Spawning Stock Size Research Articles

The role of larval transport on recruitment dynamics of red mullet (Mullus barbatus) in the Central Mediterranean Sea

Recruitment success depends on external forcing mechanisms such as ocean currents that affect the transport of eggs and larvae to favorable habitats. In this study, we investigated the role of larval transport in the recruitment of Mullus barbatus in the Central Mediterranean Sea by modeling the recruits' abundance as a function of both spawning stock size and dispersal rates of the species’ early life stages. Our analysis involved twenty years of data on recruits and spawners abundance obtained from scientific trawl surveys, and data on larval dispersal rates derived from a combination of actualized published sources and original data. By calculating the estimates of retention, import and uniformity of the contribution of the spawning areas distributed among different Geographical Sub Areas (GSAs) in the Sicilian nurseries, we assessed their contribution to recruitment using modified Ricker stock size-recruits models. In particular, our results show that a high uniform contribution from spawning areas within GSA16, mainly related to the oceanographic patterns promoting larval retention, together with spawners abundance, significantly reduced the variability of red mullet recruitment. We further highlighted that when switching from a higher to a lower level of evenness of contribution to the recruit population from different spawning areas in the GSA16, the expected spawning stock abundance per recruit for a given fishing pattern can suffer a rapid short-term decline, which is likely to have negative consequences for stock assessment and management decisions. Our results suggest that larval transport plays a crucial role in explaining the interannual variability of recruitment, thereby contributing to a better understanding of stock size variation. Additionally, our study enhances the understanding of the spatial dynamics involved in the recruitment of this species, which is of increasing interest within fisheries management frameworks.

From spawner habitat selection to stock-recruitment: Implications for assessment.

The relationship between the spawning stock size and subsequent number of recruits is a central concept in fisheries ecology. The influence of habitat selection of spawning individuals on the stock-recruitment relationship is poorly known. Here we explore how each of four different spawner behaviors might influence the stock-recruitment relationship and estimates of its parameters in the two most commonly used stock-recruitment functions (Beverton-Holt and Ricker). Using simulated stock-recruitment data generated by four different spawner behaviors applied to multiple discrete habitats, we show that when spawners were distributed proportionally to local carrying capacities, there was small or no bias in estimated recruitment and stock-recruitment parameters. For an ideal free distribution of spawners, larger bias in the estimates of recruitment and stock-recruitment parameters was obtained, whereas a random and a stepwise spawner behavior introduced the largest bias. Using stock-recruitment data corresponding to a "realistic" range of population densities and adding measurement error (20%-60%) to the simulated stock-recruitment data generated larger variation in the estimation bias than what was introduced by the spawner behavior. Thus, for exploited stocks at low population density and where spawning stock size and recruitment cannot be observed perfectly, partial observation of the possible spawner abundance range and measurement error might be of higher concern for management.

Strength and consistency of density dependence in marine fish productivity

AbstractThe correct prediction of the shape and strength of density dependence in productivity is key to predicting future stock development and providing the best possible long‐term fisheries management advice. Here, we identify unbiased estimators of the relationship between somatic growth, recruitment and density, and apply these to 80 stocks in the Northeast Atlantic. The analyses revealed density‐dependent recruitment in 68% of the stocks. Excluding pelagic stocks exhibiting significant trends in spawning stock biomass, the probability of significant density dependence was even higher at 78%. The relationships demonstrated that at the commonly used biomass limit of 0.2 times maximum spawning stock size, only 32% of the stocks attained three quarters of their maximum recruitment. This leaves 68% of the stocks with less than three quarters of their maximum recruitment at this biomass limit. Significantly lower recruitment at high stock size than at intermediate stock size was seen in 38% of the stocks. Density dependence in late growth occurred in 54% of the stocks, whereas early growth was generally density‐independent. Pelagic stocks were less likely to exhibit density dependence in recruitment than demersal and benthic stocks. We recommend that both the degree to which productivity is related to density and the degree to which the relationship changes over time should be investigated. Both of these aspects should be considered in evaluations of whether sustainability and yield can be improved by including density dependence in forecasts of the effects of different management actions.

Climate change or mismanagement?

Climate change and deoxygenation are affecting fish stocks on a global scale, but disentangling the impacts of these stressors from the effects of overfishing is a challenge. This study was conducted to distinguish between climate change and mismanagement as possible causes for the drastic decline in spawning stock size and reproductive success in cod (Gadus morhua) and herring (Clupea harengus) in the Western Baltic Sea, when compared with the good or satisfactory status and reproductive success of the other commercial species in the area. Available data on water temperature, wind speed, and plankton bloom during the spawning season did not reveal conclusive correlations between years with good and bad reproductive success of cod or herring. Notably, the other commercial species in the area have very similar life history traits suggesting similar resilience against stress caused by climate change or fishing. The study concludes that severe, sustained overfishing plus inappropriate size selectivity of the main fishing gears have caused the decline in spawning stock biomass of cod and herring to levels that are known to have a high probability of impaired reproductive success. It is pointed out that allowed catches were regulated by management and adhered to by the fishers, meaning that unregulated fishing did not occur. Thus, mismanagement (quotas that were too high and gears that selected too small sizes) and not climate change appears to be the primary cause of the bad status of cod and herring in the Western Baltic Sea.

Was historical cisco Coregonus artedi yield consistent with contemporary recruitment and abundance in Lake Superior?

AbstractHistorically, cisco Coregonus artedi Lesueur was the predominant prey fish and target of commercial fisheries throughout Lake Superior, but most spawning stocks collapsed by the mid‐1900s. Stocks partially recovered by the early 1990s, but contemporary abundance is considered to be below historical levels and driven by intermittent recruitment. Stochastic, age‐structured simulation models were used to determine whether historical (pre‐1955) cisco yield in Lake Superior was consistent with contemporary (1992–2015) abundance, life‐history characteristics and recruitment dynamics. When compared to contemporary stocks, the findings suggest historical stocks had: (1) similar recruits per spawner at low spawning stock sizes; (2) lower rates of compensatory density dependence; (3) similar or lower recruitment variation depending on the area and (4) higher median adult and age‐1 density. These findings are consistent with the hypothesis that eutrophication during the historical period supported greater recruitment and adult abundance and that re‐oligotrophication during the contemporary period may be limiting full recovery.

Rapid species level identification of fish eggs by proteome fingerprinting using MALDI-TOF MS

Quantifying spawning biomass of commercially relevant fish species is important to generate fishing quotas. This will mostly rely on the annual or daily production of fish eggs. However, these have to be identified precisely to species level to obtain a reliable estimate of offspring production of the different species. Because morphological identification can be very difficult, recent developments are heading towards application of molecular tools. Methods such as COI barcoding have long handling times and cause high costs for single specimen identifications. In order to test MALDI-TOF MS, a rapid and cost-effective alternative for species identification, we identified fish eggs using COI barcoding and used the same specimens to set up a MALDI-TOF MS reference library. This library, constructed from two different MALDI-TOF MS instruments, was then used to identify unknown eggs from a different sampling occasion. By using a line of evidence from hierarchical clustering and different supervised identification approaches we obtained concordant species identifications for 97.5% of the unknown fish eggs, proving MALDI-TOF MS a good tool for rapid species level identification of fish eggs. At the same time we point out the necessity of adjusting identification scores of supervised methods for identification to optimize identification success. SignificanceFish products are commercially highly important and many societies rely on them as a major food resource. Over many decades stocks of various relevant fish species have been reduced due to unregulated overfishing. Nowadays, to avoid overfishing and threatening of important fish species, fish stocks are regularly monitored. One component of this monitoring is the monitoring of spawning stock sizes. Whereas this is highly dependent on correct species identification of fish eggs, morphological identification is difficult because of lack of morphological features.

Egg production methods applied to Eastern Baltic cod provide indices of spawning stock dynamics

Egg production methods (EPM) provide fishery independent estimates of spawning stock sizes and dynamics of fish populations. Such methods are commonly used for short-lived pelagic species, less so for demersal fish such as cod. In this paper, we apply EPMs on cod in the eastern Baltic Sea, using a long time series of ichthyoplankton data. Stock assessment of Eastern Baltic cod has been challenged due to changing productivity of the stock invalidating some of the standard procedures, e.g. age determination and input variables, e.g. natural mortality. We demonstrate that EPMs, based on other data and assumptions than standard stock assessments, provide useful information on stock status and dynamics. We apply both the annual and daily egg production methods, which yielded similar results and were in line with stock trends derived from bottom trawl surveys. However, the EPM based spawning stock estimates were consistently lower compared to results from the latest analytical stock assessment. We identified processes introducing uncertainties in EPM estimates and their effects on the resulting estimates, and conclude that they mainly affect the absolute estimates but less the relative trends in stock dynamics. Therefore, we consider that EPMs are useful for providing relative indices for stock assessment purposes, with the Eastern Baltic cod being the first case where such indices are included in an official stock assessment of a demersal gadoid species. We also identify knowledge gaps in order to be able to derive absolute stock size estimates from EPMs in the future.

Oceanographic drivers of petrale sole recruitment in the California Current Ecosystem

AbstractThis paper investigates environmental drivers of U.S. West Coast petrale sole (Eopsetta jordani) recruitment as an initial step toward developing an environmental recruitment index that can inform the stock assessment in the absence of survey observations of age‐0 and age‐1 fish. First, a conceptual life history approach is used to generate life‐stage‐specific and spatio‐temporally specific mechanistic hypotheses regarding oceanographic variables that likely influence survival at each life stage. Seven life history stages are considered, from female spawner condition through benthic recruitment as observed in the Northwest Fisheries Science Center West Coast Groundfish Bottom Trawl Survey (age‐2 fish). The study area encompasses the region from 40 to 48°N in the California Current Ecosystem. Hypotheses are tested using output from a regional ocean reanalysis model outputs and model selection techniques. Four oceanographic variables explained 73% of the variation in recruitment not accounted for by estimates based exclusively on the spawning stock size. Recruitment deviations were (a) positively correlated with degree days during the female precondition period, (b) positively correlated with mixed‐layer depth during the egg stage, (c) negatively correlated with cross‐shelf transport during the larval stage, and (d) negatively correlated with cross‐shelf transport during the benthic juvenile stage. While multiple mechanisms likely affect petrale sole recruitment at different points during their life history, the strength of the relationship is promising for stock assessment and integrated ecosystem assessment applications.

Winter monsoon promotes the transport of Japanese temperate bass Lateolabrax japonicus eggs and larvae toward the innermost part of Tango Bay, the Sea of Japan

AbstractNorthwesterly cold winds characteristic of the East Asian Winter Monsoon (EAWM) dictate winter climatic conditions over the Japanese Archipelago. Japanese temperate bass Lateolabrax japonicus is a commercially important coastal fish that spawns offshore in winter and uses shallow waters as nursery habitats. To investigate the effects of EAWM on the planktonic period of L. japonicus, eggs, larvae, and juveniles were quantitatively collected in Tango Bay on the Sea of Japan side in winter and spring from 2007 to 2017. Although eggs occurred close to the mouth of the bay, planktonic larvae occurred further inside as they developed. The horizontal distribution of planktonic larvae, combined with water velocity data obtained from mooring observations, indicated that planktonic larvae are transported south‐ to westward through Ekman current and an anticyclonic circulation, which are driven by northwesterly winds. To evaluate survival during the planktonic period in each year class, the abundance of benthic larvae/juveniles was divided by winter total landings of Lateolabrax spp. (proxy of the spawning stock size). This survival index exhibited a positive correlation with the northwesterly component of winter winds, and a negative correlation with winter air temperature (average from December to February, Spearman's correlation, p < .05). There was, however, no significant correlation with winter water temperature or winter freshwater discharge in the bay. We conclude that northwesterly cold winds of EAWM play a critical role in transporting L. japonicus eggs and larvae toward nursery habitats, specifically beaches and estuaries fringing the innermost part of Tango Bay.

Effects of water temperature and pikeperch (Sander lucioperca) abundance on the stock\u2013recruitment relationship of Eurasian perch (Perca fluviatilis) in the northern Baltic Sea

How spawning stock size, environmental conditions and recruitment relate to each other is an essential question in understanding population dynamics of exploited fish stocks. We estimated the recruitment of Eurasian perch (Perca fluviatilis), one of the most important species in coastal fisheries in northern Baltic Sea, and examined the factors that determine perch recruitment success. We hypothesized that perch spawning population biomass and summer water temperature would increase perch recruitment, with potential density dependence, while the effect of the population size of pikeperch (Sander lucioperca) would be negative. Different forms of general stock–recruitment functions, with and without density dependence, and with and without water temperature and pikeperch population size as environmental factors were fitted to long-term (1981–2014) stock assessment data of perch and pikeperch in the Archipelago Sea, southwestern coast of Finland. Perch spawning stock biomass (ages 5–14), water temperature in June–July and pikeperch stock size (ages ≥ 1) at spawning year best explained variation in perch recruitment. The results supported the predictions: perch recruitment increased with spawning stock in density-dependent manner, pikeperch effect on perch recruitment was negative and summer temperature effect was positive suggesting environmentally affected competitive interaction between these two percids.

Genetic Stock Identification Reveals That Angler Harvest Is Representative of Cryptic Stock Proportions in a High‐Profile Kokanee Fishery

AbstractEstimating fishery harvest and spawning escapement (spawning stock size) are critical components of fisheries management; however, they can be particularly challenging to measure in systems where visually indistinguishable, but reproductively isolated populations mix within a single fishing area. Genetic stock identification is a common tool used in such mixed‐stock fisheries to improve estimates of spawning escapement and productivity; however, there are few references for management applications, particularly for inland recreational fisheries. The kokanee Oncorhynchus nerka population in Wood Lake, British Columbia, is a highly productive and valuable mixed‐stock fishery that includes two reproductively distinct ecotypes: shore‐ and stream‐spawning. Enumeration of shore‐spawning kokanee is logistically challenging, as the spawning population is not confined to a defined area or depth like stream‐spawners. Here, we combined in‐lake sampling (angler harvest and age‐0 trawl samples) over a 9‐year period (2008–2016) with genetic stock identification and Bayesian statistics to develop a new method for enumerating shore‐spawning kokanee. Our results suggest that angler‐harvested kokanee are representative of the spawner age structure and stock proportions. Therefore, we used the angler harvest sample combined with known stream‐spawner escapement to reconstruct the shore‐spawner escapement time series. Shore‐spawner abundance varied between 2,040 spawners and 13,460 spawners across years, which is over four times that previously predicted using the peak estimate of visual survey counts. Our results demonstrate the recovery of both the shore‐ and stream‐spawning kokanee in Wood Lake following a well‐documented crash in 2011 and suggest that a larger harvestable surplus is available for this high‐value kokanee fishery.

When does fishing forage species affect their predators?

This paper explores the impact of fishing low trophic level “forage” species on higher trophic level marine predators including other fish, birds and marine mammals. We show that existing analyses using trophic models have generally ignored a number of important factors including (1) the high level of natural variability of forage fish, (2) the weak relationship between forage fish spawning stock size and recruitment and the role of environmental productivity regimes, (3) the size distribution of forage fish, their predators and subsequent size selective predation (4) the changes in spatial distribution of the forage fish as it influences the reproductive success of predators. We show that taking account of these factors generally tends to make the impact of fishing forage fish on their predators less than estimated from trophic models. We also explore the empirical relationship between forage fish abundance and predator abundance for a range of U.S. fisheries and show that there is little evidence for a strong connection between forage fish abundance and the rate of change in the abundance of their predators. We suggest that any evaluation of harvest policies for forage fish needs to include these issues, and that models tailored for individual species and ecosystems are needed to guide fisheries management policy.

Population Structure of Adult Blue Crabs, Callinectes sapidus, in Relation to Physical Characteristics in Barnegat Bay, New Jersey

Blue crabs (Callinectes sapidus) are an important species in coastal or lagoonal estuaries where adult population characteristics may differ as compared to drowned-river estuaries. Barnegat Bay, in southern New Jersey, is composed of two large embayments: one without and one with a salinity gradient. We tested the influence of physical characteristics on the abundance, sex ratio, and size of adult blue crabs and examined variation in measures of reproductive potential (e.g., sperm stores) in both sexes in Barnegat Bay from June to September, 2008–2009. Population structure was distinct between the embayments due to sex-specific responses to salinity: male abundance was negatively correlated with salinity whereas adult females were more abundant in high salinity because of proximity to Barnegat Inlet. This produced high sex ratios in low salinity areas and low sex ratios in high salinity areas. Summer was a growing season for adult males while in late summer-early fall, juvenile males recruited to the adult size class. The spawning season lasted from May to August and ovigerous females were concentrated near the inlets. Information on female sperm stores and ovarian development identified two cohorts of adult females: females that will spawn in the current summer and females that will not spawn until the following summer. Thus, not all adult females near the spawning grounds were members of the current spawning stock. This suggests that annual estimates of spawning stock size which overlook the proximity of females to spawning are overestimating the current spawning stock in Barnegat Bay and other estuaries.

An explicit solution for calculating optimum spawning stock size from Ricker's stock recruitment model.

Stock-recruitment models have been used for decades in fisheries management as a means of formalizing the expected number of offspring that recruit to a fishery based on the number of parents. In particular, Ricker’s stock recruitment model is widely used due to its flexibility and ease with which the parameters can be estimated. After model fitting, the spawning stock size that produces the maximum sustainable yield (SMSY) to a fishery, and the harvest corresponding to it (UMSY), are two of the most common biological reference points of interest to fisheries managers. However, to date there has been no explicit solution for either reference point because of the transcendental nature of the equation needed to solve for them. Therefore, numerical or statistical approximations have been used for more than 30 years. Here I provide explicit formulae for calculating both SMSY and UMSY in terms of the productivity and density-dependent parameters of Ricker’s model.

Impact of the Three Gorges Dam on the spawning stock and natural reproduction of Chinese sturgeon in Changjiang River, China

Chinese sturgeon (Acipenser sinensis) is the flagship species of the Changjiang River. The migration route of this species is blocked by the first dam, the Gezhou Dam, and its reproduction is affected by the Three Gorges Dam (TGD), one of the largest dams in the world. We studied the impact of the impoundment of the Three Gorges Reservoir (TGR) since 2003 on the spawning stock and the natural reproduction of the Chinese sturgeon by using our monitoring data from 1997 to 2013. Results indicate that TGR impoundment has delayed the first spawning dates of the fish from middle-late October to late November, decreased the amount of spawning activities from twice to only once each year, and significantly reduced egg production. In particular, the fish did not demonstrate any spawning activities in 2013. Therefore, TGR impoundment significantly affects the natural reproduction of the fish downstream of the TGD. The spawning stock size of the fish is also predicted to further decrease in the future, which will lead to a risk of population extinction. Ecological regulations must be imposed on decreasing the water temperature to 20°C before mid-October and increasing water discharge downstream of the TGD in October to induce spawning of the Chinese sturgeon.

Changing recruitment capacity in global fish stocks

Marine fish and invertebrates are shifting their regional and global distributions in response to climate change, but it is unclear whether their productivity is being affected as well. Here we tested for time-varying trends in biological productivity parameters across 262 fish stocks of 127 species in 39 large marine ecosystems and high-seas areas (hereafter LMEs). This global meta-analysis revealed widespread changes in the relationship between spawning stock size and the production of juvenile offspring (recruitment), suggesting fundamental biological change in fish stock productivity at early life stages. Across regions, we estimate that average recruitment capacity has declined at a rate approximately equal to 3% of the historical maximum per decade. However, we observed large variability among stocks and regions; for example, highly negative trends in the North Atlantic contrast with more neutral patterns in the North Pacific. The extent of biological change in each LME was significantly related to observed changes in phytoplankton chlorophyll concentration and the intensity of historical overfishing in that ecosystem. We conclude that both environmental changes and chronic overfishing have already affected the productive capacity of many stocks at the recruitment stage of the life cycle. These results provide a baseline for ecosystem-based fisheries management and may help adjust expectations for future food production from the oceans.

The effects of stock size and environmental variability on Cape hake recruitment in Namibia: an unsolved puzzle (a comment on Kainge et al. 2013)

Measuring and forecasting recruitment are central to the understanding and management of fish stocks. Kainge et al. (2013) studied the effect of spawning stock size and environmental fluctuations on the recruitment levels of the Cape hake Merluccius capensis in Namibia. However, their study contains some flaws that undermine the conclusion that Cape hake recruitment is under the influence of upwelling in summer. Until those flaws are properly addressed, this conclusion, in our view, should be treated with caution.

Different maturity scales affect estimations of fecundity, TEP and spawning stock size of Greenland halibut, Reinhardtius hippoglossoides (Walbaum, 1792)

Based on 138 samples from female Northeast Arctic Greenland halibut (Reinhardtius hippoglossoides) taken in November–December 2011 at spawning grounds on the continental slope of the Barents Sea, the relationships between fecundity and fish size were established and found to be in the same range as those of estimates from 1996, 1997 and 1998. Ovarian maturity stages were determined by using a scale proposed by Kennedy et al. in 2011 based on microscopic oocyte diameter measurements. These data were compared to the maturity stages determined at sea in a routine manner, based on a standard macroscopic scale and a macroscopic scale previously developed for Greenland halibut. Maturity ogives were derived based on the three different maturity scales and an overestimation of both spawning stock size and total egg production (TEP) of approximately 20% was found when the macroscopic scales were used in the conventional way to determine maturity ogives. Most accurate ogives are assumed to be derived based on the microscopic scale, but this method can be impractical at sea on routine surveys. Thus, it is proposed to use the special macroscopic scale for Greenland halibut females, but consider both truly immature (stage 1) and early maturing (stage 2) females as non-spawning. This adjustment gives maturity ogives that do not deviate significantly from those based on the microscopic scale, and results in estimates of spawning stock size and TEP that we consider appropriate for stock assessment purposes.

A retrospective evaluation of the Barents Sea capelin management advice

Since 1998, the assessment model framework Bifrost/Captool has been used for advice on total allowable catch for the Barents Sea capelin (Mallotus villosus) stock. However, since the management is based on a target escapement strategy, and most capelin die after spawning, viewed in retrospect, there is hardly any possibility of checking whether the forecast underpinning the quota was actually realistic. The forecast using Captool for the period from 1 October up to capelin spawning time at 1 April relies on a forecast of cod abundance during this period. This estimate of cod abundance is, in turn, based on a cod assessment from the previous spring. By rerunning the Captool model, where the cod forecast is replaced with the actual amount of cod from the cod assessment model run later in time, we show that considerably smaller annual quotas would have been advised if the true amount of cod had been known when the capelin quota was set. We discuss this fact in light of the present knowledge about recruitment success of capelin during the period. Our conclusion is that either the acoustic stock size estimates of capelin have been consistently underestimating the true stock size, the predation model overestimates the natural mortality of capelin during winter, the minimum spawning stock size considered necessary to uphold normal recruitment (Blim) built into the harvest control rule is higher than needed to secure good recruitment, or a combination of these.

Using sound propagation modeling to estimate the number of calling fish in an aggregation from single-hydrophone sound recordings

Many fishes make sounds during spawning events that can be used to estimate abundance. Spawning stock size is a measure of fish population size that is used by fishery biologists to manage harvests levels. It is desirable that such an estimate be assessed easily and remotely using passive acoustics. Passive acoustics techniques (hydrophones) can be used to identify sound-producing species, but it is difficult to count individual sound sources in the sea, where it is dark, background noise levels can be high, but species can be identified by their sounds. We have developed a method that can estimate the density of calling fish in an aggregation from single-hydrophone recordings. Our method requires a sound propagation model for the area in which the aggregation is located. We generate a library of modeled sounds of virtual Monte-Carlo generated distributions of fish to determine the range of fish population densities that match the characteristics of single-hydrophone sound recording. Such a model could be used from a fixed station (e.g., an observatory) to estimate the population size of the sound producers. In this presentation, we will present some calculations made using this method and will examine the benefits and limitations of the technique.