eighteen taxa were analysed, of which the Myctophidae Diogenychthys atlanticus , Diogenichthys laternatus , and the anchovy Engraulis ringens , were most abundant. Our data showed little evidence for diel vertical migration and larvae were more abundant at depth (>80 m) under low temperature (∼12°C) and low chl a (∼2 mg m^–3), below the highly advective upper layer. The exploratory K -means analysis allowed the separation of data into two distinct habitats: upwelling and nonupwelling types. Most taxa were allocated in nonupwelling waters, i.e. outside the cold plume. However, short-term variations (<24 h) in the position of the upwelling plume influenced both horizontal and vertical occurrence as well as abundance of taxa, and caused variability in temperature, oxygen and chl a . These changes in oceanographic conditions, caused by upwelling circulation and the dynamics of the cold plume, may sharply modify the habitat of fish larvae and have an important role in survivorship and recruitment success.     

Spatial variability in red sea urchin (Strongylocentrotus franciscanus) recruitment in northern California

To better understand the spatial distribution of recruitment in the northern California red sea urchin ( Strongylocentrotus franciscanus ) population, we sampled size distributions at each of 12 locations in 1995 and 1996, two of those locations in 1994, and 5 of those locations in 1997. An index of recent recruitment in these size distributions and an estimate of density of recent recruitment reflect a similar spatial pattern of recruitment. This pattern appears to be determined by the effect of coastal circulation on larval delivery during relaxation of upwelling, and not the result of the positive effect of the adult spine canopy on juvenile survival. Recent recruitment of red sea urchins in northern California is higher in areas more frequently subjected to onshore and poleward flow during relaxation of upwelling. These results are consistent with a mechanism by which alongshore spatial variability in southward, offshore flow during upwelling winds allows larvae to maintain latitudinal position, whereas flows during event-scale relaxations in upwelling winds serve to distribute settling larvae alongshore, favouring areas north of promontories. The consequent spatial pattern of red sea urchin settlement varies from year to year, and there are not yet sufficient data to demonstrate the degree to which this spatial pattern in recruitment determines a spatial pattern in fishable adult abundance.     

Simulation and quantification of enrichment and retention processes in the southern Benguela upwelling ecosystem

Important environmental processes for the survival and recruitment of early life stages of pelagic fishes have been synthesized through Bakun's fundamental triad as enrichment, concentration and retention processes (A. Bakun, 1996, Patterns in the Ocean. Ocean Processes and Marine Population Dynamics. San Diego, CA, USA: University of California Sea Grant). This conceptual framework states that from favourable spawning habitats, eggs and larvae would be transported to and/or retained in places where food originating from enrichment areas would be concentrated. We propose a method for quantifying two of the triad processes, enrichment and retention, based on the Lagrangian tracking of particles transported within water velocity fields generated by a three‐dimensional hydrodynamic model. We apply this method to the southern Benguela upwelling ecosystem, constructing putative maps of enrichment and retention. We comment on these maps regarding main features of the circulation in the region, and investigate seasonal variability of the processes. We finally discuss the results in relation to available knowledge on the reproductive strategies of two pelagic clupeoid species abundant in the southern Benguela, anchovy (Engraulis encrasicolus) and sardine (Sardinops sagax). Our approach is intended to be sufficiently generic so as to allow its application to other upwelling systems.     

Method for estimating buoyancy of midwater float required to standardize hook depth in pelagic longline

ABSTRACT:   A midwater float is a small float attached to the mainline of pelagic longlines to standardize the hook depth. In this study theoretical equations are presented for estimating the buoyancy of the midwater float required to lift the joints of the midwater float line on the mainline to the target depth. Sea trials using full scale tuna longline gear with midwater floats were carried out in the Indian Ocean in December 2004 and 2005, in order to examine the validity of the theoretical equations. In the sea trials, two types of midwater float settings, single midwater float setting and double midwater float setting, were tested and compared with the conventional setting. As a result, the joints of the midwater float line on the mainline were successfully lifted to the target depth as expected, demonstrating the validity of the theoretical equation. The range of hook depths in the midwater float setting was less spread over depths than in the conventional setting, and therefore, use of long float lines (100 m) with the midwater float setting allows all hooks to avoid entering the sea turtle habitat of shallower than 100 m depth. Factors affecting shoaling of the longline with the midwater float are also discussed.     

The importance of retention processes in upwelling areas for recruitment of Octopus vulgaris: the example of the Arguin Bank (Mauritania)

The relationship between recruitment of octopus ( Octopus vulgaris ) populations and environmental conditions off the Arguin Bank (Mauritania) in the main nursery ground was investigated between 1990 and 1996. Three environmental indices (coastal upwelling intensity, coastal retention and wind-induced turbulence indices) are derived from satellite infrared imagery (METEOSAT) and from the COADS (Comprehensive Ocean–Atmosphere Data Set) data to quantify the coastal retention process and productivity, both believed to be important factors for recruitment success. The octopus recruitment index as well as environmental indices fluctuate annually and seasonally. Linear regression and generalized additive models (GAM) are used to relate recruitment indices and the environmental conditions prevailing during the early-life planktonic stage, known to be the main critical period of the octopus life cycle. Coastal retention especially appears to be a key factor for recruitment success. Owing to seasonal variations in enrichment and mixing processes, larvae benefit from retention in the spring but are negatively affected by a breakdown in retention in the autumn. Increasing upwelling intensity is beneficial for octopus recruitment, as the Arguin Bank limits the detrimental dispersive effects linked to upwelling.     

Interannual variation in the coastal distribution of a juvenile gadid in the northeast Pacific Ocean: The relevance of wind and effect on recruitment

Drift of propagules occurs within many populations inhabiting flow fields. This affects the number of propagules that rejoin their source population (recruitment) and plays a role in adaptive spatial redistribution. We focus on the cause and consequence of interannual variation in geographic distribution of population density among five cohorts of young‐of‐the‐year (age‐0) juvenile walleye pollock Gadus chalcogrammus in the western Gulf of Alaska (GOA). The coastal GOA is a wind‐driven advective system. Walleye pollock spawn during spring and their eggs and larvae drift southwestward; by late summer, age‐0 juveniles are variously distributed over the shelf. We found that high population densities of age‐0 juveniles (ca. 6 months old) near the southwestward exit of the Alaska Coastal Current from the GOA corresponded with high abundance of larvae from the major spawning area upstream, but did not translate into high abundance at older ages. Further, offshore and upwelling‐favorable winds were associated with the high downstream abundance and presumed export. In contrast, downwelling‐favorable (northeasterly) wind during and shortly after spawning (April–May) was associated with high recruitment at age 1. Finally, we found that recruitment also increased with apparent retention of age‐0 juveniles in favorable habitat upstream near the main spawning area. We hypothesize that wind‐related retention in superior upstream habitat favors recruitment. Our results argue for including wind‐driven transport in future walleye pollock recruitment models. We encourage more work on the juvenile stage of marine fishes aimed at understanding how transport and species‐specific habitat suitability interact to affect population response to large‐scale forcing.     

Variation in the distribution of ocean shrimp (Pandalus jordani) recruits: links with coastal upwelling and climate change

In this analysis, an atypical northward shift in the distribution of age‐1 ocean shrimp (Pandalus jordani) recruits off Oregon in 2000 and 2002–2004 was linked to anomolously strong coastal upwelling winds off southern Oregon (42°N latitude) in April–July of the year of larval release (t?1). This is the first clear evidence that strong upwelling winds can depress local recruitment of ocean shrimp. Regression analysis confirmed a long‐term negative correlation between log_e of ocean shrimp recruitment and April sea level height (SLH) at Crescent City, California, in the year of larval release, for both northern and southern Oregon waters. The regional pattern of ocean shrimp catches and seasonal upwelling winds showed that, although the timing of the spring transition as reflected in April SLH drives ocean shrimp recruitment success off Oregon generally, the strength and consistency of spring upwelling limits the distribution of large concentrations of ocean shrimp at the southern end of the northern California/Oregon/Washington area. A northward shift in 1999 and 2001–03 in the northern edge of this ‘zone of maximum upwelling’ is the likely cause of the weak southern Oregon recruitment and resulting atypical distribution of ocean shrimp observed off Oregon in 2000 and 2002–04, with a return to a more typical catch distribution as spring upwelling moderated in subsequent years. It is noted that a northward shift in the conditions that produce strong and steady spring upwelling winds is consistent with many predictions of global climate models under conditions of global warming.     

Spatial correlation patterns in coastal environmental variables and survival rates of salmon in the north-east Pacific Ocean

We examined spatial correlations for three coastal variables [upwelling index, sea surface temperature (SST), and sea surface salinity (SSS)] that might affect juvenile salmon ( Oncorhynchus spp.) during their early marine life. Observed correlation patterns in environmental variables were compared with those in survival rates of pink ( O. gorbuscha ), chum ( O. keta ), and sockeye ( O. nerka ) salmon stocks to help identify appropriate variables to include in models of salmon productivity. Both the upwelling index and coastal SST were characterized by strong positive correlations at short distances, which declined slowly with distance in the winter months, but much more rapidly in the summer. The SSS had much weaker and more variable correlations at all distances throughout the year. The distance at which stations were no longer correlated (spatial decorrelation scale) was largest for the upwelling index (> 1000 km), intermediate for SST (400–800 km in summer), and shortest for SSS (< 400 km). Survival rate indices of salmon showed moderate positive correlations among adjacent stocks that decreased to zero at larger distances. Spatial decorrelation scales ranged from approximately 500 km for sockeye salmon to approximately 1000 km for chum salmon. We conclude that variability in the coastal marine environment during summer, as well as variability in salmon survival rates, are dominated by regional scale variability of several hundred to 1000 km. The correlation scale for SST in the summer most closely matched the observed correlation scales for survival rates of salmon, suggesting that regional-scale variations in coastal SST can help explain the observed regional-scale covariation in survival rates among salmon stocks.     

Global climate change and ocean upwelling

The influence of global greenhouse warming on the ocean's biological productivity may be more complicated and weaker than that proposed by Bakun (1990). A doubled carbon dioxide simulation made with the Canadian Climate Centre atmospheric general circulation model coupled to a simplified mixed layer ocean model suggests that the midlatitude continents do not all follow the Bakun scenario in developing anomalous low pressure in summer and enhancing coastal winds favorable to upwelling. In the open ocean the equatorial and subpolar zonal upwelling bands and the subtropical downwelling bands generally weaken as winds diminish owing to the weakening of the equator-to-pole temperature gradient in the lower troposphere under global warming. With a weakening of open ocean upwelling and an absence of enhanced coastal upwelling, the overall effect of global warming could be to decrease the global biological productivity.     

Fine-scale movement patterns, site fidelity, and habitat selection of ocean whitefish (Caulolatilus princeps)

The fishery for California groundfishes is managed using broad species complexes, although some non-groundfish species are managed similarly due to the perception of shared behavioral characteristics. This study integrates acoustic telemetry and a GIS to quantify movement patterns of one such species, the ocean whitefish (Caulolatilus princeps) in a marine protected area. Seventeen ocean whitefish were tagged and actively tracked over multiple 24-h periods to measure fine-scale movement patterns. Home ranges based on 95% kernel utilization distributions averaged 20,439 ± 28,492 (±S.D.) m². Fish were active during the day, foraging over sand habitat at depths averaging 21 ± 8 m, but were inactive at night, taking refuge near rocky reefs at depths averaging 15 ± 7 m. Seventeen additional fish were tagged with coded acoustic transmitters and passively tracked using automated underwater acoustic receivers for up to 1 year. Approximately 75% of these fish exhibited long-term (1 year) fidelity to home ranges in the study area. Results suggest that MPAs can be an effective means of protecting populations of ocean whitefish and based on their habitat associations, ocean whitefish can be managed separately from other reef associated groundfishes.     

Influence of episodic upwelling on capelin, Mallotusvillosus, and Atlantic cod, Gadusmorhua, catches in Newfoundland coastal waters

Wind-induced upwelling in the north-west Atlantic has been hypothesized to influence catch rates of fish by gear fishing at fixed locations. Passive movement to shallow depths during upwelling has been proposed to increase encounter rates of fish with net leaders set at the coast. However, supporting evidence is not conclusive, possibly because fish respond to strong events rather than all events. We investigated whether catch rates of capelin, Mallotusvillosus, and Atlantic cod, Gadusmorhua, were related to the strength of upwelling, as measured by the rates of water temperature change, or to stage of upwelling. Capelin trap catches were positively related to increases in water temperature, representing the relaxation phase. This result was due primarily to increases in catch after strong (> 4°C change) rather than after typical (< 4°C change) upwelling events. Cod trap catches were not related to upwelling strength but did increase 1–2 days after typical events. The data suggest that upwelling increases capelin movement, while the return of warmer surface water after an upwelling event increases cod movement.     

Evaluating upwelling estimates off the west coasts of North and South America

Both low‐ and high‐resolution studies of wind and upwelling were conducted off the west coasts of North and South America between August 1999 and December 2001. For the low‐resolution study, done at 25 regional sites spanning both coasts, weekly mean winds were calculated from satellite measurements, from geostrophic estimates, and from an operational 100‐km‐resolution global atmospheric model. The satellite‐measured winds, used as the reference, showed that the coastal regions of both North and South America were divided into fairly uniform climatic domains located at higher, mid, and lower latitudes. All three weekly‐mean wind data sets were compared at each of the 25 sites, and then used to estimate upwelling based on the method employed by NOAA's Pacific Fisheries Environmental Laboratory. Within each of the wind domains, model‐derived wind and upwelling estimates agreed with satellite‐derived values better than those from geostrophic‐derived estimates. To investigate variability between the regional sites, a 9‐km‐resolution atmospheric model was run for an area off California which spanned four of the regional sites. This high‐resolution model, verified with satellite measurements, revealed jets of wind curving and intensifying around coastal promontories. These near‐shore wind jets, undetected in the global model, resulted in strong upwelling bands on the order of 50 km alongshore by 20 km offshore. These bands are critical for calculating local upwelling. Our results indicate that regional upwelling estimates for fisheries research can be improved by replacing geostrophic estimates of winds with those from a global atmospheric model. For localized coastal upwelling estimates, however, models with resolution an order of magnitude higher are required.     

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.     

Swimming depths of the giant jellyfish <Emphasis Type="Italic">Nemopilema nomurai</Emphasis> investigated using pop-up archival transmitting tags and ultrasonic pingers

The swimming depths of 12 individual Nemopilema nomurai with bell diameters of 0.8–1.6 m were investigated using pop-up archival transmitting tags and ultrasonic pingers, and the validity of the research method was evaluated. The N. nomurai studied frequently showed vertical movement, with the swimming depth ranging from 0 to 176 m, The mean swimming depths of most individuals were less than 40 m. The swimming depths of N. nomurai in the northern Japan Sea in the winter were mostly deeper than those of this species in the southern Japan Sea in the autumn. This result suggests that the range of the depths almost depends on the vertical structure of the ocean. Swimming depths during the nighttime were significantly deeper than those during the daytime. More specifically, during the daytime, the swimming depths in the afternoon tended to be shallower than those in the morning, while during the nighttime, the swimming depths after midnight were deeper than those before midnight.     

Environmental conditions, satellite imagery, and clupeoid recruitment in the northern Benguela upwelling system

The relationship between oceanographic conditions and clupeoid (pilchard, Sardinops sagax, and anchovy, Engraulis capensis ) recruitment in the northern Benguela upwelling system was investigated from 1981 to 1987 using a time-series of mean weekly SST images. Two approaches were taken. The first involved correlating recruitment success with the number of weekly coastal `SST events' above various cut-off temperatures during the main reproductive season. The second involved constructing a multiple regression model of recruitment success with two independent environmental variables: namely, the number of coastal `SST events' greater than 19°C, and an onshore retention index for the early life-history stages. The retention index was derived from a spatial time-series analysis of the SST images using principal components analysis. In general, pilchard recruitment showed a positive relationship with the `number of SST events' whilst anchovy recruitment had a negative relationship; 1987 was an outlier year, during which there were exceptionally high levels of both pilchard and anchovy recruitment. The multiple regression R ² values were high and significant for both species (pilchard R ² = 0.88, anchovy R ² = 0.96). The regression model also accounted for the 1987 outlier according to levels of onshore retention which, despite low inshore SSTs, were particularly high during the 1986/87 reproductive season. Although these results need to be validated with data from a longer time period, they show how satellite data might be used for predicting clupeoid recruitment success in the northern Benguela.     

Population dynamics of Calanus chilensis in the Chilean Eastern Boundary Humboldt Current

A study of the population dynamics of Calanus chilensis was carried out in the coastal area of the Mejillones Peninsula (23°S), which is an upwelling site in the Eastern Boundary Humboldt Current. Information was derived from time-series sampling (15 day interval) at three stations, and from two broad-scale cruises: under active upwelling, and under 1997/1998 El Niño conditions. Calanus chilensis showed continuous reproduction throughout the year. In the study area, upwelling takes place intermittently year-round, suggesting that Calanus production is not limited by food. Advection played an important role in structuring the spatial distribution of Calanus , both during upwelling and under El Niño conditions. Population losses from upwelling and offshore transport may be compensated by rapid turnover rates of cohorts, growing at temperature-dependent rates and hence allowing population recovery, even in abnormally warm waters during El Niño conditions.     

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.     

Comparison of oxygen consumption patterns between wild and cultured black rockfish Sebastes schlegeli

ABSTRACT:   To compare metabolic activity rhythms between wild and cultured black rockfish Sebastes schlegeli , we measured long-term oxygen consumption rates (OCR) using an automatic intermittent-flow respirometer under constant temperature and darkness. Oxygen consumption rates peaked among wild black rockfish at 12.4 h intervals, which corresponded to a circatidal rhythm. The wild fish were probably exhibiting responses that corresponded to tidal events in their natural environment. However, when captured wild black rockfish were kept under laboratory conditions (12 h light [L] : 12 h dark [D]) for 30 days, the OCR shifted to a circadian rhythm (24.1–24.9 h). The OCR of cultured black rockfish that had been reared in a tank for 9 months peaked at approximately 24 h intervals, corresponding to a circadian rhythm. The results of this study suggest that the differences in OCR patterns between wild and cultured fish were mainly due to differences in the environmental conditions between tidal and non-tidal habitats.