Behaviour and thermal experience of adult sockeye salmon migrating through stratified lakes near spawning grounds: the roles of reproductive and energetic states

Roscoe DW, Hinch SG, Cooke SJ, Patterson DA. Behaviour and thermal experience of adult sockeye salmon migrating through stratified lakes near spawning grounds: the roles of reproductive and energetic states. Ecology of Freshwater Fish 2010: 19: 51–62. © 2009 John Wiley & Sons A/S
Abstract –  Little is known about physiological factors underlying thermal behaviour in Pacific salmon ( Oncorhynchus spp.). We implanted acoustic transmitters and temperature loggers into migrating adult sockeye salmon ( Oncorhynchus nerka ) and nonlethally assessed their reproductive hormone levels and energetic states immediately prior to their passing through natal lakes en route to spawning grounds. We tested the hypothesis that energetic and reproductive status influence thermoregulatory and other in-lake behaviours. More reproductively advanced females with lower levels of energy transited through cooler temperatures compared to less mature females with high levels of energy, possibly to reduce metabolic energy expenditure and delay final maturation. Transit temperatures of males were not related to physiological variables. Salmon travelled on average 13.6 km·day⁻¹ through two lakes, and often circled or held for more than 1 day before moving upstream, but these behaviours were generally not related to physiological variables.     

Dietary fatty acid enrichment during the spawning season increases egg viability and quality in Hippocampus hippocampus

Control of the reproductive cycle is essential to achieve a successful aquaculture industry. Seahorse reared in captivity often shows eggs of poor quality and low offspring survival due to unsuitable rearing and nutritional protocols. This study aimed to evaluate the effect of dietary enrichment, added exclusively during the spawning season, on the egg quality of Hippocampus hippocampus. The effect of enriching mysis throughout the spawning season was analysed in terms of female fecundity, egg viability, egg size and fatty acid profile. The results showed that fecundity was not affected by the enrichment of the diet, but egg viability was increased when females were fed the enriched mysis (84.9 ± 4.1% compared to 62.2 ± 9.4%). Moreover, the content of several PUFA (20:4n‐6, 18:2n‐6, 22:5n‐3, 22:5n‐6, sum of n‐6) decreased throughout the spawning season, especially in the last clutch, in eggs originated from females fed non‐enriched mysis. Eggs from females fed the enriched diet maintained the fatty acid profile throughout the spawning season. In conclusion, enrichment of the diet of H. hippocampus improved egg viability and avoided a decrease in the quality of eggs by maintaining PUFA content in the last clutches of the spawning season.     

Environmental characteristics of spawning and nursery grounds of Japanese sardine and mackerels in the Kuroshio and Kuroshio Extension area

The sustainable use of marine resources requires understanding the surrounding ecosystem and elucidating mechanisms of variation. However, we still lack a comprehensive understanding of environmental variation in the spawning and nursery grounds of important fisheries species Japanese sardine (Sardinops melanostictus) and mackerels (Scomber japonicus and Scomber australasicus) in the northwest Pacific. Here, we investigate detailed physical, chemical, and biological environment variations in the spawning and nursery grounds along the Kuroshio and Kuroshio Extension area from intensive investigation in spawning season (April) of 2013. We found similar water mass property and copepod community in the egg‐rich Kuroshio area and the larvae‐rich downstream Kuroshio Extension area, indicating environmental variability is small during transportation and development processes. The egg‐rich northern Izu Islands region showed high copepod abundance, although low nutrient and chlorophyll concentrations were observed. Eggs were scarce or absent in the second survey 10 days after abundant eggs were observed in the region, along with differences in water property and copepod community. This indicates that not only the location but also the specific water characteristic and copepod community are a determining factor for spawning. Indicator communities of copepod found in our study (indicator community of transportation process from spawning ground, of non‐spawning ground, and of reproductive area in the Kuroshio Extension area) would be a key factor for recruitment prediction.     

Modelling transport of inshore and deep‐spawned chokka squid (Loligo reynaudi) paralarvae off South Africa: the potential contribution of deep spawning to recruitment

The South African chokka squid, Loligo reynaudi, spawns both inshore (≤70 m) and on the mid‐shelf (71–130 m) of the Eastern Agulhas Bank. The fate of these deep‐spawned hatchlings and their potential contribution to recruitment is as yet unknown. Lagrangian ROMS‐IBM (Regional Ocean Modelling System‐Individual‐Based Model) simulations confirm westward transport of inshore and deep‐spawned hatchlings, but also indicate that the potential exists for paralarvae hatched on the Eastern Agulhas Bank deep spawning grounds to be removed from the shelf ecosystem. Using a ROMS‐IBM, this study determined the transport and recruitment success of deep‐spawned hatchlings relative to inshore‐hatched paralarvae. A total of 12 release sites were incorporated into the model, six inshore and six deep‐spawning sites. Paralarval survival was estimated based on timely transport to nursery grounds, adequate retention within the nursery grounds and retention on the Agulhas Bank shelf (<200 m). Paralarval transport and survival were dependent on both spawning location and time of hatching. Results suggest the importance of the south coast as a nursery area for inshore‐hatched paralarvae, and similarly the cold ridge nursery grounds for deep‐hatched paralarvae. Possible relationships between periods of highest recruitment success and spawning peaks were identified for both spawning habitats. Based on the likely autumn increase in deep spawning off the Tsitsikamma coast, and the beneficial currents during this period (as indicated by the model results) it can be concluded that deep spawning may at times contribute significantly to recruitment.     

Annually twice induced spawnings provide multiple benefits: Experimental evidence from an Indian major carp (Catla catla,Hamilton 1822)

Multiple spawnings are special reproductive tactics that many fish evolved to obtain different types of direct and/or indirect spawning benefits. This has led to two pertinent questions: (a) what types of benefit does a commercially important fish provide during multiple induced spawnings annually? and (b) is there any trade‐off between traits during these spawnings? To address these questions, the present study was carried out with two separate experiments during two spawning events of an Indian major carp (Catla catla) to assess: (a) male's spawning performances and (b) female's spawning performances. The male's spawning data revealed that they produced significantly higher amount of milt and hatchlings during the first spawning, while they significantly reduced hatchling deformation rate, enhanced offspring survival rate and enlarged offspring body size during the second spawning. The female's spawning information unveiled no significant differences in total egg weight, hatching rate and hatchling deformation rate between the first and second spawning. However, females had significantly higher offspring survival rate during the first spawning and enhanced offspring body size during the second spawning. Interestingly, the analysis of trait associations revealed that males allocated energy with a trade‐off between milt weight and offspring survival rate, whereas females showed a trade‐off between hatching rate and offspring survival rate. Thus, the study provides some important information for carp breeders and associated stakeholders to know which spawning season is preferable for the higher production of eggs, milt and good quality larvae.     

Impact of long‐term dietary exposure to lead on some reproductive parameters of a female Common carp (Cyprinus carpio L.)

During a long‐term 3‐year dietary exposure of mature carp females to lead, its bioaccumulation in the brain, changes in the neurohormonal activity at the level of the hypothalamus and the pituitary gland as well as the growth and maturation of ovaries were analysed. Moreover, an analysis of the effectiveness of hormone‐stimulated spawning during two subsequent spawning seasons was carried out. The results of the analyses show that chronic exposure of maturing and mature carp females to lead in feed results in its accumulation in the brain (1.365 μg g^?1). This impairs the endocrine activity of the hypothalamus, which is manifested by, among others, an increased secretion of dopamine and impaired spontaneous secretion of luteinizing hormone (LH). The observed decrease in the concentration of gonadotropin in blood inhibits vitellogenesis, which is manifested by a lower degree of the maturity of oocytes, lower fecundity (as measured by egg number, egg weight, egg maturity and egg survival) and lower gonadosomatic index values. The final negative impact of lead is the impairment of reproductive functions, as manifested by a smaller number of spawning females and in their lower fecundity during spawning.     

Variations in reproductive potential between nearshore and offshore spawning contingents of hogfish in the eastern Gulf of Mexico

Hogfish, Lachnolaimus maximus (Walbaum), have multiple traits that confound measuring reproductive potential: they are protogynous, relatively long‐lived harem‐forming fish that spawn daily for months. Additionally, recent evidence demonstrates that size, age and timing of sex change vary on a spatial scale within the study area (West Florida shelf, USA). This study investigates the effect of this spatial variation on hogfish reproductive potential by evaluating spawning seasonality, spawning frequency and batch fecundity using an indeterminate egg production model. Offshore females were larger than nearshore females, and batch fecundity was related in a log‐linear manner to female size. Gonad histology demonstrated a more protracted reproductive period for females offshore (8 months) than nearshore (4 months). Spatial variations in size coincide with ontogeny because hogfish move offshore with growth; however, even after accounting for fish size, offshore females spawned more. In areas where male removal rates are elevated, spawning harems are disrupted; thus, greater fishing effort nearshore may further reduce the reproductive potential of these females. These nearshore and offshore spawning components of the population are not genetically distinct, but instead represent two contingent spawning strategies that likely enhance total population stability and resilience of this stock in the eastern Gulf of Mexico.     

Skipped spawning in female iteroparous fishes

It is often assumed that iteroparous fishes spawn annually once reaching sexual maturity, but this is not always the case. This paper reviews available information on skipped spawning in female teleost fishes. All instances of non‐annual spawning are described as one of three types (retaining, reabsorbing, resting), depending on where in the normal spawning cycle development has been interrupted. Retaining ripe eggs is caused by conditions experienced during the spawning season (fish density, mate availability, pollution), whereas failure to start vitellogenesis (resting) or the breakdown of all oocytes that enter into vitellogenesis (reabsorbing) is caused by factors experienced prior to the spawning season (primarily temperature and poor nutrition). It is speculated that the relative shortage of data on non‐annual spawning may be because of difficulties in identifying non‐reproductive individuals. In an attempt to rectify this situation, the criteria needed to identify females undergoing the three forms of spawning omission are presented in terms of external appearance of gonads, gonad indices, and histological analysis. The energy saved by not spawning in a poor year may lead to increased survival and the probability of spawning in subsequent years. As the cumulative number of progeny gained by surviving to spawn in multiple subsequent years outweighs the number of progeny lost by not spawning in a given single year, occasional omission of spawning may constitute an adaptive trait in long‐lived iteroparous fishes.     

Effects of Varying Dietary Protein Levels and Feeding Frequencies on Condition and Reproductive Performance of Channel Catfish to Produce Hybrid Catfish

The interspecific hybridization of channel catfish, Ictalurus punctatus, females with blue catfish, I. furcatus, males has been identified as a method to further improve production; however, lack of spawning success has affected its commercial application. To facilitate our understanding of the interaction of brood stock nutrition and reproductive performance, we evaluated the interaction of feed quality and feeding frequency. Channel catfish females were classified into two genetic groups, namely, high and low spawning. The treatments were offered during the spring season 70–90 d prior to the start of the spawning season. Induced reproduction was performed using luteinizing hormone releasing hormone analog. Condition of the fish as well as reproductive performance using spawning success, egg production, egg size, and fertilization at 48 h were determined. Changing protein level of the diet from 32 to 42% did not influence spawning, fecundity, or fertilization, but affected egg size and biochemical composition of the eggs. Increasing the feeding frequency from three to six times per week negatively affected spawning in one of the two genetics groups, did not affect egg production and egg fertilization, but had a significant effect on egg size. Older fish performed better than younger fish in terms of spawning success and egg production.     

Littoral spawning habitats of three southern Arctic charr (Salvelinus alpinus L.) populations

Abstract Arctic charr populations in southern latitudes are nonmigratory, with all life‐stages limited to freshwater lakes and in‐ or out‐flowing tributaries. Although many of these populations are reported to also spawn in lake littorals, little is known about the physical characteristics of putative spawning grounds. A total of 23 discrete spawning sites within three Irish lakes were located by fyke netting of spawning adults and snorkelling in littoral habitats. Spawning sites were found to be long, narrow strips running parallel to the shore at a maximum depth of 124 cm. Spawning sites were limited to areas of coarse mineral substrate with an adequate (c. 8 cm) depth of clean interstitial spaces. In individual lakes, combined areas of spawning sites made up 0.4–0.7% of available littoral. Egg densities varied considerably between sites (33–900·eggs m^?2) and were significantly correlated with gradient and width of spawning sites. No evidence of redd digging was found. The shallow, localised and restricted nature of spawning grounds makes such populations vulnerable to anthropogenically induced postoviposition changes in surface water level, eutrophication processes such as increased lake sedimentation and elevated nutrient status.     

Spawning season,spawning grounds,and egg production of red sea bream in Hiuchi-nada,Seto Inland Sea

The spawning season and grounds of red sea bream in Hiuchi-nada, the central part of the Seto Inland Sea, were described using a new method based on monoclonal antibodies for identifying Pagrus major eggs, and the daily egg production (standardized by the incubation time and survival rate) was estimated. At the peak of spawning (May), the ranges of sea temperature, salinity, and chlorophyll a where red sea bream eggs occurred were 14.8–17.4°C, 32.0–33.0, 0.5–4.4 μg/L, respectively. The main spawning grounds of the red sea bream were confirmed as being the areas near the Geiyo Islands, Misaki Peninsula, Saijyo, Niihama. The spatial spread of red sea bream eggs increased with the egg developmental stage. The ranges of daily egg production in 2005 and 2006 were 0.3–19.3 and 0.2–6.7 × 10⁹ eggs/day, respectively. In Hiuchi-nada, aquaculture farms are located close to the spawning grounds, and the potential spawning population from the aquaculture farms equaled or exceeded that of the estimated spawning population obtained by the egg production method. Red sea bream eggs in Hiuchi-nada might be produced by both wild and aquaculture-based spawning populations.     

Effects of Water Hardness and Calcium: Magnesium Ratios on Reproductive Performance and Offspring Quality of Macrobrachium rosenbergii

This study evaluated the effect of specific calcium and magnesium ratios at two hardness values on reproductive performance and offspring quality of Macrobrachium rosenbergii brood‐females. Hatchery water containing 110.5 mg/L CaCO3 hardness was considered baseline as it has been used successfully to hatch and rear larvae and was used as the control treatment. At each hardness value of 150 and 190 mg/L CaCO₃, four experimental water treatments were made in triplicate. Each treatment was adjusted using soluble salts (CaCl₂·H₂O, MgSO₄·7H₂O, and MgCl₂·6H₂O) to provide the hardness and calcium to magnesium ratios of 0:20, 20:80, 50:50, and 80:20 needed. Each of the experimental tanks (140 L) were stocked with six females and one male. The results indicated that both hardness and Ca:Mg ratios affect brood‐female reproductive qualitative parameters such as intermolt period, egg hatchability, egg dry weight, and egg‐clutch somatic index (ESI) parameters (P < 0.05) but not fecundity and eggs per spawn. The results revealed that brood‐females at 150 mg/L hardness showed greater reproductive performance than at 190 and 110.5 mg/L hardness. The findings also demonstrated that the treatment 50Ca50Mg at 150 mg/L hardness with 38.8 mg/L calcium and 12.9 mg/L magnesium had optimum reproductive performance and offspring quality for M. rosenbergii brood‐females.     

Effect of density on brood size in noble crayfish,Astacus astacus L., subjected to indoor rearing conditions*

Abstract. The effects of density, cover, and individual versus communal holding on brood size and female survival during the breeding period were examined in a large-scale experiment, involving 3591 mature noble crayfish, Astacus astacus L., females. Crayfish density during both mating/spawning and winter incubation had major effects on egg survival. In the high density experiment (67 and 44 individuals per m² during mating/spawning and winter incubation respectively) mean pleopod egg number in April was 18-20. Lowering the density to 50 during mating/spawning (still 44 individuals per m² during winter incubation) gave mean pleopod egg numbers of 31-38. In the low density experiment (50 and 33 individuals per m² during mating/spawning and winter incubation) mean pleopod egg numbers were 44-46. Egg-bearing females held in individual compartments with perforated bottom had a mean pleopod egg number of 25. Female survival during winter incubation was high, ranging from 91.8 to 98.3%. Covering the basins had no distinct effect on egg and female survival. In Astacus astacus culture we recommend that crayfish density should not exceed about 50 individuals per m² during mating/spawning and about 33 individuals per m² during winter incubation.     

Migration,homing and spatial ecology of common carp in interconnected lakes

The common carp, Cyprinus carpio, is a large, long-lived, fecund and mobile cyprinid, which evolved in complex inter-braided Ponto-Caspian rivers that experience both springtime flooding and freezing winters. Studies suggest adults often move to productive, shallow lakes and floodplains to spawn because they often lack egg predators and then return to deeper normoxic waters to overwinter. Whether these movements involve individuals consistently selecting, or homing to, the same spawning and refuge lakes as part of a strategy benefiting their reproductive success is unknown. To address this question, we examined the movements of 67 radio-tagged adult carp for 3 years in a watershed with 11 interconnected lakes. Carp were tagged and released into a centrally located, normoxic deep lake in spring and fall. Each spring over 95% of its adults left via a single stream and swam into one of 5 shallow lakes, with most individuals (84%) selecting the same lake(s) in which to spawn each year (median Bhattacharyya affinity coefficient of similarity of 0.82). Young were later found in those lakes without egg predators, which cannot survive winter anoxia. After spawning, carp spent summers moving between productive lakes in an individualistic fashion, presumably foraging, with most (89%) eventually returning to the same deep lake to overwinter (median Bhattacharyya affinity of 1.0). These movements appear to reveal a life-history tactic involving seasonal homing migration, first to a spawning location and later to winter refuges, that is well adapted to productive but highly heterogeneous and interconnected freshwater environments.     

A spatial model of population dynamics of the early life stages of Japanese sardine, Sardinops melanostictus, off the Pacific coast of Japan

We constructed a numerical model reproducing the transport, survival and individual growth of the early life stages of Japanese sardine, Sardinops melanostictus, off the Pacific coast of Japan during 1978–93. The causes of early life stage mortality, including the influence of the effects of the spatial relationship between the spawning grounds and the Kuroshio on the mortality rate, were investigated. Survival and transport from egg stage to 60 days after spawning were modelled daily in a 1 × 1 degree mesh cell and individual growth in the period was modelled in each region (Kuroshio, Inshore, Offshore and Transition regions). Individual growth and survival from 60 to 180 days after spawning were modelled daily in the Transition region. Environmental data were taken from outside the model system. Our simulation indicates that survival variability in the larval stage (5–25 mm in standard length) is the key factor in determining the year‐class strength. The simulation revealed that strong year classes occurred with good survival in the spawning ground and whilst entrained in the Kuroshio current being transported to the main feeding grounds in the Transition region. The simulation also indicated that survival rates in 1988–93 were low in the Inshore, Kuroshio and Offshore regions, which depressed the year‐class strength during that period.     

Characteristics and rehabilitation of the spawning habitats of the sea trout, Salmo trutta, in Gotland (Sweden)

Characteristics of the natural spawning habitat of sea trout, Salmo trutta L., were studied in Själsöån, a small stream of Gotland, Sweden, from 1992 to 1999. Each year, trout spawned on 17 ± 7 different areas (156 places ha^?1). Most of the spawning grounds were used every year. Overcutting was evident for at least 60% of the spawning grounds. Fish spawned on a gravel of 19 ± 7 mm in diameter. From 1978 to 1992, 242 artificial spawning grounds were constructed by the Gotland Sport Fishermen Association in four Gotland streams. A sediment trap was dug upstream to the spawning grounds. The artificial spawning ground comprised of a downstream V‐shape stream deflector of large stones with a log weir at the narrowest point of the deflector. Upstream of the dam, 15–60 mm diameter gravel was deposited to constitute the spawning ground substratum. To keep the installation efficient, maintenance is needed every year before the spawning season.     

Suspending Mammalian LHRHa‐injected Channel Catfish,Ictalurus punctatus,in Individual Soft Mesh Bags Reduces Stress and Improves Reproductive Performance

Hormone‐induced spawning of channel catfish held communally in tanks is a reliable method to produce channel catfish, Ictalurus punctatus ♀ × blue catfish, Ictalurus furcatus ♂, F₁ hybrid catfish fry. However, mature catfish are crowded, and repeatedly handled during the process of induced ovulation. Repeated handling of gravid females is stressful and may impair ovulation, egg quality, and reproductive performance. Three trials were conducted to evaluate the effects of two methods of confining post‐hormone‐injected female channel catfish on stress response (cortisol concentrations) and reproductive performance: fish were either held individually while suspended in soft, nylon‐mesh bags or communally in a concrete tank. Percent of females ovulated to hormone treatment, relative fecundity, percent egg viability, and latency of channel catfish did not differ for fish in the two treatments. However, percent hatch and fry/kg of females was higher (P < 0.05) for fish held in bags that for fish held communally in tanks. Mean plasma cortisol response immediately prior to the first hormone injection (0 h) did not differ among fish groups in the two treatments. However, mean plasma cortisol concentrations were significantly lower (P < 0.05) for fish in the bag treatment at 16 and 36 h compared to fish held communally in tanks. Plasma estradiol levels (measure of oocyte maturation) were assessed at 0, 16, and 36 h after hormone injection; concentrations were (P < 0.05) higher at 16 h compared to 0 and 36 h; however, estradiol concentrations did not differ for fish held in the two treatments (P > 0.05). Suspending hormone‐injected broodfish individually in soft bags reduced stress response, improved egg hatching rate, and increased hybrid fry produced per kg weight of female broodfish. Using this simple technology, farmers can improve the efficiency of hatcheries producing hybrid catfish fry.     

Larval dispersal and population connectivity among a network of marine reserves

A major challenge in marine ecology is describing patterns of larval dispersal and population connectivity, as well as their underlying processes. We used a biophysical model to simulate dispersal of eastern oyster, Crassostrea virginica, larvae and connectivity among a network of 10 no‐take reserves in a shallow, wind‐driven estuary to assess the relative importance of spawning location, spawning date, larval behavior, larval mortality, and adult reproductive output to predicted dispersal and connectivity patterns. The location (i.e., natal reserve) and date of spawning relative to physical processes, particularly frequency of wind reversals, were the dominant drivers of dispersal and connectivity patterns. To a lesser extent, larval behavior (i.e., 3D vertical advection and ontogenetic depth regulation) and mortality modified dispersal and connectivity, whereas spatiotemporal variability in adult reproductive output was of minimal importance. Over a 21‐day larval duration, mean dispersal distance of passive surface particles ranged from 5 to 40 km. Reserves were too small (1 km²) relative to mean dispersal distances to promote extensive local retention (median 2%) and spaced too far apart (typically ~50 km) to promote extensive inter‐reserve connectivity (median 2%). Limited connectivity and local retention may preclude the network from being self‐sustainable, thereby limiting its long‐term conservation and management benefits. In reserve systems characterized by limited connectivity, management efforts should focus on increasing connectivity by increasing the number or size of reserves to realize the benefits of improved adult demographics within reserves.     

The settlement and reproductive success of Lepeophtheirus salmonis (Krøyer 1837; Copepoda: Caligidae) on atypical hosts

The reproductive success of Lepeophtheirus salmonis settled on host and non‐host fish has been compared. Triplicate single species tanks of Atlantic salmon, marine three‐spined sticklebacks, saithe and Atlantic cod were exposed to 10 adult female L. salmonis per tank (n=30 lice per species). Adult female L. salmonis settlement and egg string production occurred only on salmon and cod, with no egg production occurring on saithe and three‐spined sticklebacks. The number of eggs in egg strings, hatching success of eggs and the survival of all larval stages to the copepodid stage were severely affected by the species of fish on which female L. salmonis had settled. L. salmonis settled on cod produced significantly fewer eggs, lower hatching rates and lower survival rates of larvae than females on Atlantic salmon. The production of egg strings by L. salmonis females infecting cod, which successfully hatch and moult through to the infective copepodid stage, albeit in small numbers, is discussed in terms of the implications to aquaculture and salmon and cod farming scenarios.     

The growth of larval cod and haddock in the Irish Sea: a model with temperature,prey size and turbulence forcing

We applied a physiological individual‐based model for the foraging and growth of cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae, using observed temperature and prey fields data from the Irish Sea, collected during the 2006 spawning season. We used the model to estimate larval growth and survival and explore the different productivities of the cod and haddock stocks encountered in the Irish Sea. The larvae of both species showed similar responses to changes in environmental conditions (temperature, wind, prey availability, daylight hours) and better survival was predicted in the western Irish Sea, covering the spawning ground for haddock and about half of that for cod. Larval growth was predicted to be mostly prey‐limited, but exploration of stock recruitment data suggests that other factors are important to ensure successful recruitment. We suggest that the presence of a cyclonic gyre in the western Irish Sea, influencing the retention and/or dispersal of larvae from their spawning grounds, and the increasing abundance of clupeids adding predatory pressure on the eggs and larvae; both may play a key role. These two processes deserve more attention if we want to understand the mechanisms behind the recruitment of cod and haddock in the Irish Sea. For the ecosystem‐based management approach, there is a need to achieve a greater understanding of the interactions between species on the scale a fish stock is managed, and to work toward integrated fisheries management in particular when considering the effects of advection from spawning grounds and prey–predator reversal on the recovery of depleted stocks.