Life stage‐specific,stochastic environmental effects overlay density dependence in an Atlantic salmon population

Atlantic salmon populations appear to fluctuate stochastically through time. It is suspected that both density‐dependent and density‐independent factors cause these fluctuations but the relative importance of each, and the life stages at which they operate, is not well known. In this study, a long‐term data set on Atlantic salmon migrants returning to the Foyle catchment, Ireland, was used to determine the role of density‐dependent and life stage‐specific environmental factors regulating population size. A Ricker density‐dependent model showed that spawning adult population size significantly predicted variation in the resultant filial generation; however, a large amount of variation (ca. 68%) remained unexplained. It was shown that environmental factors were significant in explaining some of the remaining variance and that these influences were linked to specific life stages. Three life stages—spawning and incubation, fry emergence and marine survival—were shown to have significant environmental effects that resulted in changes in the returning cohort strength. It is concluded that these life stage‐specific environmental effects are likely to contribute to the stochastic variation in population size resulting from the application of traditional stock–recruitment models. The identification and quantification of these effects should allow improved model accuracy.     

Environmentally driven forecasts of northern rock sole (Lepidopsetta polyxystra) recruitment in the eastern Bering Sea

Northern rock sole recruitment in the eastern Bering Sea has been hypothesized to (a) depend on wind‐driven surface currents linking spawning and nursery areas, (b) be density‐dependent, and (c) be negatively impacted by cold bottom temperatures over a large nursery area during the first summer of life. A suite of models was developed to test these hypotheses. Data included 32 years of recruitment and spawning biomass estimates derived from a stock assessment model and wind and temperature indices customized to the environmental exposure of age‐0 northern rock sole in the eastern Bering Sea. The predictive ability of the models was evaluated, and the models were used to forecast recruitment to age‐4 for recent year classes which are poorly retained by the standard multi‐species bottom trawl survey gear. Models which included wind and temperature indices performed better than a naïve forecast based on the running mean. The best‐performing model was a categorical model with wind and temperature thresholds, which explained 49% of the variation in recruitment. Ricker models performed more poorly than models without a spawning biomass term, providing no evidence that recruitment is related to stock size. The models forecast higher recruitment for the most recent year classes (2015–2018) than for prior year classes with observed poor recruitment (2006–2013). These environment‐based recruitment forecasts may improve recruitment estimates for the most recent year classes and facilitate study of the effects of future climate change on northern rock sole population dynamics.     

Interannual differences in growth and hatch‐date distributions of early juvenile European anchovy in the Bay of Biscay: implications for recruitment

In order to understand better the recruitment variability in European anchovy in the Bay of Biscay, it is important to investigate the processes that affect survival during the early life stages. Anchovy juvenile growth trajectories and hatch‐date distributions were inferred over a 3‐year period based on otolith microstructure analysis. Otolith growth trajectories showed a characteristic shape depending on their hatch‐date timing. Earlier‐born juveniles had notably broader maximum increments than later born conspecifics, resulting in higher growth rates. This observation suggests that early hatching would be beneficial for larval and juvenile growth, and, therefore, survival. The estimated juvenile hatch‐date distributions were relatively narrow compared with the extended anchovy spawning season (March–August) in the Bay of Biscay and indicated that only individuals originated mainly from the summer months (June–August) survived until autumn. Hatch‐date distributions were markedly different among years and seemed to influence the interannual recruitment strength of anchovy. We conclude that years characterized by juvenile survivors originating from the peak spawning period (May and June) would lead to considerable recruitment success. Downwelling events during the peak spawning period seem to affect larval survival. Furthermore, size‐dependent overwinter mortality would be an additional process that regulates recruitment strength in the anchovy population in the Bay of Biscay.     

Density‐ and size‐dependent mortality in fish early life stages

The importance of survival and growth variations early in life for population dynamics depends on the degrees of compensatory density dependence and size dependence in survival at later life stages. Quantifying density‐ and size‐dependent mortality at different juvenile stages is therefore important to understand and potentially predict the recruitment to the population. We applied a statistical state‐space modelling approach to analyse time series of abundance and mean body size of larval and juvenile fish. The focus was to identify the importance of abundance and body size for growth and survival through successive larval and juvenile age intervals, and to quantify how the dynamics propagate through the early life to influence recruitment. We thus identified both relevant ages and mechanisms (i.e. density dependence and size dependence in survival and growth) linking recruitment variability to early life dynamics. The analysis was conducted on six economically and ecologically important fish populations from cold temperate and sub‐arctic marine ecosystems. Our results underscore the importance of size for survival early in life. The comparative analysis suggests that size‐dependent mortality and density‐dependent growth frequently occur at a transition from pelagic to demersal habitats, which may be linked to competition for suitable habitat. The generality of this hypothesis warrants testing in future research.     

Environmental factors influencing larval sprat Sprattus sprattus feeding during spawning time in the Baltic Sea

The management of Baltic sprat is challenged by highly variable recruitment success and hence large stock fluctuations. Recent studies have identified the larval and early juvenile life stages to be critical for the survival rate of a sprat year class. Although prey abundance was found to be linked to larval survival success, an analysis identifying the functional relationship and relative importance of other environmental factors is still missing. Sprat larval feeding was investigated in 2002 during three cruises, covering the main spawning time in the Bornholm Basin, Baltic Sea. The aim of the study was to identify the key environmental factors determining the feeding success of larval sprat taking their potential interactions explicitly into account. An extension of generalized additive models (GAMs) was adopted that allows the inclusion of interaction terms in a non‐parametric regression model. The final model of sprat larval feeding success explained ～80% of the variance in the data and was based on the following environmental factors: bottom depth, cubed wind speed as proxy for small‐scale turbulence rates, degree of cloudiness as proxy for light conditions and prey density in combination with a feeding period–cloudiness interaction term. Our study demonstrates that the feeding success of sprat larvae in the Baltic Sea is controlled by a number of simultaneously acting key environmental factors.     

Recruitment of the threatened mussel Margaritifera margaritifera in relation to mussel population size,mussel density and host density

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Riparian restoration offsets predicted population consequences of climate warming in a threatened headwater fish

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Factors influencing recruitment of walleye and white bass to three distinct early ontogenetic stages

Determining the factors that influence recruitment to sequential ontogenetic stages is critical for understanding recruitment dynamics of fish and for effective management of sportfish, particularly in dynamic and unpredictable environments. We sampled walleye (Sander vitreus) and white bass (Morone chrysops) at 3 ontogenetic stages (age 0 during spring: ‘age‐0 larval’; age 0 during autumn: ‘age‐0 juvenile’; and age 1 during autumn: ‘age‐1 juvenile’) from 3 reservoirs. We developed multiple linear regression models to describe factors influencing age‐0 larval, age‐0 juvenile and age‐1 juvenile walleye and white bass abundance indices. Our models explained 40–80% (68 ± 9%; mean ± SE) and 71%–97% (81 ± 6%) of the variability in catch for walleye and white bass respectively. For walleye, gizzard shad were present in the candidate model sets for all three ontogenetic stages we assessed. For white bass, there was no unifying variable in all three stage‐specific candidate model sets, although walleye abundance was present in two of the three white bass candidate model sets. We were able to determine several factors affecting walleye and white bass year‐class strength at multiple ontogenetic stages; comprehensive analyses of factors influencing recruitment to multiple early ontogenetic stages are seemingly rare in the literature. Our models demonstrate the interdependency among early ontogenetic stages and the complexities involved with sportfish recruitment.     

A study on relationships between large‐scale climate indices and estimates of North Pacific albacore tuna productivity

A logistic production model was used to examine potential relationships between three climate indices, the North Pacific Gyre Oscillation (NPGO), the Pacific Decadal Oscillation (PDO), and the Multivariate El Niño‐Southern Oscillation Index (MEI), and productivity estimates of the North Pacific albacore tuna (Thunnus alalunga) population. Catch and standardized catch‐per‐unit‐effort data from three longline fisheries (Japan, US, and Taiwan) were used in the model. The climate indices were incorporated into the model by correlating time‐varying intrinsic population growth rate (r_y) of the production model with the annual mean value for each index. The estimated probability that the NPGO is positively correlated with stock productivity, as measured by r_y, was 0.99, and the calculated probability that MEI is negatively correlated with the productivity was 0.95. The time lag for these correlations is 4 yr, which is consistent with the timing of recruitment to the Japan longline fishery. The PDO did not seem to have any detectable relationship with stock productivity. However, it remains uncertain if there is a conclusive linkage between the albacore productivity and the NPGO or the MEI index, because model fit to the data is about the same as that of a base model which does not use any climate index and assumes a time‐invariant r.     

Marine environment‐based forecasting of coho salmon (Oncorhynchus kisutch) adult recruitment

Generalized additive models (GAMs) were used to investigate the relationships between annual recruitment of natural coho salmon (Oncorhynchus kisutch) from Oregon coastal rivers and indices of the physical ocean environment. Nine indices were examined, ranging from large‐scale ocean indicators, e.g., Pacific Decadal Oscillation (PDO), to indicators of the local ecosystem (e.g., coastal water temperature near Charleston, OR). Generalized additive models with two and three predictor variables were evaluated using a set of performance metrics aimed at quantifying model skill in short‐term (approximately 1 yr) forecasting. High explanatory power and promising forecast skill resulted when the spring/summer PDO averaged over the 4 yr prior to the return year was used to explain a low‐frequency (multi‐year) pattern in recruitment and one or two additional variables accounted for year‐to‐year deviations from the low‐frequency pattern. More variance was explained when averaging the predictions from a set of models (i.e., taking the ensemble mean) than by any single model. Making multiple forecasts from a set of models also provided a range of possible outcomes that reflected, to some degree, the uncertainty in our understanding of how salmon productivity is driven by physical ocean conditions.     

Diverse juvenile life‐history behaviours contribute to the spawning stock of an anadromous fish population

Habitat quality often varies substantially across space and time, producing a shifting mosaic of growth and mortality trade‐offs across watersheds. Traditional studies of juvenile habitat use have emphasised the evolution of single optimal strategies that maximise recruitment to adulthood and eventual fitness. However, linking the distribution of individual behaviours that contribute to recruitment at the population level has been elusive, particularly for highly fecund aquatic organisms. We examined juvenile habitat use within a population of sockeye salmon (Oncorhynchus nerka) that spawn in a watershed consisting of two interconnected lakes and a marine lagoon. Otolith microchemical analysis revealed that the productive headwater lake accounted for about half of juvenile growth for those individuals surviving to spawn in a single river in the upper watershed. However, 47% of adults had achieved more than half of their juvenile growth in the downstream less productive lake, and 3% of individuals migrated to the estuarine environment during their first summer and returned to freshwater to overwinter before migrating back to sea. These results describe a diversity of viable habitat‐use strategies by juvenile sockeye salmon that may buffer the population against poor conditions in any single rearing environment, reduce density‐dependent mortality and have implications for the designation of critical habitat for conservation purposes. A network of accessible alternative habitats providing trade‐offs in growth and survival may be important for long‐term viability of populations.     

Roles of discharge and temperature in recruitment of a cold‐water fish,the European grayling Thymallus thymallus,near its southern range limit

Recruitment of salmonids is a result of density‐dependent factors, specifically egg production in the previous year, and density‐independent environmental processes driven by discharge and temperature. With the plethora of knowledge on major drivers of Atlantic salmon Salmo salar and brown trout Salmo trutta recruitment, there is a requirement to explore less known species, such as European grayling Thymallus thymallus, whose postemergence time coincides with period of increasing temperature and low discharge. This study assessed drivers of grayling recruitment in a southern English chalk stream, a system vulnerable to discharge and temperature alterations under future climate change predictions. The analyses explored age 0+ grayling survival in relation to conspecific and heterospecific densities and discharge‐ and temperature‐derived factors. The final mixed‐effects model revealed a positive relationship between age 0+ grayling survival and incubation temperature anomaly and age 0+ trout abundance. Similarly, postincubation temperature anomaly had a positive effect on 0+ grayling survival, but only up to a threshold temperature of 13.5°C, beyond which it had a negative effect. In contrast, increasing number of days with low discharge postincubation negatively influenced age 0+ grayling survival, with no evidence of an effect of elevated discharges following spawning. Our results emphasise the importance of maintaining natural discharge regimes in salmonid rivers by tackling multiple stressors operating at the catchment scale, including land and water use to mitigate for predicted climate driven changes. In addition, further research on recruitment drivers in less stable, rain‐fed systems, is required.     

Abiotic and biotic correlates of yellow perch recruitment to age‐2 in southern Lake Michigan, 1984–2007

Abstract Seven abiotic and biotic variables were tested to determine whether they influence the recruitment to age two of yellow perch, Perca flavescens (Mitchill), in waters of southern Lake Michigan, USA. Recruitment was analysed using a Ricker stock‐recruitment relationship that accounts for density dependency. Significant model variables tested individually and ranked by total variance explained included the abundance of sexually mature yellow perch, alewife, Alosa pseudoharengus (Wilson) and spottail shiner, Notropis hudsonius (Clinton). Variables found unrelated to recruitment or generally less explanatory included the abundance of round goby, Neogobius melanostomus (Pallus), mean water temperature, variation in water temperature and water clarity. The best‐fitting multivariable model explained 69% of the recruitment variability and included abundances of sexually mature yellow perch, alewife and spottail shiner. These results suggest that yellow perch recruitment in southern Lake Michigan is regulated in part by biotic interactions with other species of the near‐shore community, including alewife and spottail shiners, in addition to the abundance of reproductively mature yellow perch.     

Spatial variability in ocean‐mediated growth potential is linked to Chinook salmon survival

Early ocean survival of Chinook salmon, Oncorhynchus tshawytscha, varies greatly inter‐annually and may be the period during which later spawning abundance and fishery recruitment are set. Therefore, identifying environmental drivers related to early survival may inform better models for management and sustainability of salmon in a variable environment. With this in mind, our main objectives were to (a) identify regions of high temporal variability in growth potential over a 23‐year time series, (b) determine whether the spatial distribution of growth potential was correlated with observed oceanographic conditions, and (c) determine whether these spatial patterns in growth potential could be used to estimate juvenile salmon survival. We applied this method to the fall run of the Central Valley Chinook salmon population, focusing on the spring and summer period after emigration into central California coastal waters. For the period from 1988 to 2010, juvenile salmon growth potential on the central California continental shelf was described by three spatial patterns. These three patterns were most correlated with upwelling, detrended sea level anomalies, and the strength of onshore/offshore currents, respectively. Using the annual strength of these three patterns, as well as the overall growth potential throughout central California coastal waters, in a generalized linear model we explained 82% of the variation in juvenile salmon survival estimates. We attributed the relationship between growth potential and survival to variability in environmental conditions experienced by juvenile salmon during their first year at sea, as well as potential shifts in predation pressure following out‐migration into coastal waters.     

Factors affecting the population viability of the burbot,Lota lota

Abstract Population viability analysis (PVA) is a frequently used conservation tool for the assessment of long‐term survival of populations. Based on demographic information from published literature, a PVA model using VORTEX was constructed for the burbot Lota lota (L.), a freshwater fish species believed extinct in the United Kingdom since the 1970s. The model was constructed to evaluate the persistence of a single burbot population over a 100‐year period as part of an assessment of the feasibility of reintroducing the species to rivers of its former distribution in England. Population persistence was highly variable and dependent on juvenile survival and spawning regularity. Under high juvenile survival scenarios, all populations persisted and were maintained at the carrying capacity. At low juvenile survival, population persistence was highly influenced by spawning regularity. Sensitivity analysis indicated that females must successfully spawn approximately every second year to guarantee 95% population survival. The analysis suggests that the long‐term viability of any potential reintroduced burbot population will be dependent on the environmental conditions related to spawning success.     

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.     

海冰与南半球环状模对南极布兰斯菲尔德海峡南极磷虾种群结构的影响

南极磷虾(Euphausia superba)是南大洋生态系统的关键物种,其生活史各阶段的生理过程与海冰密切相关。本研究基于开放获取数据库计算了布兰斯菲尔德海峡内反映南极磷虾种群结构特征的5个参数,包括补充指数和幼体、雌性、雄性平均体长以及雌性个体占比,并结合布兰斯菲尔德海峡及周围3个水域的海冰面积、水深、海底复杂度以及南半球环状模(SAM)逐日指数构建广义加性模型,探讨了海冰与气候事件对磷虾种群动态的影响。结果表明,磷虾幼体、雄性磷虾平均体长与60日前的威德尔海西北部海域海冰面积呈显著的负相关,磷虾补充指数与当日和60日前的SAM呈显著的正相关。随着海底复杂性的增加,磷虾补充指数显著下降。4个海域的海冰面积变化对磷虾补充指数和其他种群特征指标的影响存在差异。     

Oceanographic drivers of sablefish recruitment in the California Current

Oceanographic processes and ecological interactions can strongly influence recruitment success in marine fishes. Here, we develop an environmental index of sablefish recruitment with the goal of elucidating recruitment‐environment relationships and informing stock assessment. We start with a conceptual life‐history model for sablefish Anoplopoma fimbria on the US west coast to generate stage‐ and spatio‐temporally‐specific hypotheses regarding the oceanographic and biological variables likely influencing sablefish recruitment. Our model includes seven stages from pre‐spawn female condition through benthic recruitment (age‐0 fish) for the northern portion of the west coast U.S. sablefish stock (40°N–50°N). We then fit linear models and use model comparison to select predictors. We use residuals from the stock‐recruitment relationship in the 2015 sablefish assessment as the dependent variable (thus removing the effect of spawning stock biomass). Predictor variables were drawn primarily from ROMS model outputs for the California Current Ecosystem. We also include indices of prey and predator abundance and freshwater input. Five variables explained 57% of the variation in recruitment not accounted for by the stock‐recruitment relationship in the sablefish assessment. Recruitment deviations were positively correlated with (i) colder conditions during the spawner preconditioning period, (ii) warmer water temperatures during the egg stage, (iii) stronger cross‐shelf transport to near‐shore nursery habitats during the egg stage, (iv) stronger long‐shore transport to the north during the yolk‐sac stage, and (v) cold surface water temperatures during the larval stage. This result suggests that multiple mechanisms likely affect sablefish recruitment at different points in their life history.     

Disentangling density‐dependent effects on egg production and survival from egg to recruitment in fish

Understanding of density‐dependent effects is key to achieving sustainable management of self‐regulating biological resources such as fish stocks. Traditionally, density‐dependent effects on population abundance in fish have been considered to occur from hatching to recruitment, based on the paradigm of proportionality between spawning stock biomass and total egg production. Here, we demonstrate how the existence of intraspecific and interspecific density dependence in egg production changes the current understanding of density‐dependent processes in the life history of fish, by disentangling density‐dependent effects on egg production and survival from egg to recruitment, using sardine (Sardinops melanostictus, Clupeidae) and anchovy (Engraulis japonicus, Engraulidae) as model species. For sardine, strong intraspecific density‐dependent effects occurred in egg production, but no density‐dependent effects occurred or if any they were weak enough to be masked by environmental factors from hatching to recruitment. In contrast, for anchovy, interspecific density‐dependent effects occurred in egg production. In the survival after hatching, anchovy experienced stronger intraspecific density‐dependent effects than currently recognized. This analysis could overturn the current understanding of density‐dependent effects in the life history, highlighting contrasts between the effects on individual quality and population abundance and between the model species. We propose to reconsider the basis of fisheries management and recruitment studies based on the revised understanding of density‐dependent effects in the life history of the respective species.     

Population dynamics of the exploited sea cucumber Isostichopus fuscus in the western Galápagos Islands, Ecuador

Long-term density data for the sea cucumber, Isostichopus fuscus, from Canal Bolivar (separating western Isabela and eastern Fernandina Islands), Galápagos, were compared with catch statistics and used in a stock depletion model to determine the extent of the effects of the fishery on the existing population and the potential for the population to recover after each fishing season. Recruitment indices were found to be low from 1994 until mid-1999, when a mass recruitment event was registered and numbers of small individuals increased dramatically. The increase in density of juveniles peaked in 2001 and has declined since. Recruitment in this species of sea cucumber appears to be highly sporadic and may depend on intense climatic events such as the warm El Niño. Current levels of fishing activity are unsustainable. The management implications of pulse recruitment in this fishery are discussed.