Maturation Reaction Norms Research Articles

A multi-population approach supports common patterns in marine growth and maturation decision in Atlantic salmon (Salmo salar L.) from southern Europe.

This study provides a regional picture of long-term changes in Atlantic salmon growth at the southern edge of their distribution, using a multi-population approach spanning 49 years and five populations. We provide empirical evidence of salmon life history being influenced by a combination of common signals in the marine environment and population-specific signals. We identified an abrupt decline in growth from 1976 and a more recent decline after 2005. As these declines have also been recorded in northern European populations, our study significantly expands a pattern of declining marine growth to include southern European populations, thereby revealing a large-scale synchrony in marine growth patterns for almost five decades. Growth increments during their sea sojourn were characterized by distinct temporal dynamics. At a coarse temporal resolution, growth during the first winter at sea seemed to gradually improve over the study period. However, the analysis of finer seasonal growth patterns revealed ecological bottlenecks of salmon life histories at sea in time and space. Our study reinforces existing evidence of an impact of early marine growth on maturation decision, with small-sized individuals at the end of the first summer at sea being more likely to delay maturation. However, each population was characterized by a specific probabilistic maturation reaction norm, and a local component of growth at sea in which some populations have better growth in some years might further amplify differences in maturation rate. Differences between populations were smaller than those between sexes, suggesting that the sex-specific growth threshold for maturation is a well-conserved evolutionary phenomenon in salmon. Finally, our results illustrate that although most of the gain in length occurs during the first summer at sea, the temporal variability in body length at return is buffered against the decrease in post-smolt growth conditions. The intricate combination of growth over successive seasons, and its interplay with the maturation decision, could be regulating body length by maintaining diversity in early growth trajectories, life histories, and the composition of salmon populations.

Bioen-OSMOSE: A bioenergetic marine ecosystem model with physiological response to temperature and oxygen

1.Marine ecosystem models have been used to project the impacts of climate-induced changes in temperature and oxygen on biodiversity mainly through changes in species spatial distributions and primary production. However, fish populations may also respond to climatic pressures via physiological changes, leading to modifications in their life history that could either mitigate or worsen the consequences of climate change.2.Building on the individual-based multispecies ecosystem model OSMOSE, Bioen-OSMOSE has been developed to account for high trophic levels’ physiological responses to temperature and oxygen in future climate projections. This paper presents an overview of the Bioen-OSMOSE model, mainly detailing the new developments. These consist in the implementation of a bioenergetic sub-model that mechanistically describes somatic growth, sexual maturation and reproduction as they emerge from the energy fluxes sustained by food intake under the hypotheses of a biphasic growth model and plastic maturation age and size represented by a maturation reaction norm. These fluxes depend on temperature and oxygen concentration, thus allowing plastic physiological responses to climate change.3.To illustrate the capabilities of Bioen-OSMOSE to represent realistic ecosystem dynamics, the model is applied to the North Sea ecosystem. The model outputs are confronted with population biomass, catch, maturity ogive, mean size-at-age and diet data of each species of the fish community. The model succeeds in reproducing observations, with good performances for all indicators. A first exploration of current spatial variability in species’ bioenergetic fluxes resulting from temperature, oxygen, and food availability is presented in this paper, highlighting the role of temperature.4.This new model development opens the scope for new fields of research such as the exploration of seasonal or spatial variation in life history in response to biotic and abiotic factors at the individual, population and community levels. Understanding such variability is crucial to improve our knowledge on potential climate change impacts on marine ecosystems and to make more reliable projections under climate change scenarios.

Decades of warming alters maturation and reproductive investment in fish

AbstractHow does warming affect maturation and reproductive investment in ectotherms? Younger age and smaller size at maturation, as well as altered reproduction processes, have been found in a few species subjected to elevated temperatures. These observations, however, come from studies that do not distinguish effects of warming on maturation from those on growth, are also restricted to single generation responses to warming, or have additional stressors besides warming in the study system. Here, we study warming effects on maturation and reproductive investment in wild, unexploited fish populations using a whole‐ecosystem heating experiment. The experiment is conducted on Eurasian perch (Perca fluviatilis) in a heated and control area (with >5°C temperature difference) in the Baltic Sea. We compare female perch size at maturation using estimated probabilistic maturation reaction norms (PMRNs) and the gonado‐somatic index over 17 years of heating, spanning approximately five to eight perch generations. Using the PMRN approach, we show that warming has substantial effects on maturation size independent of warming‐induced changes in body growth. We found that young fish mature at a smaller size and invest more in developing their gonads in the heated population than in the unheated population. Our findings suggest that warming effects on reproductive investment may initially compensate for the cost of warming‐induced decrease in maturation size caused by the trade‐off between early maturation and size‐dependent fecundity. After multiple additional generations of warming, maturation and reproduction traits in perch differed from those in the first generations following the onset of warming, which suggests that warming‐induced evolution may have occurred. Our study is particularly relevant in the context of climate change because of the unusually large temperature difference between the areas and the fact that the heating occurred on an ecosystem level. We call for experimental studies resolving mechanisms of trait responses to warming across generations, complemented with genomic analyses, to aid understanding of organisms' long‐term responses to climate change.

Stock-specific variation in the probability of precocious male maturation in hatchery Chinook salmon (Oncorhynchus tshawytscha)

Age of maturation in many salmonid species is phenotypically plastic and dependent on exceeding a genetically set threshold in growth, often described as a probabilistic maturation reaction norm (PMRN). Hatchery supplementation programs for Chinook salmon (Oncorhynchus tshawytscha) in the Pacific Northwest US have been developed to minimize differences between hatchery and wild fish by integrating natural-origin adults into broodstock, potentially affecting PMRNs. We raised fish from 10 Chinook salmon stocks with variable levels of natural-origin integration in a common garden environment to explore potential genetic variation in PMRNs for precocious male maturation as age 2 minijacks. Proportion minijacks varied ≈10-fold (0.043 to 0.443) and the PMRN WP50 (predicted weight at 50% maturation) varied by ≈18 g (24.1 to 41.7 g). The propensity for minijack maturation was generally higher in stocks with higher levels of natural-origin integration. These findings demonstrate the effect of genotype by environment interactions on life history of salmonids and the need for stock-specific tailoring of rearing regimes to regulate differences between hatchery and wild fish, when wild fish are used in broodstocks.

Maturation reaction norm evolution under varying conditions of eco-evolutionary change

Probabilistic maturation reaction norms (PMRNs) are commonly used to infer evolution of maturation age and size in wild fish stocks, but how well estimates from phenotypic data actually reflect underlying genotypes is debated. We used an eco-genetic model of a commercially harvested freshwater fish to simulate populations undergoing various levels of fisheries-induced evolution and density-dependent feedback and evaluated effects on the estimation of PMRNs. We estimated PMRNs from phenotypic data sampled from simulated populations (age, length, and maturation status of individuals), as is done for wild stocks, and compared estimates with the known maturation genotypes of individuals in the simulated population. PMRN estimates were robust to changes in the strength of density-dependent growth and high levels of fisheries-induced evolution. However, our ability to detect slower rates of evolution was limited, especially when individuals matured within a narrow range of ages. Furthermore, low numbers of immature or mature individuals within a given age class limited our ability to robustly estimate midpoints for more than one or two ages, despite our large sample sizes (n = 200 randomly sampled individuals of each age per year), highlighting a challenge with this approach. Nonetheless, this study suggests that the widely applied method of estimating PMRNs from readily available phenotypic data to detect underlying evolution of maturation schedule is robust to some key factors that vary in wild populations.

Determinism of Temporal Variability in Size at Maturation of Sardine Sardina pilchardus in the Bay of Biscay

Age and size at maturation appear as key parameters governing the dynamics of a population as they affect growth rate, fecundity, and survival. The expression of such life history traits is determined by genetic make-up and modulated by environmental factors mainly through phenotypic plasticity. Moreover, fishing, besides decreasing population size and changing demographic composition can alter allelic frequencies through fisheries-induced evolution by selecting for some particular traits. In the Bay of Biscay, a decreasing trend in both sardine body condition and size-at-age has recently been pointed out at the population level. The Probabilistic Maturation Reaction Norm (PMRN) approach was applied to help disentangle phenotypic plasticity and genetic changes. Based on the analysis of sardine spawning seasonality, PMRN was estimated by considering body condition as additional life-history state variable to predict the onset of maturation. The resulting PMRN was then used to investigate temporal trends in reaction norm midpoints to test whether changes in length at maturation can be explained by plastic and/or evolutionary adaptive change. Overall, our results emphasize for the first time that including sardine body condition as explanatory variable improves predictions of maturation probability. We found that better individual condition increases maturation probability. The assessment of temporal changes in length at maturation confirms the low plasticity in this trait for a species maturing mostly at age-1 and advocates for the use of a monthly time scale when investigating PMRNs for this species. Beside environmental variables included in this analysis (water temperature, chlorophyll-a, and population biomass) that only show a weak correlation with PMRN midpoints, our results reveal no evidence for recent fisheries-induced evolution in the sardine stock of the Bay of Biscay. They suggest that the short-term variability in length at maturation is strongly dependent upon individual growth which is likely driven by environmental factors. For sardine fisheries management, our study highlights the need to consider both the length-composition data and the seasonality within a stock assessment model. Finally, we discuss the fact that considering individual growth trajectories should improve our understanding of the relationship between environmental variability and changes in maturation for sardine.

The geometry of reaction norms yields insights on classical fitness functions for Great Lakes salmon.

Life history theory examines how characteristics of organisms, such as age and size at maturity, may vary through natural selection as evolutionary responses that optimize fitness. Here we ask how predictions of age and size at maturity differ for the three classical fitness functions–intrinsic rate of natural increase r, net reproductive rate R0, and reproductive value Vx−for semelparous species. We show that different choices of fitness functions can lead to very different predictions of species behavior. In one’s efforts to understand an organism’s behavior and to develop effective conservation and management policies, the choice of fitness function matters. The central ingredient of our approach is the maturation reaction norm (MRN), which describes how optimal age and size at maturation vary with growth rate or mortality rate. We develop a practical geometric construction of MRNs that allows us to include different growth functions (linear growth and nonlinear von Bertalanffy growth in length) and develop two-dimensional MRNs useful for quantifying growth-mortality trade-offs. We relate our approach to Beverton-Holt life history invariants and to the Stearns-Koella categorization of MRNs. We conclude with a detailed discussion of life history parameters for Great Lakes Chinook Salmon and demonstrate that age and size at maturity are consistent with predictions using R0 (but not r or Vx) as the underlying fitness function.

Long-term changes in the maturation and growth of Cape Cod/Gulf of Maine yellowtail flounder Limanda ferruginea

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 633:169-180 (2020) - DOI: https://doi.org/10.3354/meps13175 Long-term changes in the maturation and growth of Cape Cod/Gulf of Maine yellowtail flounder Limanda ferruginea Gregory R. DeCelles1,*, Tiffany Vidal1,2 1Massachusetts Division of Marine Fisheries, Resource Assessment and Monitoring Program, New Bedford, MA 02744, USA 2The Pacific Community, BP D5, 98848 Noumea, New Caledonia *Corresponding author: gregory.decelles@mass.gov ABSTRACT: Yellowtail flounder Limanda ferruginea in the Gulf of Maine have been harvested by commercial fisheries since the 1930s, and the most recent stock assessment estimated that from 1985 to 2016 fishing mortality rates exceeded the fishing mortality reference point (F40%) in all but one year. The objective of this study was to understand changes in the maturation and growth of Cape Cod/Gulf of Maine yellowtail flounder by examining fisheries-independent data from cohorts produced over a 38 yr period (1976 to 2014). Maturity ogives demonstrated that the length and age at 50% maturity declined for cohorts throughout that time period. Probabilistic maturation reaction norms (PMRNs), which describe the probability that an immature individual will become mature at a certain age and size class, demonstrated a shift towards maturation at younger ages and smaller sizes. For age 3 female and male yellowtail flounder, the size at which the probability of becoming mature was 0.5 declined from 30.8 and 26.0 cm to 23.9 and 16.4 cm, respectively. Using von Bertalanffy growth curves, a decrease in asymptotic length (L×) and an increase in the growth rate (K) was estimated for both sexes. Taken together, these life history changes are suggestive of fisheries-induced evolution. However, although the PMRN approach is designed to account for phenotypic plasticity associated with growth and survival, further work is needed to determine whether other factors (e.g. temperature and body condition) could be responsible for the changes in maturation observed in this study. KEY WORDS: PMRN · Limanda ferruginea · Fisheries-induced evolution · Maturation · Von Bertalanffy Full text in pdf format PreviousNextCite this article as: DeCelles GR, Vidal T (2020) Long-term changes in the maturation and growth of Cape Cod/Gulf of Maine yellowtail flounder Limanda ferruginea. Mar Ecol Prog Ser 633:169-180. https://doi.org/10.3354/meps13175 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 633. Online publication date: January 09, 2020 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2020 Inter-Research.

Maintaining a wild phenotype in a conservation hatchery program for Chinook salmon: The effect of managed breeding on early male maturation.

In many salmonid species, age and size at maturation is plastic and influenced by the interaction between genetic and environmental factors. Hatchery reared salmon often mature at an earlier age and smaller size than wild fish. Modern salmon conservation efforts have focused on managing the level of gene flow between hatchery and natural origin fish to minimize potential genotypic and phenotypic change. In salmonids, maturation probability is dependent on exceeding a genetically set threshold in growth rate and energetic status (and by association, body size) referred to as the probabalisitic maturation reaction norm (PMRN). Over fourteen years, we monitored the frequency of age-2 precocious male maturation (common term: age-2 minijack rate) and the PMRN of natural founder (FNDR), integrated natural-hatchery (INT), and segregated hatchery (SEG) broodlines of spring Chinook salmon, Oncorhynchus tshawytscha. The average age-2 minijack rate (± SEM) of the FNDR, INT and SEG broodlines was 48.2 ± 5.2%, 41.9 ± 3.6% and 30.9 ± 4.7%, respectively. Additionally, the PMRN WP50 (predicted weight at 50% maturation) of the SEG broodline was significantly greater (20.5 g) than that of the FNDR/INT broodlines (18.2 g). We also conducted a common garden experiment exploring the effects of less than one [INT (0–1)], one [SEG (1)] or two [SEG (2)] generations of hatchery culture on the age-2 minijack rate and PMRN WP50. Growth was not significantly different among broodlines, but age-2 minijack rates were significantly lower following two consecutive generations of hatchery culture: [INT (0–1): 68.3 ± 1.7%], [SEG (1): 70.3 ± 1.8%] and [SEG (2): 58.6 ± 0.4%] and the PMRN WP50 was significantly higher by 6.1 g after two generations of SEG culture. These results indicate that managed gene flow reduces phenotypic divergence, but may serve to maintain potentially undesirably high age-2 minijack rates in salmon conservation hatchery programs.

Population density and temperature correlate with long-term trends in somatic growth rates and maturation schedules of herring and sprat.

We examine long-term trends in the average growth rates and maturation schedules of herring and sprat populations using survey data collected from the North Sea and west of Scotland since the 1960s and 1980s respectively. Otolith age data and maturity data are used to calculate time series of mean lengths at age, von Bertalanffy growth parameters, and probabilistic maturation reaction norms. As the growth and maturation of fish is known to be influenced by temperature and stock abundances, we account for these variables using Generalised Additive Models. Each of the herring populations displayed either steady declines in mean length across multiple age groups, or declines in length followed years later by some recovery. Depending on region, lengths at age of sprat increased or decreased over time. Varying temporal trends in maturation propensity at age and length were observed across herring populations. Many of the trends in growth rate and maturation were correlated to population abundance and/or temperature. In general, abundance is shown to be negatively correlated to growth rates in herring and sprat, and positively correlated with maturation propensity in herring. Temperature is also shown to be correlated to growth and maturation, and although the effect is consistent within species, the temperature effects differ between herring and sprat. This study provides detailed information about long-term trends in growth and maturation, which is lacking for some of these pelagic stocks, especially in the west of Scotland.

Reaction norm analysis reveals rapid shifts toward delayed maturation in harvested Lake Erie yellow perch (Perca flavescens).

Harvested marine fish stocks often show a rapid and substantial decline in the age and size at maturation. Such changes can arise from multiple processes including fisheries‐induced evolution, phenotypic plasticity, and responses to environmental factors other than harvest. The relative importance of these processes could differ systematically between marine and freshwater systems. We tested for temporal shifts in the mean and within‐cohort variability of age‐ and size‐based maturation probabilities of female yellow perch (Perca flavescens Mitchill) from four management units (MUs) in Lake Erie. Lake Erie yellow perch have been commercially harvested for more than a century, and age and size at maturation have varied since sampling began in the 1980s. Our analysis compared probabilistic maturation reaction norms (PMRNs) for cohorts when abundance was lower and harvest higher (1993–1998) to cohorts when abundance was higher and harvest lower (2005–2010). PMRNs have been used in previous studies to detect signs of evolutionary change in response to harvest. Maturation size threshold increased between the early and late cohorts, and the increases were statistically significant for the youngest age in the western MU1 and for older ages in the eastern MU3. Maturation envelope widths, a measure of the variability in maturation among individuals in a cohort, also increased between early and late cohorts in the western MUs where harvest was highest. The highest rates of change in size at maturation for a given age were as large or larger than rates reported for harvested marine fishes where declines in age and size at maturation have been observed. Contrary to the general observation of earlier maturation evolving in harvested stocks, female yellow perch in Lake Erie may be rapidly evolving delayed maturation since harvest was relaxed in the late 1990s, providing a rare example of possible evolutionary recovery.

Ecological change alters the evolutionary response to harvest in a freshwater fish.

Harvesting can induce rapid evolution in animal populations, yet the role of ecological change in buffering or enhancing that response is poorly understood. Here, we developed an eco-genetic model to examine how ecological changes brought about by two notorious invasive species, zebra and quagga mussels, influence harvest-induced evolution and resilience in a freshwater fish. Our study focused on lake whitefish (Coregonus clupeaformis) in the Laurentian Great Lakes, where the species supports valuable commercial and subsistence fisheries, and where the invasion of dreissenid (zebra and quagga) mussels caused drastic shifts in ecosystem productivity. Using our model system, we predicted faster rates of evolution of maturation reaction norms in lake whitefish under pre-invasion ecosystem conditions when growth and recruitment of young to the population were high. Slower growth rates that occurred under post-invasion conditions delayed when fish became vulnerable to the fishery, thus decreasing selection pressure and lessening the evolutionary response to harvest. Fishing with gill nets and traps nets generally selected for early maturation at small sizes, except when fishing at low levels with small mesh gill nets under pre-invasion conditions; in this latter case, evolution of delayed maturation was predicted. Overall, the invasion of dreissenid mussels lessened the evolutionary response to harvest, while also reducing the productivity and commercial yield potential of the stock. These results demonstrate how ecological conditions shape evolutionary outcomes and how invasive species can have a direct effect on evolutionary responses to harvest and sustainability.

Changing maturation reaction norms and the effects of growth history in Alaskan Chinook salmon

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 595:187-202 (2018) - DOI: https://doi.org/10.3354/meps12564 Changing maturation reaction norms and the effects of growth history in Alaskan Chinook salmon Jared E. Siegel1,*, Milo D. Adkison2, Megan V. McPhee2 1South Fork Research, Inc., 44842 SE 145th St., North Bend, WA 98045, USA 2College of Fisheries and Ocean Sciences, University of Alaska Fairbanks, 17101 Point Lena Loop Road, Juneau, AK 99801, USA *Corresponding author: derajlegeis@gmail.com ABSTRACT: We explored the nature of declines in size and age at maturity in 2 populations of Chinook salmon Oncorhynchus tshawytscha in western Alaska, USA, using multidimensional probabilistic maturation reaction norms (PMRNs) accounting for growth history. Individual growth histories informed by retrospective scale analysis were used to construct PMRNs and to describe the relative influence of different life-history periods on age at maturity. Similar results were found in both populations. Models accounting for growth history uniformly outperformed size-at-age models, suggesting the importance of growth history for the determination of maturation. The second year of marine growth was found to have a disproportionate influence on the age at maturity in both sexes. Males tended to grow more than females in length during the second year at sea, possibly as a consequence of females storing more energy in preparation for the high cost of female gonad development. Finally, we found that growth thresholds for maturation have shown a long-term decline in both sexes. This suggests that declines in the average age at maturity of western Alaskan Chinook salmon may have been caused in part by adaptation to environmental or fisheries-induced selection. KEY WORDS: Oncorhynchus tshawytschka · Size at age · PMRN · Age at maturity · Life history · Plasticity · Evolution · Retrospective scale analysis Full text in pdf format Supplementary material PreviousNextCite this article as: Siegel JE, Adkison MD, McPhee MV (2018) Changing maturation reaction norms and the effects of growth history in Alaskan Chinook salmon. Mar Ecol Prog Ser 595:187-202. https://doi.org/10.3354/meps12564 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 595. Online publication date: May 14, 2018 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2018 Inter-Research.

Adaptive and plastic variation in growth and maturation of the cutlassfish Trichiurus japonicus in the subtropical Pacific Ocean

We investigate d va ria- t ion in growth and maturation in response to 1) spatial variation in climate and 2) exploitation of the subtropical cutlassfish Trichiurus japonicus , an important fishery species whose population ecology i s virtually unknown . Individuals of this cutlassfish species were sampled monthly at 2 primary landing sites: Kengfang, in northeast (NE) Taiwan, and Tsukuan, in southwest (SW ) Taiwan, during 2013–2015. Habitat temperatures were about 1–4°C lower at the NE site t han at the SW site, and the length at age of adult fish had a n inverse pattern with temperature (NE lengths > SW lengths at age). The probabilistic maturation reaction norms did not differ significantly between the 2 areas, but ages and lengths a t maturation were higher for males from NE than for males from SW. Differences in asymptotic lengths (NE>SW) and growth coefficients (NE<SW), tog eth- er with narrower length distribution at the SW site than at the NE site, indicate potentially different mortality between the areas (NE<SW). These results indicate plastic variation in maturation and potentially adaptive variation in growth for this species that are related to the differences in temperature and mortality between the areas, although reliable proxies for mortality are lacking. The observed life-history variation was consistent with the tem perature–size rule that organisms tend to grow faster, mature earlier, and reach smal ler asymptotic sizes at warmer temperatures and may indicate an adaptive divergence of cutlassfish populati

Sex-based trade-offs among growth, mortality, and maturation in Great Lakes yellow perch stocks

Trade-offs among growth, mortality, and reproduction form the basis of life history theory but may vary among populations owing to local ecological conditions. We examined life history trade-offs driving variation in maturation among 13 yellow perch (Perca flavescens) stocks in the Great Lakes using sex-specific age and length at 50% maturity (A50 and L50, respectively) and probabilistic maturation reaction norm midpoints (Lp50,a). Both sexes exhibited positive correlations between growth and mortality, and faster-growing stocks were mature at younger ages but larger sizes. Male and female A50 and L50 were positively correlated among stocks, but Lp50,a estimates were negatively correlated among stocks, indicating stocks that matured at large sizes for a given age in females matured at smaller age-specific sizes in males. Female Lp50,a estimates were negatively related to growth and mortality, while male Lp50,a estimates were positively related to growth. These results suggest that (i) sex-based life history trade-offs sometimes act to differentially structure maturation schedules in males and females and (ii) males may be less responsive to changes in mortality than females.

A deterministic eco-genetic model for the short-term evolution of exploited fish stocks

Eco-genetic models describe the contemporary evolution of quantitative genetic traits by integrating the key ecological aspects with the modes of inheritance. Because of the flexibility in incorporating physiological, ecological, and genetic detail, eco-genetic models are typically individual-based and stochastic. Here we propose a deterministic eco-genetic population model to study the evolution of maturation schedules in exploited fish stocks. It is the first deterministic model addressing the evolution of the probabilistic maturation reaction norm (PMRN)—the genetic trait recognized to control maturation in fish species. The PMRN is the probability of interseasonal maturation as a function of age and size of juvenile individuals. Being independent of the growth trajectory followed by the individual, it can be considered a non-plastic phenotype, under limited change in temperature and climate. We consider the continuous age and size structures of the fish stock and explicitly represent foraging on a single food resource and mating under 1:1 sex ratio. Inheritance follows a single-locus-two-alleles Mendelian rule to limit the number of genotypes. We adapt the standard escalator boxcar train method for the simulation of physiologically structured population models to the case of two-dimensional age distributions—mature individuals being also distributed according to the age at maturation. The model simulations qualitatively reproduce empirical trends, specifically the genotypic redistribution toward early-maturing types under increasing exploitation and the eventual collapse of the stock. Interestingly, increased effort targeted at large sizes seems to erode the genetic diversity, an effect missed by stochastic models that might serve as warning to fishery managers and policy makers.

Fish life history dynamics: shifts in prey size structure evoke shifts in predator maturation traits

A bioenergetic framework is developed to predict optimal life history responses to environmentally driven changes in the rate of energy production by a predator. This framework is used to predict the responses of age at maturation, size at maturation, and asymptotic size to changes in the predator–prey size ratio. Predators feeding on relatively smaller prey (i.e., having larger predator–prey size ratios) have lower growth efficiency and are predicted as a consequence to mature earlier, at smaller sizes, and reach smaller asymptotic sizes. This prediction was tested using a 78-year time series (1936–2013) of data from a natural population of lake trout (Salvelinus namaycush) in Lake Opeongo, Algonquin Park, Ontario, Canada. A large decrease in the predator–prey size ratio for this population occurred over the period 1950–1965 when a preferred prey (cisco, Coregonus artedii) was introduced to the lake. This decrease was followed by ∼20 years of constancy in the size ratio and then 25 years of progressive increase. Lake trout life history responded plastically during both periods and consistently with our predictions. Extensive analysis of available data provided little empirical support for alternative explanations for the observed changes in lake trout size and maturity (e.g., changes in cisco and (or) lake trout density and harvest rates). The framework developed here derives plastic life history changes from fixed developmental thresholds that are based on the scaling of net production with body size and can be used to predict the shape of maturation reaction norms for the major shifts in community structure that are compactly summarized by changes in size spectrum parameters.

Reconstructing the effects of fishing on life-history evolution in North Sea plaice Pleuronectes platessa

Growing evidence suggests that fishing may induce rapid contemporary evolution in certain life-history traits. This study analyzes fisheries-induced changes in life-history traits describing growth, maturation, and reproduction, using an individual-based eco-genetic model that captures both the population dynamics and changes in genetic trait values. The model was successfully calibrated to match the observed life-history traits of female North Sea plaice 'Pleuronectes platessa' around the years 1900 and 2000. On this basis, we report the fllowing findings. First, the model indicates changes in 3 evolving life-history traits: the intercept of the maturation reaction norm decreases by 27%, the weight-specific reproductive-investment rate increases by 10%, and the weight-specific energy-acquisition rate increases by 1%. Together, these changes reduce the weight at maturation by 46% and the asymptotic body weight by 28% relative to the intensification of fishing around 1900. Second, while the maturation reaction norm and reproductive-investment rate change monotonically over time, the energy-acquisition rate follows a more complex course: after an initial increase during the first 50 yr, it remains constant for about 30 yr and then starts to decline. Third, our analysis indicates that North Sea plaice has not yet attained a new evolutionary equilibrium: it must be expected to evolve further towards earlier maturation, increased reproductive investment, and lower adult body size. Fourth, when fishing continues in our model 100 yr into the future, the pace of evolution slows down for the maturation reaction norm and the rate of energy acquisition, whereas no such slowing down is expected for the rate of reproductive investment.

Rapidly shifting maturation schedules following reduced commercial harvest in a freshwater fish.

Size-selective harvest of fish stocks can lead to maturation at smaller sizes and younger ages, which may depress stock productivity and recovery. Such changes in maturation may be very slow to reverse, even following complete fisheries closures. We evaluated temporal trends in maturation of five Great Lakes stocks of yellow perch (Perca flavescens Mitchill) using indices that attempt to disentangle plastic and evolutionary changes in maturation: age at 50% maturity and probabilistic maturation reaction norms (PMRNs). Four populations were fished commercially throughout the time series, while the Lake Michigan fishery was closed following a stock collapse. We documented rapid increases in PMRNs of the Lake Michigan stock coincident with the commercial fishery closure. Saginaw Bay and Lake Huron PMRNs also increased following reduced harvest, while Lake Erie populations were continuously fished and showed little change. The rapid response of maturation may have been enhanced by the short generation time of yellow perch and potential gene flow between northern and southern Lake Michigan, in addition to potential reverse adaptation following the fishing moratorium. These results suggest that some fish stocks may retain the ability to recover from fisheries-induced life history shifts following fishing moratoria.

Non-random sampling for reproductive status induces bias in probabilistic maturation reaction norm midpoints

The midpoint of the probabilistic maturation reaction norm (PMRN), which is the age-specific size at which the probability of maturing is 50%, has been used as an index of age and size at maturation of fish populations. The most important data requirement for the PMRN method is random sampling for reproductive status (mature or immature) at each age or size class. When age at first maturation varies between sexes, randomness of sampling for reproductive status can be evaluated by the sex ratio at the age of first maturity of the precocious sex (i.e., if the sex ratio at the age of first maturity of the precocious sex [sex of which age at first maturation is younger than the other] is biased, the randomness of sampling for reproductive status is not ensured). We examined the effect of the randomness of sampling for reproductive status on PMRN midpoints of precocious males in 10 tributaries containing two salmonid species. The sex ratio (the proportion of males) of fish at the age of first maturity varied among tributaries. The PMRN midpoints for males decreased with an increase in the sex ratio in both species. The biased sex ratio was thought to be caused by different movement patterns of precocious mature males and immature males/females. Our results emphasize that non-random sampling for reproductive status greatly affects estimated PMRN midpoint values.