Stock–environment–recruitment models for North Atlantic albacore (Thunnus alalunga)

Different stock–recruitment models were fitted to North Atlantic albacore (Thunnus alalunga) recruitment and spawning stock biomass data. A classical density dependence hypothesis, a recent environmental‐dependence hypothesis and a combination of both were considered. For the latter case, four stock–environment–recruitment models were used: Ricker, Beverton‐Holt, Deriso's General Model (modified to take into account environmental effects) and conditioned Neural Networks. Cross‐validation analysis showed that the modified Deriso model had the best predictive capability. It detected an inverse effect of the North Atlantic Oscillation (NAO) on recruitment, a Ricker‐type behaviour with density dependent overcompensation when environmental conditions are unfavourable and a Beverton–Holt‐type behaviour towards an asymptotic recruitment carrying capacity with favourable environmental conditions. The Neural Network model also detected that under favourable environmental conditions high spawning stock biomass does not necessarily have a depensatory effect on recruitment. Moreover, they suggest that under extremely favourable environmental conditions, albacore recruitment could increase well above the asymptotic carrying capacity predicted by Beverton–Holt‐type models. However, the general decrease in spawning stock biomass in recent years and increasing NAO trends suggest that there is low probability of exceptionally large recruitment in the future and instead there is a danger of recruitment overfishing.     

Predicting recruitment density dependence and intrinsic growth rate for all fishes worldwide using a data‐integrated life‐history model

Fisheries scientists use biological models to determine sustainable fishing rates and forecast future dynamics. These models require both life‐history parameters (mortality, maturity, growth) and stock‐recruit parameters (juvenile production). However, there has been little research to simultaneously predict life‐history and stock‐recruit parameters. I develop the first data‐integrated life‐history model, which extends a simple model of evolutionary dynamics to field measurements of life‐history parameters as well as historical records of spawning output and subsequent recruitment. This evolutionary model predicts recruitment productivity (steepness) and variability (variance and autocorrelation in recruitment deviations) as well as mortality, maturity, growth, and size, and uses these to predict intrinsic growth rate (r) for all described fishes. The model confirms previous analysis showing little correlation between steepness and either natural mortality or asymptotic maximum size (). However, it does reveal taxonomic patterns, where family Sebastidae has lower steepness () and Salmonidae has elevated steepness () relative to the prediction for bony fishes (class Actinopterygii, ). Similarly, genus Sebastes has growth rate (0.09) approaching that of several shark families (Lamniformes: 0.02; Carcharhiniformes: 0.02). A cross‐validation experiment confirms that the model is accurate, explains a substantial portion of variance (32%–67%), but generates standard errors that are somewhat too small. Predictive intervals are tighter for species than for higher‐level organizations (e.g. families), and predictions (including intervals) are available for all fishes worldwide in R package FishLife. I conclude by outlining how multivariate predictions of life‐history and stock‐recruit parameters could be useful for stock assessment, decision theory, ensemble modelling and strategic management.     

Life‐history variation in a species complex of nonmigratory galaxiids

Life‐history theory predicts that the optimal strategy in the trade‐off between egg size and number varies in relation to resource availability and environmental disturbance. We assessed interspecific differences in egg size, fecundity and other life‐history traits in a species complex of stream‐resident galaxiid fish, which are distributed across a range of contrasting habitat types on the South Island, New Zealand. Oocyte size, fecundity and reproductive effort were measured from gravid females collected immediately before spawning. Proxy measures of stream productivity, flow variability and predation pressure were extracted from modelled data sets. A suite of different egg sizes were identified across species within the Galaxias vulgaris complex, with mean oocyte volume differing by up to 133% between species. The species with the smallest eggs showed mean size‐relative fecundities 246% higher than the species with the largest eggs. A significant negative relationship was found between species’ mean egg size and size‐relative fecundity, suggesting a trade‐off between these traits. Species with larger eggs had larger maternal body size, lower reproductive effort and delayed maturity compared to ‘small‐egg’ species. Consistent with the predictions of life‐history theory, species with larger eggs, lower size‐relative fecundity, lower reproductive effort and delayed maturity were associated with low productivity, stable streams, whereas species exhibiting the opposite set of traits occurred in relatively productive but disturbed systems.     

Discriminating alternative stock–recruitment models and evaluating uncertainty in model structure

Fish stock–recruitment (S–R) assessment is one of the most essential keystones for fisheries management. Yet the analysis involves a variety of uncertainties. Amidst these difficulties, uncertainty in model structure is perhaps the most problematical to investigate because no rigorous statistical techniques can be used to explore the fundamental biological processes in S–R relationships. In this paper, I used computer simulations to investigate: (1) the differences between the estimated parameters of alternative S–R models as a function of stock characteristics: population growth rate, data range, fishing mortality, and process noise; and (2) the probability of selecting a correct model using information criteria. Two popular S–R functions, the Ricker and the Beverton–Holt models, were used as examples. Time series data were generated from a known S–R model and fitted by alternative models. The results show that when the two models fit the data similarly well, significant differences in parameters existed between the alternative models. The Ricker model tended to underestimate the population growth rate (initial slope) and the carrying capacity parameter, whereas the Beverton–Holt model overestimated these parameters. The management quantities (e.g., optimal virgin stock size) produced by one model were more conservative (i.e., larger optimal stock size or lower optimal harvest rate) under some conditions but became less conservative under other conditions. The differences between the alternative models were functions of the population growth rate, long-term fishing mortality, and data range of the stock size. The correct and incorrect models were statistically indistinguishable. For typical fishery data the probability of selecting the correct model based on information criteria was approximately 0.70 for the Ricker model and 0.61 for the Beverton–Holt model.     

Stock assessment of Japanese eels using Japanese abundance indices

The paper compiles a catch history of Japanese eels Anguilla japonica in East Asia and some Japanese relative abundance series. Maximum likelihood estimates of stock abundance of eels have been obtained using the abundance series and various biological parameters, such as growth, maturity and natural mortality. Age- and sex-structured models have been used to express the dynamics of stock abundance, and the Beverton and Holt model has been used to express the relationship between stock and recruitment. Data for estimations are standardized catch per unit effort of commercial fishery for exploitable stock (1954–2006 and 1968–2008) and for glass eel (1954–2010, 1972–2004, and 1973–1997). From the results of the base case scenario of estimations, the estimated stock size of individuals aged ≥1 year was 18.7 thousand tons in 2010, which was 24 % of the carrying capacity. The estimated stock size has recovered since 1990. Maximum sustainable yield was 4,180 tons if only the exploitable stock were utilized, and 266 tons if only the glass eel were utilized. These results and issues relating to estimation and management for reducing the fishery impact on stock are discussed.     

Density‐ and size‐dependent mortality in fish early life stages

The importance of survival and growth variations early in life for population dynamics depends on the degrees of compensatory density dependence and size dependence in survival at later life stages. Quantifying density‐ and size‐dependent mortality at different juvenile stages is therefore important to understand and potentially predict the recruitment to the population. We applied a statistical state‐space modelling approach to analyse time series of abundance and mean body size of larval and juvenile fish. The focus was to identify the importance of abundance and body size for growth and survival through successive larval and juvenile age intervals, and to quantify how the dynamics propagate through the early life to influence recruitment. We thus identified both relevant ages and mechanisms (i.e. density dependence and size dependence in survival and growth) linking recruitment variability to early life dynamics. The analysis was conducted on six economically and ecologically important fish populations from cold temperate and sub‐arctic marine ecosystems. Our results underscore the importance of size for survival early in life. The comparative analysis suggests that size‐dependent mortality and density‐dependent growth frequently occur at a transition from pelagic to demersal habitats, which may be linked to competition for suitable habitat. The generality of this hypothesis warrants testing in future research.     

A common environment experiment reveals plastic and genetic contributions to the fast life‐history strategy of an invasive fish

Invasive species often exhibit a suite of life‐history traits that promote rapid population growth, including early age and small size at maturation, and high reproductive investment. The common expression of these “fast” life‐history traits in invasive populations could be the result of plastic and/or genetic responses to the non‐native environment, or in response to the process of range expansion. To determine the relative importance of plastic and genetic contributions to the expression of life‐history traits, we reared two native Canadian and two invasive Spanish populations of Pumpkinseed sunfish (Lepomis gibbosus) in a common environment in central Ontario, Canada. In the wild, European Pumpkinseed tend to exhibit faster juvenile growth rates, younger age and smaller size at maturity, and higher reproductive investment than native North American populations. When reared in a common environment, both native and invasive populations exhibited similar juvenile growth rates, and similar age and size at maturity, suggesting that the differences seen among wild populations are a plastic response to the warmer non‐native environment. However, reproductive investment was consistently higher in the Spanish populations regardless of rearing environment, suggesting a genetic difference in reproductive investment between native and invasive populations. Selection for greater reproductive investment in non‐native Pumpkinseed may have contributed to their widespread success in Europe.     

Rethinking length‐based fisheries regulations: the value of protecting old and large fish with harvest slots

Managing fisheries using length‐based harvest regulations is common, but such policies often create trade‐offs among conservation (e.g. maintaining natural age‐structure or spawning stock biomass) and fishery objectives (e.g. maximizing yield or harvest numbers). By focusing harvest on the larger (older) fish, minimum‐length limits are thought to maximize biomass yield, but at the potential cost of severe age and size truncation at high fishing mortality. Harvest‐slot‐length limits (harvest slots) restrict harvest to intermediate lengths (ages), which may contribute to maintaining high harvest numbers and a more natural age‐structure. However, an evaluation of minimum‐length limits vs. harvest slots for jointly meeting fisheries and conservation objectives across a range of fish life‐history strategies is currently lacking. We present a general age‐ and size‐structured population model calibrated to several recreationally important fish species. Harvest slots and minimum‐length limits were both effective at compromising between yield, numbers harvested and catch of trophy fish while conserving reproductive biomass. However, harvest slots consistently produced greater numbers of fish harvested and greater catches of trophy fish while conserving reproductive biomass and a more natural population age‐structure. Additionally, harvest slots resulted in less waste in the presence of hooking mortality. Our results held across a range of exploitation rates, life‐history strategies and fisheries objectives. Overall, we found harvest slots to represent a valuable option to meet both conservation and recreational fisheries objectives. Given the ubiquitous benefits of harvest slots across all life histories modelled, rethinking the widespread use of minimum‐length limits is warranted.     

Intrinsic and extrinsic drivers of life‐history variability for a south‐western cutthroat trout

The impacts of climate change on cold‐water fishes will likely negatively manifest in populations at the trailing edge of their distributions. Rio Grande cutthroat trout (Oncorhynchus clarkii virginalis, RGCT) occupy arid south‐western U.S. streams at the southern‐most edge of all cutthroat trout distributions, making RGCT particularly vulnerable to the anticipated warming and drying in this region. We hypothesised that RGCT possess a portfolio of life‐history traits that aid in their persistence within streams of varying temperature and stream drying conditions. We used otolith and multistate capture–mark–recapture data to determine how these environmental constraints influence life‐history trait expression (length‐ and age‐at‐maturity) and demography in RGCT populations from northern New Mexico, United States. We found evidence that RGCT reached maturity fastest at sites with warm stream temperatures and low densities. We did not find a strong relationship between discharge and any demographic rate, although apparent survival of mature RGCT decreased as stream temperature increased. Our study suggests plasticity in trait expression may be a life‐history characteristic which can assist trailing edge populations like RGCT persist in a changing climate.     

Intra‐population variability of life‐history traits and growth during range expansion of the invasive round goby,Neogobius melanostomus

Abstract Fish can undergo changes in their life‐history traits that correspond with local demographic conditions. Under range expansion, a population of non‐native fish might then be expected to exhibit a suite of life‐history traits that differ between the edge and the centre of the population’s geographic range. To test this hypothesis, life‐history traits of an expanding population of round goby, Neogobius melanostomus (Pallas), in early and newly established sites in the Trent River (Ontario, Canada) were compared in 2007 and 2008. Round goby in the area of first introduction exhibited a significant decrease in age at maturity, increased length at age 1 and they increased in GSI from 2007 to 2008. While individuals at the edges of the range exhibited traits that promote population growth under low intraspecific density, yearly variability in life‐history traits suggests that additional processes such as declining density and fluctuating food availability are influencing the reproductive strategy and growth of round goby during an invasion.     

Reproductive ecology and scientific inference of steepness: a fundamental metric of population dynamics and strategic fisheries management

The relationship between the biomass of reproductively mature individuals (spawning stock) and the resulting offspring added to the population (recruitment), the stock–recruitment relationship, is a fundamental and challenging problem in all of population biology. The steepness of this relationship is commonly defined as the fraction of recruitment from an unfished population obtained when the spawning stock biomass is 20% of its unfished level. Since its introduction about 20 years ago, steepness has become widely used in fishery management, where it is usually treated as a statistical quantity. Here, we investigate the reproductive ecology of steepness, using both unstructured and age-structured models. We show that if one has sufficient information to construct a density-independent population model (maximum per capita productivity and natural mortality for the unstructured case or maximum per capita productivity, natural mortality and schedules of size and maturity at age for the structured model) then one can construct a point estimate for steepness. Thus, steepness cannot be chosen arbitrarily. If one assumes that the survival of recruited individuals fluctuates within populations, it is possible, by considering the early life history, to construct a prior distribution for steepness from this same demographic information. We develop the ideas for both compensatory (Beverton–Holt) and over-compensatory (Ricker) stock–recruitment relationships. We illustrate our ideas with an example concerning bluefin tuna ( Thunnus thynnus/orientalis , Scombridae). We show that assuming that steepness is unity when recruitment is considered to be environmentally driven is not biologically consistent, is inconsistent with a precautionary approach, and leads to the wrong scientific inference (which also applies for assigning steepness any other single value).     

Early life‐history adaptation influences conservation approaches for facultatively amphidromous fish

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Length-based Beverton and Holt spawning stock biomass per-recruit-models, with application to the Lates stappersii (Boulenger) stock in Lake Tanganyika

Two kinds of length-based Beverton and Holt spawning stock biomass per-recruit ( SSB/R )-models are presented. The first allows the computation of the absolute value of SSB/R . The second estimates the index of SSB/R , also called 'relative SSB/R '. Both models are extensions of the original age-based Beverton and Holt SSB/R -models. The SSB/R and its index can be assessed with reference to variations in the exploitation characteristics. These models were applied to the Lates stappersii (Boulenger) stock in Lake Tanganyika using fishing mortality as the independent variable. They produce an absolute value of SSB/R of 12.95 g, representing 22.4% of that of the 'virgin' stock for the 1993–1994 equilibrium-fishing regime. This level is indicative of an unsafe state of the stock in question. Moreover, the SSB for L. stappersii seems to be fragile under increased fishing pressure as, for example, doubling or tripling the current fishing regime would reduce the current SSB/R by 57.3 and 77.4%, respectively. In general, the use of length-based Beverton and Holt SSB/R -models is handicapped by underlying hypotheses and sub-models which are restrictive.     

Trends and management implications of human‐influenced life‐history changes in marine ectotherms

Evidence is accumulating that many marine ectotherms are undergoing rapid changes in their life‐history characteristics. These changes have been variously attributed to fisheries‐induced evolution, inhibited adult growth rate due to oxygen limitation at higher temperatures, and plastic responses to density dependence or changes in ocean productivity. Here, we review the diverse underlying mechanisms by which plastic and evolutionary responses to climate change and fisheries are likely to produce similar life‐history trends in harvested marine ectotherms, leading to faster life‐histories with earlier maturation and smaller adult size‐at‐age. While mechanistically understanding these growth and maturation changes may be difficult, it is becoming clear that changing life‐histories will lead to modified population dynamics, productivity and natural mortality of the affected species. We discuss how the observed and expected life‐history changes could affect the assumptions and uncertainty within single and multispecies models currently used in marine ecosystem management, highlighting that models which allow for dynamic life‐history traits often report significantly different estimates of stock biomass. Given that both climate‐ and harvest‐induced life‐history changes are likely to intensify and possibly amplify each other, there is an urgent need to adequately assess the implications of faster life‐histories for marine ecosystem management. This is especially true for data‐poor stocks, where growth and maturation are not regularly assessed. Targeted monitoring can be used to inform responsive management, but for improved sustainability outcomes, a precautionary approach to management that is robust to life‐history trends is advised.     

Assessment of winter size‐selective mortality of young‐of‐the‐year Atlantic salmon (Salmo salar) using otolith microstructure analysis*

Abstract – The objectives of this study were, first, to assess the usefulness of otolith microstructure analysis to examine winter size‐selective mortality of young‐of‐the‐year (YOY) Atlantic salmon and, secondly, to validate various hypotheses relating to the dynamics of two populations with different winter survival. By examining otolith microstructure, we back‐calculated body size at hatching and at emergence of YOY salmon sampled in fall 2000 and in early summer 2001 on the Petite Cascapédia River and the Bonaventure River (Québec, Canada). The results of the study did not reveal any size‐selective mortality of YOY salmon in the Petite Cascapédia River, while in the Bonaventure River, size‐selective mortality of the smaller individuals of the cohort was detected. This case study allowed not only a better comprehension of the population dynamics of those rivers but demonstrated the usefulness of otolith analysis to detect winter size‐selective mortality under a natural environment.     

Selectivity matters: Rules of thumb for management of plate‐sized,sex‐changing fish in the live reef food fish trade

Effective management of fisheries depends on the selectivity of different fishing methods, control of fishing effort and the life history and mating system of the target species. For sex‐changing species, it is unclear how the truncation of age‐structure or selection of specific size or age classes (by fishing for specific markets) affects population dynamics. We specifically address the consequences of plate‐sized selectivity, whereby submature, “plate‐sized” fish are preferred in the live reef food fish trade. We use an age‐structured model to investigate the decline and recovery of populations fished with three different selectivity scenarios (asymptotic, dome‐shaped and plate‐sized) applied to two sexual systems (female‐first hermaphroditism and gonochorism). We parameterized our model with life‐history data from Brown‐marbled grouper (Epinephelus fuscoguttatus) and Napoleon fish (Cheilinus undulatus). “Plate‐sized” selectivity had the greatest negative effect on population trajectories, assuming accumulated fishing effort across ages was equal, while the relative effect of fishing on biomass was greatest with low natural mortality. Fishing such sex‐changing species before maturation decreased egg production (and the spawning potential ratio) in two ways: average individual size decreased and, assuming plasticity, females became males at a smaller size. Somatic growth rate affected biomass if selectivity was based on size at age because in slow growers, a smaller proportion of total biomass was vulnerable to fishing. We recommend fisheries avoid taking individuals near their maturation age, regardless of mating system, unless catch is tightly controlled. We also discuss the implications of fishing post‐settlement individuals on population dynamics and offer practical management recommendations.     

Boom‐bust like dynamics of invasive black bullhead (Ameiurus melas) in Lake Sava (Serbia)

The life‐history traits and population dynamics of the black bullhead Ameiurus melas (Rafinesque) were studied monthly from August 2009 to December 2012 in Lake Sava in Belgrade (Serbia). This period included a mass mortality event in May 2011 and subsequent population recovery during 2012. Available evidence suggests the mass mortality was caused by European catfish virus that only affected the black bullhead. Changes in key life‐history traits and the recovery potential (increase in catch‐per‐unit‐effort, doubling of young‐of‐the‐year to adult ratio, earlier maturation and increased fecundity) following the mass mortality event suggest population resilience typical of a recurring boom‐bust pattern and demonstrate the species’ capacity to recover rapidly and re‐establish following disturbance. Repeated systematic mass removals of both juveniles and adults (using fyke nets) will be required if future fishery management measures aimed at limiting the population size are to be effective.     

Total population density during the first year of life as a major determinant of lifetime body‐length trajectory in marble trout

Abstract – The conditions experienced early in life can strongly influence life‐history trajectories in a variety of animal species. Here, we use data from four isolated populations of the endangered stream‐dwelling salmonid marble trout (Salmo marmoratus Cuvier 1817) living in the Soca and Idrijca river basins (Slovenia) to explore the influence of the total density experienced during and after the first year of life by marble trout year‐classes on body length of marble trout through the lifetime. Analyses were performed by pooling together the stream‐specific datasets to cover a wider range of densities. Mean body length of marble trout year‐classes through the lifetime (from age 1+ to 5+) was negatively related to total density of marble trout during the first year of life. The relationship between density during the first growth period and body length through the lifetime was well described by negative power curves. Total population density after the first year of life was not correlated with body length, thus suggesting that body growth trajectories are heavily determined early in life. Given size‐dependent sexual maturity and egg production in marble trout, the relationship between density early in life and lifetime individual growth may have strong implications in terms of population dynamics and regulation of population size.     

Predicting non‐native fish dispersal under conditions of climate change: case study in England of dispersal and establishment of pumpkinseed Lepomis gibbosus in a floodplain pond

Predictions of future climate change include shifts in patterns of precipitation, evapotranspiration and water run‐off, resulting in increased periods of drought as well as variability and intensity of rainfall events. In the United Kingdom, the non‐native North American sunfish, pumpkinseed Lepomis gibbosus (L.), is expected to benefit from these changes. We examine how hydrological variability induced by predicted changes in climate will affect the dispersal and spread of pumpkinseed in England by: (i) determining the relationship between discharge regime and pumpkinseed propagule pressure; (ii) examining a newly‐established pumpkinseed population following a flood event in 2007; and (iii) comparing the growth and life‐history traits of this new population with fish collected from the source population to demonstrate how the pumpkinseed's life‐history plasticity contributes to its success as a coloniser. Using Bayesian modelling, we determined that the number of pumpkinseed escapees is likely to increase with increasing discharge. The newly‐established pumpkinseed population showed fast juvenile growth, early age at maturity and small size at maturity. These traits differed significantly from the source population, specifically total length (TL) means at ages 1 and 2 were significantly greater in the new population, whereas TL at age 4 was significantly greater in the source population, and a significantly higher proportion of mature females were found at smaller size classes in the newly established pumpkinseed population. This study demonstrates the potential link between hydrological variability (current and future) and the dispersal of non‐native pumpkinseed, leading to the establishment of new populations.     

Understanding the life histories of amphidromous fish by integrating otolith‐derived growth reconstructions,post‐larval migrations and reproductive traits

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