The recent decline in many fish stocks has been accompanied by marked changes in age at maturity. However, there is little information on whether other components of reproductive investment have changed and the relative roles of phenotypic plasticity or genetic selection in such change. This study explored how reproductive investment in cod Gadus morhua from different regions around Scotland has changed between a period of high (1969, 1970) and low (2002, 2003) spawning-stock size. Maturity ogives indicated that inshore North Sea cod now mature at a smaller size and younger age than they did around 1970. The relationship between potential fecundity and size indicated that cod in the inshore region are now more fecund than 30 yr ago. Cod from the inshore region of the North Sea now have a higher fecundity at a given size and age than cod from the offshore region and the west of Scotland. There was no such spatial trend around 1970. Similarly, whilst the relative fecundity of cod in recent times is positively correlated with age, no such relationship was evident in 1969 and 1970. Nutritional influences on maturity and fecundity, particularly liver condition, were also evident from the recent study. However, the temporal and spatial differences in fecundity-size relationships were not consistent with a change in growth conditions, as there has been a significant decrease in body condition in recent times. Our findings regarding the temporal change in reproductive investment in the inshore North Sea region appear consistent with genotypic changes in life-history traits predicted to arise from intense periods of size-selective mortality.
A paradigm of fisheries science holds that spawning stock biomass (SSB) is directly proportional to total egg production (TEP) of fish stocks. This “SSB–TEP proportionality” paradigm has been a basic premise underlying the spawner–recruitment models for fisheries management and numerous studies on recruitment mechanisms of fish. Studies on maternal effects on reproductive potential of a stock have progressed during the last few decades, leading to doubt concerning the paradigm. Nonetheless, a direct test of the paradigm at multidecadal scales has been difficult because of data limitations in the stock assessment systems worldwide. Here, we tested the paradigm for marine fish based on a novel combination of two independent 38‐year time series: fishery‐dependent stock assessment data and fishery‐independent egg survey data. Through this approach, we show that the SSB–TEP proportionality is distorted by density dependence in total egg production per spawner individual (TEPPS) or spawner unit weight (TEPPSW) at a multidecadal scale. The TEPPS/TEPPSW exponentially declined with biomass and thus was density‐dependent for Japanese sardine, a small pelagic species exhibiting a high level of population fluctuation, in the western North Pacific. By contrast, the TEPPS/TEPPSW was sardine‐density‐dependent for Japanese anchovy, another small pelagic species exhibiting a moderate level of population fluctuation well‐known for being out of phase with sardine. Our analysis revealed intraspecific (sardine) and interspecific (anchovy) density dependence in TEPPS/TEPPSW, which was previously unaccounted for in spawner–recruitment relationships. Such density‐dependent effects at the time of spawning should be considered in fisheries management and studies on recruitment mechanisms.
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.
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