Saving large fish through harvest slots outperforms the classical minimum‐length limit when the aim is to achieve multiple harvest and catch‐related fisheries objectives

We address the problem of optimal size‐selective exploitation in an age‐structured fish population model by systematically examining how density and size dependency in growth, mortality and fecundity affect optimal harvesting patterns when judged against a set of fisheries objectives. The study offers five key insights. First, while minimum‐length limits often maximize the biomass yield, exploitation using harvest slots (i.e. regulations that protect both immature and very large individuals) can generate within 95% of maximum yield; harvest slots also generally maximize the number of fish that are harvested. Second, density dependence in growth and size‐dependent mortality predict more liberal optimal size limits than those derived under assumptions of no density and size dependence. Third, strong density dependence in growth maximizes the catch of trophy fish only when modest harvest is introduced; the same holds for numbers harvested, when the stock–recruitment function follows the Ricker type. Fourth, the inclusion of size‐dependent maternal effects on fecundity or egg viability has only limited effects on optimal size limits, unless the increase in fecundity with mass (“hyperallometry”) is very large. However, large hyperallometry in fecundity shifts the optimal size limit for biomass yield from the traditional minimum‐length limit to a harvest slot. Fifth, harvest slots generally provide the best compromises among multiple objectives. We conclude that harvest slots, or more generally dome‐shaped selectivity to harvest, can outperform the standard minimum‐length selectivity. The exact configuration of optimal size limits crucially depends on objectives, local fishing pressure, the stock–recruitment function, and the density and size dependency of growth, mortality and fecundity.     

Long-term dynamics of pelagic fish density and vendace (Coregonus albula (L.)) stocks in four zones of a lake differing in trawling intensity

Abstract – An 11-year time series of hydroacoustic fish density estimates and fisheries statistics of vendace ( Coregonus albula (L.)) from four zones of a lake differing in trawling intensity was analyzed in order to test the hypothesis that intensive trawling has detrimental effects on pelagic fish stocks, especially vendace recruitment. The standardized fish density estimate in trawled zones showed no decrease in comparison to the non-trawled zone. No signs of recruitment failure associable with trawling intensity were found. The growth of vendace at the end of the study period was slower than that at the beginning, indicating a higher density, most clearly so in the zone with highest trawling intensity. No significant correlations were detected between 3-year mean trawling intensity and yield per unit effort of over-1-year-old vendace or fish density in any zone. Thus, no evidence to support the hypothesis was found. This was suggested to be due to density-dependent compensatory processes in recruitment and/or natural mortality effectively counteracting the population change induced by exploitation. ^Note     

Growth and viability of hatchery-reared Argyrosomus japonicus released into open and semi-closed systems

Abstract  Hatchery-reared Argyrosomus japonicus (Temminck & Schlegel) were released into estuaries of varying habitat between 1996 and 2004, and growth and recruitment to the fishery evaluated. Fish stocked earlier in summer had significantly faster growth rates ( P  < 0.01), and post-stocking mortality ranged from 1.1% to 11.7% within 7 day post-release. Argyrosomus japonicus stocked in the Georges River in 2003 and 2004 yielded fishery independent recapture rates up to 0.2%. For A. japonicus stocked in Smith's Lake in 1997 and 2004, growth rates, timing of recaptures and increases in commercial catches suggested recruitment to the fishery by 18 months. The 1997 stocking led to a 30-fold increase in A. japonicus catch, but stocking was not financially viable, with a cost:benefit ratio of 2.1. Small, shallow coastal lagoons may represent experimental units for refining stocking strategies, but are not optimal for A. japonicus . The potential for density-dependent effects, and complex relationships between growth, habitat and mortality, highlighted the need for a thorough understanding of species – system interactions. Pilot releases can contribute to this understanding, particularly assessment of habitat requirements, and season, site and size-of-release.     

Post-settlement growth and mortality of brackishwater clam Corbicula japonica in the Kiso estuaries, central Japan

Different cohorts of Corbicula japonica benthic individuals (from new settlers just after larval settlement up to recruits with 12-mm shell length and more) in the Kiso estuaries, Japan were separated based on temporal changes in shell length histograms. The post-settlement growth and mortality were examined for cohorts that were successful in fishery recruitment. Three growth models (Logistic, Gompertz and ALOG) were fitted to the growth data on average shell lengths of different cohorts, the best-fitting growth model varied depending on cohort and further on cohorts that were grouped according to estuary and to either overwintering or non-overwintering. Mortality rates were estimated based on regression models fitted separately to the data on temporal changes in the log-transformed density against sampling dates of individuals before and after fishery recruitment (i.e. attaining 12-mm shell length). For most cohorts, however, there was no significant difference in mortality rates between individuals before and after fishery recruitment, although significant differences were detected in initial densities between individuals of these two groups. It may be suggested that daily mortality rate after fishery recruitment was compensated with fishery mortality, so that total mortality rate was not different between individuals of these two groups.     

Cohort survival patterns of walleye pollock, Theragra chalcogramma, in Shelikof Strait, Alaska: a critical factor analysis

A series of age-specific life tables for walleye pollock ( Theragra chalcogramma ) in the western Gulf of Alaska was compiled for the 1980-91 year classes. The life tables were utilized to perform an exploratory key factor analysis to examine the timing of critical periods in the recruitment process, evidence of density-dependence at different stages and trends in mortality rates. Early larval mortality was significantly correlated with generational mortality (In recruits/spawning bio-mass), but patterns in juvenile mortality also were similar to generational mortality and in some years were clearly dominant in determining the fate of a cohort. Density-dependent mortality, based on the correlation between mortality and initial abundance, was indicated only for the late larval to early juvenile stage. Time trends were marginally significant for juvenile mortality. It is speculated that the observed increase in juvenile mortality is associated with increasing abundance of arrowtooth flounder. Weaknesses in the data base are discussed; these along with the short time series involved make our conclusions tentative and subject to further study. We hypothesize that pollock recruitment levels can be established at any life stage depending on sufficient supply from prior stages, a type of dynamics which can be termed supply dependent multiple life stage control.     

Effects of early life mass mortality events on fish populations

Mass mortality events are ubiquitous in nature and can be caused by, for example, diseases, extreme weather and human perturbations such as contamination. Despite being prevalent and rising globally, how mass mortality in early life causes population-level effects such as reduced total population biomass, is not fully explored. In particular for fish, mass mortality affecting early life may be dampened by compensatory density-dependent processes. However, due to large variations in year-class strength, potentially caused by density-independent variability in survival, the impact at the population level may be high in certain years. We quantify population-level impacts at two levels of mass mortality (50% and 99% additional mortality) during early life across 40 fish species using age-structured population dynamics models. The findings from these species-specific models are further supported by an analysis of detailed stock-specific models for three of the species. We find that population impacts are highly variable between years and species. Short-lived species that exhibit a low degree of compensatory density dependence and high interannual variation in survival experience the strongest impacts at the population level. These quantitative and general relationships allow predicting the range of potential impacts of mass mortality events on species based on their life history. This is critical considering that the frequency and severity of mass mortality events are increasing worldwide.     

Population trends of bycatch species reflect improving status of target species

Synthesis studies of fish stocks worldwide suggest improving status of mainly target species that are fully assessed. Other analyses, primarily based on catch data alone, but which include a wider range of species as well as bycatch, present a different view. Catch‐only analyses could be more robust if fishery‐independent data were used and discards accounted for. We develop a model that uses only survey biomass at length and landings data to estimate fishing mortality, spawning stock biomass (SSB) and discards. An analysis of species from the North Sea shows the model results compare well with most fully assessed stocks. When applied to bycatch species with limited data, trends in fishing mortality and SSB typically reflect those of the target species. In the last decade, mean fishing mortality rates have tended to decline, while mean SSB has increased. Despite increasing SSB, recent mean recruitment appears to have been lower than previously which may limit future biomass recovery. Species usually associated with more northerly distributions appear to show the greatest effect of weaker recruitment, which may be linked to climate. Estimated discards have tended to decline in magnitude as a result of reduced fishing mortality and associated lower total catches. The model offers a simple way to use both landings and survey data to obtain more detailed population trends for data limited species.     

Spawning success and early life history of longnose suckers in Great Lakes tributaries

Fish eggs and larvae are often subject to very high mortality, and variation in early life survival can be important for population dynamics. Although longnose suckers (Catostomus catostomus) are widespread in northern North America, little is known about their early life history. We examined fecundity and early larval survivorship during sucker spawning events in three small Lake Michigan tributaries. Although egg deposition varied 25% among spawning events, estimated larval export to the lake varied over 25,000‐fold from around 1000 to 26 million. Based on variation in environmental conditions across years, it appears that spring flow and temperature may be important determinants of egg survival to larval outmigration. Larval age data suggest that most individuals that survived to outmigration hatched during a 2‐day period despite adult spawning across at least 10 days. Most larvae spent <2 weeks in the stream and emigrated around the time of transition from endogenous to exogenous feeding before substantial growth occurred. In two of three cases, larvae drifted exclusively at night; however, high drift rates occurred during both day and night in the case where larvae were very abundant, suggesting density‐dependent drift behaviour. Our results indicate that survival in tributary streams from egg deposition to larval export is highly variable in longnose suckers. These large differences in early life survival may translate into variability in recruitment, thereby influencing population structure and dynamics.     

Ecologically and evolutionarily sustainable fishing of the pikeperch Sander lucioperca: Lake Oulujärvi as an example

Due to the multitude of participants and a diverse range of fishing gear used freshwater fisheries are often managed using minimum size limits (MSL) rather than regulations of total fishing effort. However, a concern has arisen whether attempts to improve ecological sustainability of fisheries by increasing MSLs would induce undesired adaptations to selective fishing. We examined the ecological and evolutionary impacts of varying fishing mortality rates under varying MSLs, with and without stockings, in an age-, size-, and maturity-structured evolutionary model which was parameterized for the Lake Oulujärvi pikeperch, Sander lucioperca. We found that at the current level of harvesting (fishing mortality rate, F = 0.7) and stockings (430 000 year⁻¹), and under the assumption of strongly density-dependent growth, the nation-wide MSL of 370 mm maximizes theoretical biomass yield in a deterministic model but does not prevent severe recruitment overfishing under further increased fishing pressures or stochasticity in recruitment success. The recently imposed, local MSL of 450 mm better ensures stable yields, and even increases them if individual growth is density-independent, but further increase of MSL to 500 mm would already reduce yield especially if there was discard mortality for undersized fish. Given density-dependent growth, equal survival between wild and stocked fish, and sustainable fishing mortality rate, stockings do not increase yield or significantly improve the stability of yields. Evolutionarily stable size at maturation decreases under strong fishing mortality, but increased MSLs reduce the magnitude of this undesired effect. Negatively size-dependent natural mortality was found to have a positive effect on the otherwise negative selection for length-at-age. Increased MSLs also reduce the total selection for decreased length-at-age. Our results support the intentions to increase MSLs in order to improve both ecological and evolutionary sustainability of recreational fisheries.     

Density-dependent growth of young-of-the-year Atlantic salmon (Salmo salar) revisited

Imre I, Grant JWA, Cunjak RA. Density-dependent growth of young-of-the-year Atlantic salmon ( Salmo salar ) revisited. Ecology of Freshwater Fish 2010: 19: 1–6. © 2009 John Wiley & Sons A/S
Abstract –  The length of individual young-of-the-year (YOY) Atlantic salmon ( Salmo salar ) in Catamaran Brook decreases with increasing population density following a negative power curve. Because most of this decrease in growth rate occurs at low densities (<1 fish·m⁻²), ( Imre et al. 2005 ; Journal of Animal Ecology, 74: 508–516) suggested that exploitation competition for drifting prey rather than space limitation might be responsible for this pattern. Recently, ( Ward et al. 2007 ; Journal of Animal Ecology, 76: 135–138) showed that the negative power curve of growth rate versus density can be caused by other mechanisms and suggested that Imre et al.'s evidence for density-dependent growth would have been stronger if we had analysed final size versus initial density rather than final density. We examined (i) whether the negative power curve of size versus density was also apparent in an analysis of final size versus initial density and tested two predictions that emerge from Ward et al.'s model, (ii) the variance in body size increases with population density, and (iii) the maximum fish size at a site is density-independent. The final size of YOY salmon decreased with increasing initial density following a negative power curve. Our data did not provide strong support for the above predictions emerging from Ward et al.'s model. Our analyses of different years, sites and seasons were consistent with the hypothesis of density-dependent growth of YOY salmon.     

Population dynamics of the brown shrimp Crangon crangon L., in shallow areas of the German Wadden Sea

Settlement, growth and mortality of the brown shrimp, Crangon Crangon L., were studied in the north Frisian Wadden Sea over a period of 19 months to fill existing gaps regarding the species' life cycle, and thereby to clarify points related to the fishery. Settlement of postlarval shrimp in the shallow areas occurred in three batches: in May; during summer; and an overwintering batch. Compared with an earlier model based on laboratory-derived data, a new growth model based on field data predicts similar growth rates for the earlier phase of the shrimp life cycle, but higher rates for the later stages of the cohorts. Mean growth rate of the different cohorts over the size range sampled was 0.23 ± 0.08 mm day^?1. Of the factors tested, only temperature significantly influenced growth rates. Preliminary estimates of instantaneous total mortality values for shrimp cohorts from these shallow areas ranged from 0.03 to 0.12 day^?1. Salinity explained most of the variation in mortality rates, but this most likely indicates an indirect effect of depth on emigration. The cumulative effects of growth rates estimated from the new model and lower losses through predation could imply a higher contribution of winter eggs to the autumn fishery. This, however, needs further evaluation to properly quantify the trade-offs between settlement and recruitment to the fishery, as well as the growth and mortality rates.     

Over-wintering growth and losses in a small population of the threespine stickleback, Gasterosteus aculeatus (L.), in mid-Wales

Abstract –  Mark-recapture was used to estimate the population abundance of threespine stickleback, Gasterosteus aculeatus , in a backwater of Afon Rheidol in mid-Wales in autumn and early spring, from 1984 to 1998. For the 0+ cohort, there was no evidence of density-dependent losses (mortality and emigration) or growth in the over-wintering period. No systematic size-dependent loss was detected. There was an inverse relationship between autumn abundance and the annual rate of change in abundance of the 0+ cohort.     

长江鲥鱼资源评估与管理

本文根据1973～1975年对长江鲥鱼调查的生物学资料以及1970～1985年的渔业统计资料,应用Ricker和Fox模式进行评析,评析的结果是: 1、如果开捕年龄t_(?)为1.9龄,其最佳捕捞死亡系数F为0.6。 2、根据对雌鲥单位补充量繁殖力的估算,当t_(?)=1.9时,F>0.5,资源将处在高危险区;而当t(?)>3.9时,下论F值多大,资源将不会处在高危险区。 3、最大持续产量MSY的269.4吨,其相应捕捞努力量f_(MSY)为12938流刺网网片数。 4、最适产量Y_(?)为262吨,其相应的捕捞努力量f_(?)为10111流刺网网片数。 5、当前(1985年)的状况是:t_(?)=1.9龄,F=2.003,总捕捞努力量已超过30000流刺网网片数。雌鲥单位补充量繁殖力的相对值低于2.61％。其资源量已从1974年的1862吨减少到1985年的39.8吨。显然,当前长江鲥鱼资源的利用已出现补充型捕捞进度,资源处在高危险区中的最危险点,资源已严重衰竭。为此,作者提出为恢复,保护和管理该珍贵渔业资源的八点建议。计算和绘图均用自编程序在电子计算机ABACUS—11和HP7475A X—Y绘图仪上进行。     

A modelling approach to evaluate potential management actions designed to increase growth of white perch in a high‐density population

Abstract A deterministic, age‐structured population model was developed to simulate potential management scenarios designed to increase growth of individuals (quantified by maximum length) in a hypothetical population of white perch, Morone americana (Gmelin). Four scenarios were developed that included non‐selective mortality of adult white perch, increased mortality of age groups most influential on population growth, increased age‐0 mortality and inhibiting recruitment after spawning. The greatest increase in maximum length occurred with non‐selective adult mortality when population biomass was reduced by 97%; lesser increases in maximum length were achieved with the other management scenarios. Populations returned to their original state after control efforts ceased for each scenario.     

Increasing connectivity of Great Lakes tributaries: Interspecific and intraspecific effects on resident brook trout and brown trout populations

Understanding resident fish population responses to restored connectivity would enhance decision-making on dam removal and fish passage. Since such evaluations are limited in the Great Lakes region of North America, we compared abundance, survival, and growth of resident brook trout and brown trout between sets of Michigan streams where populations were or were not interacting with salmonid species that might be present if connectivity existed. We analysed data from 34 electrofishing index sites to compare resident trout populations between streams without versus with Great Lakes access (and migratory Pacific salmonids), and brook trout populations in Great Lakes inaccessible (land-locked) streams where brown trout were present versus absent. Great Lakes accessibility effects on fish density became increasingly positive for older age groups of brown trout while generally negative for all age classes of brook trout. Brown trout had consistently negative effects on brook trout density in land-locked streams. Increased connectivity had significant effects on annual survival for only one of seven trout age classes modelled, while intraspecific density-dependent effects on survival were significant in six models. Significant intraspecific effects on resident trout growth occurred for seven of eleven age classes examined. Negative interspecific effects of Great Lakes access on resident trout growth were most noticeable for age-0 and age-1 resident trout, age classes that likely compete with juvenile Pacific salmonids. Our findings provide a more robust understanding of how Great Lakes connectivity affects resident trout populations, highlighting negative influences of brown trout on brook trout and intraspecific density-dependent effects.     

A comparison of no-take zones and traditional fishery management tools for managing site-attached species with a mixed larval pool

No‐take zones (NTZs) can generate higher larval production by sessile, sedentary and site‐attached species per unit area than in exploited areas, and may increase recruitment and yield compared to status quo management. Whilst NTZs may be considered an essential part of optimal management, few studies have specifically compared the effects of NTZs with those of correctly applied gear and effort controls. A yield‐per‐recruit (YPR) population model, based on the sedentary abalone Haliotis laevigata, was used to compare the effects of management by minimum landing size (MLS), effort limitation and NTZs, either singularly or in combination. Initially, a minimum basic YPR model was used. Three additional assumptions were sequentially added to the model to see if they affected conclusions drawn from the model. The additional assumptions were the inclusion of: (i) a length–fecundity relationship; (ii) an age‐dependent natural mortality function; and (iii) mortality of undersized individuals due to fishery operations. In the absence of undersized mortality caused by fishing, under virtually all conditions the population is best managed with a combination of MLS and effort control, without any NTZs. For simulations that included mortality of undersized individuals in the fished area, under nearly all circumstances NTZs were considered an essential part of optimal fishery management, and management incorporating NTZs greatly increased the sustainable yield that could be taken.     

The daily egg production method: recent advances, current applications and future challenges

More than 120 surveys over 25 years suggest that appropriate use of the daily egg production method (DEPM) provides unbiased but rather imprecise estimates of spawning biomass (coefficient of variation generally above 30%). Knowledge of species reproductive biology and early life history and a survey design adapted to local population dynamics are required to optimize DEPM performance in terms of bias, precision and cost. Clupeoid applications dominate worldwide (mainly for anchovies and sardines) and estimates are often used to tune indirect assessment models or to calibrate other fisheries‐independent methods. The method seems better adapted to the life history of anchovies than of sardines, leading to more precise estimates of anchovy spawning biomass and facilitating extensions of the method to estimate total biomass and numbers at age. The continuous underway fish egg sampler is often used as a secondary sampler in the ichthyoplankton survey of the DEPM to reduce the cost and improve the precision of egg production. Multinomial models were recently developed to analyse egg incubation data and used in a Bayesian procedure for egg ageing and delimitation of daily cohorts. These were incorporated in model‐based estimators to get spatially explicit estimates of egg production, daily fecundity parameters and spawning biomass that can improve the precision of DEPM. Uncertainty in daily fecundity estimation of clupeoids is mainly because of spawning fraction estimation by the post‐ovulatory follicle (POF) method. Exploration of recent histological and molecular techniques for POF characterization and laboratory experiments to test the effect of temperature on POF degradation can help to improve spawning fraction estimation. Available estimates of egg production and mortality, daily fecundity, spawning area and biomass from different populations, species and ecosystems are being used to improve the understanding of clupeoid spawning dynamics, their relation with ecosystem productivity and to build comprehensive population models. Finally, a counter‐intuitive finding of this review is that, although the DEPM is almost exclusively applied to clupeoids, recent evidence indicates that it may be easier and cheaper to use in other teleost families, including demersal species.     

The utility of visible implant (VI) tags for marking tropical river fish

In tropical fisheries where ageing fish is often difficult, tagging programmes offer an important means to estimate growth parameters and mortality rates. The majority of tags are large and attached externally via a puncture in the muscle. These characteristics may cause significant biological effects which bias parameter estimates. Visible implant (VI) tags offer a more benign means to individually mark fish. The utility of VI tags for marking five species of tropical river fish was examined. The results indicate that the tag and tagging operation had no influence on either growth or mortality, although tag retention was poor compared with other studies. For three of the five species, the only suitable tagging location was highly inconspicuous and therefore inappropriate for tagging programmes that rely upon fishermen to return tagged fish. Alternative tagging locations may improve retention rates and the visibility of the tag.     

Interannual variability in growth of larval and juvenile walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1983–91

Interannual variability in growth of larval walleye pollock Theragra chalcogramma was examined from 1983 to 1991 and of juveniles from 1985 to 1990. ANCOVA was used to assess differences in population growth rates, and an alternate method was developed to examine variations between annual length-at-age data and average 'expected' values over different age groupings. For larvae, the years 1986, 1987, 1989 and 1990 had higher than average length-at-age, and 1988 and 1991 had lower than average values. Relationships between growth and SST and larval density were not clear. A tentative relationship between copepod nauplii abundance and larval length-at-age was noted. The consequence of larval growth to larval mortality, late larval abundance or recruitment was not clear. We conclude that larval mortality rates are highly variable and tend to mask effects of moderate variability in growth on later abundance. For juveniles, 1987 had significantly lower than average length-at-age and 1988 had higher than average values. Although there are few years of data, they tend to support the importance of juvenile growth in the recruitment process. Conditions for the large 1988 year class are documented and discussed, including warm SST, calm winds, relatively low larval growth rates, low abundances of potential predators on larvae, low larval mortality rates, and high juvenile growth rates.     

Density dependence in total egg production per spawner for marine fish

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.