Shifting dynamic forces in fish stock fluctuations triggered by age truncation?

Accumulating evidence shows that environmental fluctuations and exploitation jointly affect marine fish populations, and understanding their interaction is a key issue for fisheries ecology. In particular, it has been proposed that age truncation induced by fisheries exploitation may increase the population's sensitivity to climate. In this study, we use unique long‐term abundance data for the Northeast Arctic stock of cod (Gadus morhua) and the Norwegian Spring‐Spawning stock of herring (Clupea harengus), which we analyze using techniques based on age‐structured population matrices. After identifying time periods with different age distributions in the spawning stock, we use linear models to quantify the relative effect of exploitation and temperature on the population growth rates. For the two populations, age truncation was found to be associated with an increasing importance of temperature and a relatively decreasing importance of exploitation, while the population growth rate became increasingly sensitive to recruitment variations. The results suggested that the removal of older age classes reduced the buffering capacity of the population, thereby making the population growth rate more dependent on recruitment than adult survival and increasing the effect of environmental fluctuations. Age structure appeared as a key characteristic that can affect the response of fish stocks to climate variations and its consequences may be of key importance for conservation and management.     

Climate change impacts on mismatches between phytoplankton blooms and fish spawning phenology

Substantial interannual variability in marine fish recruitment (i.e., the number of young fish entering a fishery each year) has been hypothesized to be related to whether the timing of fish spawning matches that of seasonal plankton blooms. Environmental processes that control the phenology of blooms, such as stratification, may differ from those that influence fish spawning, such as temperature‐linked reproductive maturation. These different controlling mechanisms could cause the timing of these events to diverge under climate change with negative consequences for fisheries. We use an earth system model to examine the impact of a high‐emissions, climate‐warming scenario (RCP8.5) on the future spawning time of two classes of temperate, epipelagic fishes: “geographic spawners” whose spawning grounds are defined by fixed geographic features (e.g., rivers, estuaries, reefs) and “environmental spawners” whose spawning grounds move responding to variations in environmental properties, such as temperature. By the century's end, our results indicate that projections of increased stratification cause spring and summer phytoplankton blooms to start 16 days earlier on average (±0.05 days SE) at latitudes >40°N. The temperature‐linked phenology of geographic spawners changes at a rate twice as fast as phytoplankton, causing these fishes to spawn before the bloom starts across >85% of this region. “Extreme events,” defined here as seasonal mismatches >30 days that could lead to fish recruitment failure, increase 10‐fold for geographic spawners in many areas under the RCP8.5 scenario. Mismatches between environmental spawners and phytoplankton were smaller and less widespread, although sizable mismatches still emerged in some regions. This indicates that range shifts undertaken by environmental spawners may increase the resiliency of fishes to climate change impacts associated with phenological mismatches, potentially buffering against declines in larval fish survival, recruitment, and fisheries. Our model results are supported by empirical evidence from ecosystems with multidecadal observations of both fish and phytoplankton phenology.     

Spawning stock and recruitment in North Sea cod shaped by food and climate

In order to provide better fisheries management and conservation decisions, there is a need to discern the underlying relationship between the spawning stock and recruitment of marine fishes, a relationship which is influenced by the environmental conditions. Here, we demonstrate how the environmental conditions (temperature and the food availability for fish larvae) influence the stock–recruitment relationship and indeed what kind of stock–recruitment relationship we might see under different environmental conditions. Using unique zooplankton data from the Continuous Plankton Recorder, we find that food availability (i.e. zooplankton) in essence determines which model applies for the once large North Sea cod (Gadus morhua) stock. Further, we show that recruitment is strengthened during cold years and weakened during warm years. Our combined model explained 45 per cent of the total variance in cod recruitment, while the traditional Ricker and Beverton–Holt models only explained about 10 per cent. Specifically, our approach predicts that a full recovery of the North Sea cod stock might not be expected until the environment becomes more favourable.     

The dangers of ignoring stock complexity in fishery management: the case of the North Sea cod

The plight of the marine fisheries is attracting increasing attention as unsustainably high exploitation levels, exacerbated by more extreme climatic conditions, are driving stocks to the point of collapse. The North Atlantic cod (Gadus morhua), a species which until recently formed a major component of the demersal fisheries, has undergone significant declines across its range. The North Sea stock is typical of many, with a spawning stock biomass that has remained below the safe biological limit since 2000 and recruitment levels near the lowest on record. Cod within the North Sea are currently managed as a single stock, and yet mounting empirical evidence supports the existence of a metapopulation of regionally variable, genetically distinct, sub-stocks. Applying the same management strategies to multiple stocks that differ in their resilience to exploitation inevitably results in the overfishing and likely collapse of the weaker components. Indeed, recent studies have identified two North Sea spawning stocks that have undergone disproportionally large collapses with very substantial reductions in egg production. Similarly affected cod stocks in the northwest Atlantic have shown little evidence of recovery, despite fishery closures. The possible implications of ignoring sub-structuring within management units for biocomplexity, local adaptation and ecosystem stability are considered.     

Assessment of exploited fish species in the Lake Edward System,East Africa

The unknown status of inland fish stocks hinders their sustainable management. Therefore, increasing stock status information is important for sustainable inland fisheries. Fisheries reference points were estimated for five exploited fish species (11 stocks) in the Lake Edward system, East Africa, which is one of the most productive inland water systems. The aim was to ascertain the status of the fisheries and establish reference points for effective management. The reference points were based on four linked stock assessment approaches for data-limited fisheries. Estimates showed poor stock status with the stocks defined as either collapsed, recruitment impaired or overfished. However, higher catches could be obtained under sustainable management. Estimates of maximum sustainable yield (MSY) and supporting biomass (B_msy) are provided for 10 of the stocks as targets for rebuilding plans. The immediate target of management should be rebuilding biomass to B_msy. Applicable measures include shifting length at first capture to the length that maximizes catch without endangering size structure and biomass, and livelihood diversification out of fisheries.     

Historical DNA documents long‐distance natal homing in marine fish

The occurrence of natal homing in marine fish remains a fundamental question in fish ecology as its unequivocal demonstration requires tracking of individuals from fertilization to reproduction. Here, we provide evidence of long‐distance natal homing (>1000 km) over more than 60 years in Atlantic cod (Gadus morhua), through genetic analysis of archived samples from marked and recaptured individuals. Using a high differentiation single‐nucleotide polymorphism assay, we demonstrate that the vast majority of cod tagged in West Greenland and recaptured on Icelandic spawning grounds belonged to the Iceland offshore population, strongly supporting a hypothesis of homing. The high degree of natal fidelity observed provides the evolutionary settings for development of locally adapted populations in marine fish and emphasize the need to consider portfolio effects in marine fisheries management strategies.     

Ocean warming,a rapid distributional shift,and the hybridization of a coastal fish species

Despite increasing awareness of large‐scale climate‐driven distribution shifts in the marine environment, no study has linked rapid ocean warming to a shift in distribution and consequent hybridization of a marine fish species. This study describes rapid warming (0.8 °C per decade) in the coastal waters of the Angola‐Benguela Frontal Zone over the last three decades and a concomitant shift by a temperature sensitive coastal fish species (Argyrosomus coronus) southward from Angola into Namibia. In this context, rapid shifts in distribution across Economic Exclusive Zones will complicate the management of fishes, particularly when there is a lack of congruence in the fisheries policy between nations. Evidence for recent hybridization between A. coronus and a congener, A. inodorus, indicate that the rapid shift in distribution of A. coronus has placed adults of the two species in contact during their spawning events. Ocean warming may therefore revert established species isolation mechanisms and alter the evolutionary history of fishes. While the consequences of the hybridization on the production of the resource remain unclear, this will most likely introduce additional layers of complexity to their management.     

Climate effects on size‐at‐age: growth in warming waters compensates for earlier maturity in an exploited marine fish

Over the past 3 decades, North Sea Atlantic cod (Gadus morhua) have exhibited variable length‐at‐age along with declines in spawning stock biomass and timing of maturity. Multiple factors affecting growth and development in fish acted on this economically important stock over the same period including warming waters and an intensive fishery. Here, we employ North Sea cod as a model population, exploring how a physiologically relevant temperature metric (the growing degree‐day, GDD; °C day) can be used to compare year‐classes on a physiologically relevant time‐scale, disentangling influences of climate (thermal history) on observed length‐at‐age trends. We conclude that the trends in North Sea cod length‐at‐age observed during the last three decades can be explained by a combination of temperature‐dependent growth increases and a trend toward earlier maturation, the latter likely induced by the intensive fishing pressure, and possibly evidence of fisheries‐induced evolution.     

Forecasting future recruitment success for Atlantic cod in the warming and acidifying Barents Sea

Productivity of marine fish stocks is known to be affected by environmental and ecological drivers, and global climate change is anticipated to alter recruitment success of many stocks. While the direct effects of environmental drivers on fish early life stage survival can be quantified experimentally, indirect effects in marine ecosystems and the role of adaptation are still highly uncertain. We developed an integrative model for the effects of ocean warming and acidification on the early life stages of Atlantic cod in the Barents Sea, termed SCREI (Simulator of Cod Recruitment under Environmental Influences). Experimental results on temperature and CO₂ effects on egg fertilization, egg and larval survival and development times are incorporated. Calibration using empirical time series of egg production, temperature, food and predator abundance reproduces age‐0 recruitment over three decades. We project trajectories of recruitment success under different scenarios and quantify confidence limits based on variation in experiments. A publicly accessible web version of the SCREI model can be run under www.oceanchange.uni-bremen.de/ ;SCREI. Severe reductions in average age‐0 recruitment success of Barents Sea cod are projected under uncompensated warming and acidification toward the middle to end of this century. Although high population stochasticity was found, considerable rates of evolutionary adaptation to acidification and shifts in organismal thermal windows would be needed to buffer impacts on recruitment. While increases in food availability may mitigate short‐term impacts, an increase in egg production achieved by stock management could provide more long‐term safety for cod recruitment success. The SCREI model provides a novel integration of multiple driver effects in different life stages and enables an estimation of uncertainty associated with interindividual and ecological variation. The model thus helps to advance toward an improved empirical foundation for quantifying climate change impacts on marine fish recruitment, relevant for ecosystem‐based assessments of marine systems under climate change.     

Effects of river temperature and climate warming on stock‐specific survival of adult migrating Fraser River sockeye salmon (Oncorhynchus nerka)

Mean summer water temperatures in the Fraser River (British Columbia, Canada) have increased by ～1.5 °C since the 1950s. In recent years, record high river temperatures during spawning migrations of Fraser River sockeye salmon (Oncorhynchus nerka) have been associated with high mortality events, raising concerns about long‐term viability of the numerous natal stocks faced with climate warming. In this study, the effect of freshwater thermal experience on spawning migration survival was estimated by fitting capture–recapture models to telemetry data collected for 1474 adults (captured in either the ocean or river between 2002 and 2007) from four Fraser River sockeye salmon stock‐aggregates (Chilko, Quesnel, Stellako‐Late Stuart and Adams). Survival of Adams sockeye salmon was the most impacted by warm temperatures encountered in the lower river, followed by that of Stellako‐Late Stuart and Quesnel. In contrast, survival of Chilko fish was insensitive to the encountered river temperature. In all stocks, in‐river survival of ocean‐captured sockeye salmon was higher than that of river‐captured fish and, generally, the difference was more pronounced under warm temperatures. The survival–temperature relationships for ocean‐captured fish were used to predict historic (1961–1990) and future (2010–2099) survival under simulated lower river thermal experiences for the Quesnel, Stellako‐Late Stuart and Adams stocks. A decrease of 9–16% in survival of all these stocks was predicted by the end of the century if the Fraser River continues to warm as expected. However, the decrease in future survival of Adams sockeye salmon would occur only if fish continue to enter the river abnormally early, towards warmer periods of the summer, as they have done since 1995. The survival estimates and predictions presented here are likely optimistic and emphasize the need to consider stock‐specific responses to temperature and climate warming into fisheries management and conservation strategies.     

Marine reserves and reproductive biomass: a case study of a heavily targeted reef fish

Recruitment overfishing (the reduction of a spawning stock past a point at which the stock can no longer replenish itself) is a common problem which can lead to a rapid and irreversible fishery collapse. Averting this disaster requires maintaining a sufficient spawning population to buffer stochastic fluctuations in recruitment of heavily harvested stocks. Optimal strategies for managing spawner biomass are well developed for temperate systems, yet remain uncertain for tropical fisheries, where the danger of collapse from recruitment overfishing looms largest. In this study, we explored empirically and through modeling, the role of marine reserves in maximizing spawner biomass of a heavily exploited reef fish, Lethrinus harak around Guam, Micronesia. On average, spawner biomass was 16 times higher inside the reserves compared with adjacent fished sites. Adult density and habitat-specific mean fish size were also significantly greater. We used these data in an age-structured population model to explore the effect of several management scenarios on L. harak demography. Under minimum-size limits, unlimited extraction and all rotational-closure scenarios, the model predicts that preferential mortality of larger and older fish prompt dramatic declines in spawner biomass and the proportion of male fish, as well as considerable declines in total abundance. For rotational closures this occurred because of the mismatch between the scales of recovery and extraction. Our results highlight how alternative management scenarios fall short in comparison to marine reserves in preserving reproductively viable fish populations on coral reefs.     

Variation in egg mass within two Atlantic herring Clupea harengus stocks

The main objective of this study was to investigate if egg size (mass) at spawning is invariant for Scotia-Fundy summer and autumn (SFSH) and Icelandic summer (ISSH) spawning herring Clupea harengus. Oocyte dry mass measurements for SFSH females collected in 2001 and ISSH females collected in 1999 and 2000 showed a large variation. Difference in egg dry mass among fish was found to vary by as much as twofold in each stock. For ISSH, variation in egg mass was also apparent from oocyte volume measurements made jointly with a histological examination of the ovaries. Approximately 20% of the variation in egg mass could be explained by maternal whole-body mass or total length, indicating that length or age composition in the stocks can potentially influence the recruitment success. This implies that fisheries management strategies should aim to maintain a broad range in age composition.     

Interannual variability in reproductive traits of the Patagonian toothfish Dissostichus eleginoides around the sub‐Antarctic island of South Georgia

Commercial fisheries data, collected as part of an observer programme and covering the period 1997–2014, were utilized in order to define key reproductive traits and spawning dynamics of the Patagonian toothfish Dissostichus eleginoides at South Georgia. Multi‐year spawning site fidelity of D. eleginoides was revealed through the identification of previously unknown spawning hotspots. Timing of female spawning was shown to have shifted later, leading to a shorter spawning duration. A decrease in length and mass of female and male spawning fish and a reduced number of large spawning fish was found, evidence of a change in size structure of spawning D. eleginoides. During the study period fewer later maturity stage females (including spawning stage) were observed in conjunction with increased proportions of early stage female D. eleginoides. The findings are discussed in the context of reproductive success, with consideration of the possible effects such spawning characteristics and behaviours may have on egg and larval survival. This work presents the first long‐term assessment of D. eleginoides spawning dynamics at South Georgia and provides valuable knowledge for both the ecology of the species and for future fisheries management of this commercially important species.     

西北太平洋柔鱼栖息地环境因子分析及其对资源丰度的影响

柔鱼(Ommastrephes bartramii)是西北太平洋海域重要的经济头足类,短生命周期的生活史特征决定其资源丰度易受海洋环境变化影响。根据1998—2010年我国鱿钓船生产统计资料和环境资料,包括海表温度(SST)和叶绿素浓度(Chl-a)数据,结合Nio 3.4区海表温距平值(SSTA),分析了SST和Chl-a浓度的季节和年际变化特征,并分别探讨了SST和Chl-a浓度距平值与Nio 3.4区SSTA及柔鱼资源丰度之间的关系。结果表明,产卵场海域Chl-a浓度冬季高夏季低,SST则夏季高冬季低;育肥场Chl-a和SST均呈夏季高冬季低变化,但6—12月份Chl-a浓度波动明显。产卵场和育肥场SST及Chl-a浓度年际变化明显。同时研究发现,厄尔尼诺、拉尼娜和正常年份时Nio 3.4区SSTA对柔鱼产卵场和育肥场环境的调控机制不同:拉尼娜和正常年份产卵场和育肥场温度上升,叶绿素浓度变化幅度小,有利于资源量补充,产量较高;厄尔尼诺年份温度和叶绿素均降低,尤其育肥场叶绿素浓度,对资源量产生不利影响,产量锐减。研究利用多元线性回归分别建立了基于温度和叶绿素的柔鱼资源丰度的预测模型,两者均能很好的预测柔鱼资源丰度(P0.05),但基于叶绿素的预测模型优于温度模型。     

Watershed‐scale climate influences productivity of Chinook salmon populations across southcentral Alaska

The ecosystems supporting Pacific salmon (Oncorhynchus spp.) are changing rapidly as a result of climate change and habitat alteration. Understanding how—and how consistently—salmon populations respond to changes at regional and watershed scales has major implications for fisheries management and habitat conservation. Chinook salmon (O. tshawytscha) populations across Alaska have declined over the past decade, resulting in fisheries closures and prolonged impacts to local communities. These declines are associated with large‐scale climate drivers, but uncertainty remains about the role of local conditions (e.g., precipitation, streamflow, and stream temperature) that vary among the watersheds where salmon spawn and rear. We estimated the effects of these and other environmental indicators on the productivity of 15 Chinook salmon populations in the Cook Inlet basin, southcentral Alaska, using a hierarchical Bayesian stock‐recruitment model. Salmon spawning during 2003–2007 produced 57% fewer recruits than the previous long‐term average, leading to declines in adult returns beginning in 2008. These declines were explained in part by density dependence, with reduced population productivity following years of high spawning abundance. Across all populations, productivity declined with increased precipitation during the fall spawning and early incubation period and increased with above‐average precipitation during juvenile rearing. Above‐average stream temperatures during spawning and rearing had variable effects, with negative relationships in many warmer streams and positive relationships in some colder streams. Productivity was also associated with regional indices of streamflow and ocean conditions, with high variability among populations. The cumulative effects of adverse conditions in freshwater, including high spawning abundance, heavy fall rains, and hot, dry summers may have contributed to the recent population declines across the region. Identifying both coherent and differential responses to environmental change underscores the importance of targeted, watershed‐specific monitoring and conservation efforts for maintaining resilient salmon runs in a warming world.     

Self‐recruitment and sweepstakes reproduction amid extensive gene flow in a coral‐reef fish

Identifying patterns of larval dispersal within marine metapopulations is vital for effective fisheries management, appropriate marine reserve design, and conservation efforts. We employed genetic markers (microsatellites) to determine dispersal patterns in bicolour damselfish (Pomacentridae: Stegastes partitus). Tissue samples of 751 fish were collected in 2004 and 2005 from 11 sites encompassing the Exuma Sound, Bahamas. Bayesian parentage analysis identified two parent–offspring pairs, which is remarkable given the large population sizes and 28 day pelagic larval duration of bicolour damselfish. The two parent–offspring pairs directly documented self‐recruitment at the two northern‐most sites, one of which is a long‐established marine reserve. Principal coordinates analyses of pair‐wise relatedness values further indicated that self‐recruitment was common in all sampled populations. Nevertheless, measures of genetic differentiation (F_ST) and results from assignment methods suggested high levels of gene flow among populations. Comparisons of heterozygosity and relatedness among samples of adults and recruits indicated spatially and temporally independent sweepstakes events, whereby only a subset of adults successfully contribute to subsequent generations. These results indicate that self‐recruitment and sweepstakes reproduction are the predominant, ecologically‐relevant processes that shape patterns of larval dispersal in this system.     

Stock structure analysis of ‘Bombay duck’ (Harpadon nehereus Hamilton, 1822) along the Indian coast using truss network morphometrics

Harpadon nehereus, commonly known as ‘Bombay duck’, is a fish with a discontinuous distribution along the Indian peninsula. The fisheries are dominant on the north‐east and north‐west coast but are absent in commercial landings below 15° north latitude. Heretofore stock assessment studies had not considered the various spawning stock components that replenish this fishery, therefore the present study. Fish samples were collected from four locations: two each from the northeast and the northwest coasts. Twenty‐four morphometric variables were measured using a box‐truss network method. Factor analysis of these variables differentiated the east and the west coast fish populations. Multiple comparisons on the factor scores indicated two independent stocks on the east coast, whereas the fishery on the west coast is replenished by a single stock. The important morphometric traits that accounted for most of the stock variations were related to swimming adaptations of the fish. Future stock assessments can consider the population on the west coast as a single stock when formulating management plans. To harvest the resource in a sustainable manner, the maritime states on the west coast should adopt collaborative efforts towards managing this fishery.     

Reproductive biology and spatiotemporal patterns of spawning in striped marlin Kajikia audax

This study presents the first histology‐based assessment of the reproductive dynamics of south‐west Pacific striped marlin Kajikia audax. Maturity and reproductive status were assessed from histological sections of ovaries (n = 234) and testes (n = 243) of fish caught in commercial longline and recreational fisheries between 2006 and 2009. Spawning peaked in the Coral Sea during November and December at sea surface temperatures between 24·8 and 28·3° C. Lower jaw fork length (L_LJF) at 50% maturity (L_LJF50), a key variable for stock assessment, was estimated to be 2100 ± 102 mm (mean + s.e .) for females and 1668 ± 18 mm for males. Unlike large pelagic tunas Thunnus spp., the proportion of females increased with length and spawning fish formed multiple large‐scale aggregations within a broad latitudinal band. This study provides a starting point for biological parameters needed for stock assessment and conservation of K. audax and introduces the multiple aggregation spawning concept as a reproductive mechanism to explain genetic heterogeneity observed in some highly migratory species.     

Recruitment prediction for Pacific herring (Clupea pallasi) on the west coast of Vancouver Island, Canada

The accurate prediction of recruitment to the fishery is a very important tool within the management structure of any fish stock being exploited. In the case of the Pacific herring, Clupea pallasi, fishery in Canada, a forecast of the abundance of each herring stock is particularly important for formulating an annual catch quota. The sustainable management of the fishery and the resource is based in part on accurate recruitment forecasting because Pacific herring are short-lived and so the recruitment contributes a significant part of the total spawning run targeted by the fishery each year. Several factors are believed be important in determining the success of recruitment besides spawners biomass. Since herrings are “r” strategists, conditions related to the egg, the planktonic, or even the juvenile stage might determine the future level of recruitment. Recently a formula that defines conditions for a semi-quantitative level of recruitment forecast was elaborated using genetic algorithms and current study attempts to improve on this model. Using salinity in two quarterly periods during the planktonic and pre-recruit stages, temperature and spawning biomass for the west coast of Vancouver Island stock, classification rules that define recruitment in 3 different levels (low, medium and high) were developed with a genetic algorithm, setting low and high boundaries for each condition. A 75% success in classifying recruitment was obtained. The model was shown to be particularly effective at predicting when the recruitment would be low, which could be important from the perspective of the Precautionary Approach and the sustainable management of this stock.