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The role of postmating sexual selection as a potential reproductive barrier in speciation is not well understood. Here, we studied the effects of sperm competition and cryptic female choice as putative postmating barriers in two lamprey ecotypes with a partial reproductive isolation. The European river lamprey Lampetra fluviatilis is anadromous and parasitic of other fish species, whereas the brook lamprey Lampetra planeri is freshwater resident and nonparasitic. We measured sperm traits in both ecotypes and designed sperm competition experiments to test the occurrence of cryptic female choice. We also performed sperm competition experiments either at equal semen volume or equal sperm number to investigate the role of sperm velocity on fertilization success. We observed distinct sperm traits between ecotypes with a higher sperm concentration and a lower sperm velocity for L. planeri compared with L. fluviatilis. The outcomes of sperm competition reflected these differences in sperm traits, and there was no evidence for cryptic female choice irrespective of female ecotype. At equal semen volume, L. planeri males had a higher fertilization success than L. fluviatilis and vice versa at equal sperm number. Our results demonstrate that different sperm traits between ecotypes can influence the male reproductive success and thus gene flow between L. planeri and L. fluviatilis. However, postmating prezygotic barriers are absent and thus cannot explain the partial reproductive isolation between ecotypes.

Gut microbiome diversity and functions are jointly shaped by the host’s genetic background and environmental conditions, but the consequences of this interaction are still unclear. Unravelling the effect of the interaction between evolution and environment on the gut microbiome is particularly relevant considering the unprecedented level of human-driven disruption on the ecological and evolutionary trajectories of species. Here, we aimed to evaluate whether size-selective mortality influences the gut microbiome of medaka (Oryzias latipes), how environment conditions modulate the effect of the genetic background of medaka on their microbiota, and the association between microbiome diversity and medaka fitness. To do so, we studied two lineages of medaka that were raised under antagonistic size-selective regimes for 10 generations (i.e. the largest or the smallest breeders were removed to mimic fishing-like or natural mortality). In pond mesocosms, the two lineages were subjected to contrasting population density and light intensity (i.e. used as a proxy of primary production, hence resource availability). We observed significant differences in the gut microbiome composition and richness between the two lines, and this effect was mediated by light intensity. Indeed, the bacterial richness of fishing-like medaka (small-breeder line) was reduced by 34% under low-light conditions compared to high-light conditions, while it remained unchanged in natural mortality-selected medaka (large-breeder line). However, the observed changes in bacterial richness did not correlate with changes in growth rate or body condition, possibly due to functional redundancy among the microbial taxa residing in the gut. Given the growing evidence about the gut microbiomes importance to host health, more in-depth studies are required to fully understand the role of the microbiome in size-selected organisms and the possible ecosystem-level consequences.

Gut microbiome diversity and functions are jointly shaped by the host’s genetic background and environmental conditions, but the consequences of this interaction are still unclear. Unravelling the effect of the interaction between evolution and environment on the gut microbiome is particularly relevant considering the unprecedented level of human-driven disruption on the ecological and evolutionary trajectories of species. Here, we aimed to evaluate whether size-selective mortality influences the gut microbiome of medaka (Oryzias latipes), how environment conditions modulate the effect of the genetic background of medaka on their microbiota, and the association between microbiome diversity and medaka fitness. To do so, we studied two lineages of medaka that were raised under antagonistic size-selective regimes for 10 generations (i.e. the largest or the smallest breeders were removed to mimic fishing-like or natural mortality). In pond mesocosms, the two lineages were subjected to contrasting population density and light intensity (i.e. used as a proxy of primary production, hence resource availability). We observed significant differences in the gut microbiome composition and richness between the two lines, and this effect was mediated by light intensity. Indeed, the bacterial richness of fishing-like medaka (small-breeder line) was reduced by 34% under low-light conditions compared to high-light conditions, while it remained unchanged in natural mortality-selected medaka (large-breeder line). However, the observed changes in bacterial richness did not correlate with changes in growth rate or body condition, possibly due to functional redundancy among the microbial taxa residing in the gut. Given the growing evidence about the gut microbiomes importance to host health, more in-depth studies are required to fully understand the role of the microbiome in size-selected organisms and the possible ecosystem-level consequences.

The role of postmating sexual selection as a potential reproductive barrier in speciation is not well understood. Here, we studied the effects of sperm competition and cryptic female choice as putative postmating barriers in two lamprey ecotypes with a partial reproductive isolation. The European river lamprey Lampetra fluviatilis is anadromous and parasitic of other fish species, whereas the brook lamprey Lampetra planeri is freshwater resident and nonparasitic. We measured sperm traits in both ecotypes and designed sperm competition experiments to test the occurrence of cryptic female choice. We also performed sperm competition experiments either at equal semen volume or equal sperm number to investigate the role of sperm velocity on fertilization success. We observed distinct sperm traits between ecotypes with a higher sperm concentration and a lower sperm velocity for L. planeri compared with L. fluviatilis. The outcomes of sperm competition reflected these differences in sperm traits, and there was no evidence for cryptic female choice irrespective of female ecotype. At equal semen volume, L. planeri males had a higher fertilization success than L. fluviatilis and vice versa at equal sperm number. Our results demonstrate that different sperm traits between ecotypes can influence the male reproductive success and thus gene flow between L. planeri and L. fluviatilis. However, postmating prezygotic barriers are absent and thus cannot explain the partial reproductive isolation between ecotypes.

Evaluating the effects of climate variation on ecosystems is of paramount importance for our ability to forecast and mitigate the consequences of global change. However, the ways in which complex food webs respond to climate variations remain poorly understood. Here, we use long-term time series to investigate the effects of temperature variation on the intraguild-predation (IGP) system of Windermere (UK), a lake where pike (Esox lucius, top predator) feed on small-sized perch (Perca fluviatilis) but compete with large-sized perch for the same food sources. Spectral analyses of time series reveal that pike recruitment dynamics are temperature controlled. In 1976, expansion of a size-truncating perch pathogen into the lake severely impacted large perch and favoured pike as the IGP-dominant species. This pathogen-induced regime shift to a pike-dominated IGP apparently triggered a temperature-controlled trophic cascade passing through pike down to dissolved nutrients. In simple food chains, warming is predicted to strengthen top–down control by accelerating metabolic rates in ectothermic consumers, while pathogens of top consumers are predicted to dampen this top–down control. In contrast, the local IGP structure in Windermere made warming and pathogens synergistic in their top–down effects on ecosystem functioning. More generally, our results point to top predators as major mediators of community response to global change, and show that size-selective agents (e.g. pathogens, fishers or hunters) may change the topological architecture of food webs and alter whole ecosystem sensitivity to climate variation.

Evaluating the effects of climate variation on ecosystems is of paramount importance for our ability to forecast and mitigate the consequences of global change. However, the ways in which complex food webs respond to climate variations remain poorly understood. Here, we use long-term time series to investigate the effects of temperature variation on the intraguild-predation (IGP) system of Windermere (UK), a lake where pike (Esox lucius, top predator) feed on small-sized perch (Perca fluviatilis) but compete with large-sized perch for the same food sources. Spectral analyses of time series reveal that pike recruitment dynamics are temperature controlled. In 1976, expansion of a size-truncating perch pathogen into the lake severely impacted large perch and favoured pike as the IGP-dominant species. This pathogen-induced regime shift to a pike-dominated IGP apparently triggered a temperature-controlled trophic cascade passing through pike down to dissolved nutrients. In simple food chains, warming is predicted to strengthen top–down control by accelerating metabolic rates in ectothermic consumers, while pathogens of top consumers are predicted to dampen this top–down control. In contrast, the local IGP structure in Windermere made warming and pathogens synergistic in their top–down effects on ecosystem functioning. More generally, our results point to top predators as major mediators of community response to global change, and show that size-selective agents (e.g. pathogens, fishers or hunters) may change the topological architecture of food webs and alter whole ecosystem sensitivity to climate variation.

Predicted universal responses of ectotherms to climate warming include increased maximum population growth rate and changes in body size through the temperature-size rule. However, the mechanisms that would underlie these predicted responses are not clear. Many studies have focused on proximate mechanisms of physiological processes affecting individual growth. One can also consider ultimate mechanisms involving adaptive explanations by evaluating temperature effects on different vital rates across the life history and using the information in a population dynamical model. Here, we combine long-term data for a top predator in freshwater ecosystems (pike; Esox lucius) with a stochastic integral projection model to analyze concurrent effects of temperature on vital rates, body size, and population dynamics. As predicted, the net effect of warming on population growth rate (fitness) is positive, but the thermal sensitivity of this rate is highly size- and vital rate–dependent. These results are not sensitive to increasing variability in temperature. Somatic growth follows the temperature-size rule, and our results support an adaptive explanation for this response. The stable length structure of the population shifts with warming toward an increased proportion of medium-sized but a reduced proportion of small and large individuals. This study highlights how demographic approaches can help reveal complex underlying mechanisms for population responses to warming.

Intraspecific variation in body size is common in animals and plants. Body size affects trophic interactions like foraging ability and vulnerability to predation, which in turn affect individual fitness as well as population stability and extinction risk. Experimental and theoretical work has shown that the size distribution of individuals within cohorts is strongly influenced by intraspecific competition for resources, often leading to skewed frequency distributions. However, little is known about the effects of environmental factors such as climate and eutrophication on the cohort size-structure of natural populations. We use a long-term time series of scientific monitoring of a freshwater fish (European perch Perca fluviatilis) to investigate the effects of density dependence, predation, nutrient availability, climate and the timing of spawning on the cohort size distributions. We find that the mean length of the fish is best predicted by the extrinsic factors phosphorus concentration and summer temperature, and the densities of the different age-classes, whereas the skewness of the length distribution is best predicted by phosphorus concentration, summer temperature, abundance of small fish, and the timing of spawning. Higher nutrient levels, temperatures and densities of small fish increase food availability and thus reduce competition, which is reflected in increased mean length and decreased skewness. The timing of spawning affects skewness presumably through changes in the initial size variation of the cohort and the length of the first growth season. Our results indicate that higher temperatures increase the mean length and decrease skewness due to the concurrent eutrophication of the lake. The study thereby highlights the potential impact of human-induced environmental change on the size structure of fish populations. More studies are needed to understand better the complex mechanisms through which these factors alter the intensity of intraspecific competi

Intraspecific variation in body size is common in animals and plants. Body size affects trophic interactions like foraging ability and vulnerability to predation, which in turn affect individual fitness as well as population stability and extinction risk. Experimental and theoretical work has shown that the size distribution of individuals within cohorts is strongly influenced by intraspecific competition for resources, often leading to skewed frequency distributions. However, little is known about the effects of environmental factors such as climate and eutrophication on the cohort size-structure of natural populations. We use a long-term time series of scientific monitoring of a freshwater fish (European perch Perca fluviatilis) to investigate the effects of density dependence, predation, nutrient availability, climate and the timing of spawning on the cohort size distributions. We find that the mean length of the fish is best predicted by the extrinsic factors phosphorus concentration and summer temperature, and the densities of the different age-classes, whereas the skewness of the length distribution is best predicted by phosphorus concentration, summer temperature, abundance of small fish, and the timing of spawning. Higher nutrient levels, temperatures and densities of small fish increase food availability and thus reduce competition, which is reflected in increased mean length and decreased skewness. The timing of spawning affects skewness presumably through changes in the initial size variation of the cohort and the length of the first growth season. Our results indicate that higher temperatures increase the mean length and decrease skewness due to the concurrent eutrophication of the lake. The study thereby highlights the potential impact of human-induced environmental change on the size structure of fish populations. More studies are needed to understand better the complex mechanisms through which these factors alter the intensity of intraspecific competi

Anthropogenic factors, including climate warming, are increasing the incidence and prevalence of infectious diseases worldwide. Infectious diseases caused by pathogenic parasites can have severe impacts on host survival, thereby altering the selection regime and inducing evolutionary responses in their hosts. Knowledge about such evolutionary consequences in natural populations is critical to mitigate potential ecological and economic effects. However, studies on pathogen-induced trait changes are scarce and the pace of evolutionary change is largely unknown, particularly in vertebrates. Here, we use a time series from long-term monitoring of perch to estimate temporal trends in the maturation schedule before and after a severe pathogen outbreak. We show that the disease induced a phenotypic change from a previously increasing to a decreasing size at maturation, the most important life-history transition in animals. Evolutionary rates imposed by the pathogen were high and comparable to those reported for populations exposed to intense human harvesting. Pathogens thus represent highly potent drivers of adaptive phenotypic evolution in vertebrates.

Recently developed theoretical models of stage-structured consumer–resource systems have shown that stage-specific biomass overcompensation can arise in response to increased mortality rates. We parameterized a stage-structured population model to simulate the effects of increased adult mortality caused by a pathogen outbreak in the perch (Perca fluviatilis) population of Windermere (UK) in 1976. The model predicts biomass overcompensation by juveniles in response to increased adult mortality due to a shift in food-dependent growth and reproduction rates. Considering cannibalism between life stages in the model reinforces this compensatory response due to the release from predation on juveniles at high mortality rates. These model predictions are matched by our analysis of a 60- year time series of scientific monitoring of Windermere perch, which shows that the pathogen outbreak induced a strong decrease in adult biomass and a corresponding increase in juvenile biomass. Age-specific adult fecundity and size at age were higher after than before the disease outbreak, suggesting that the pathogen-induced mortality released adult perch from competition, thereby increasing somatic and reproductive growth. Higher juvenile survival after the pathogen outbreak due to a release from cannibalism likely contributed to the observed biomass overcompensation. Our findings have general implications for predicting population- and community-level responses to increased size-selective mortality caused by exploitation or disease outbreaks.

We use six decades of catch-at-age data for perch (Perca fluviatilis) and pike (Esox lucius) in Windermere (UK) to estimate age- and sex-specific population sizes, natural mortalities and catchabilities in both species. Population sizes are estimated by fitting age structured population models to the catch-at-age data using standard maximum likelihood methods. We validate our methods using data simulations, and use our estimates of vital rates (natural mortality, recruitment and catahability) to address important aspects of fisheries biology. Our model indicates that strong fishery selection against male perch apparently triggered a population collapse, highlighting that sex-selective fisheries can be harmful even at a reasonable exploitation rate (here ≤30%). Recruitment (R) increased with the abundance of spawners (S) in both species, but it also responded to both abiotic and other biotic factors. In particular, increased predator (pike) abundance induced a change from compensation to depensation in the prey (perch) SR relationship, thus favouring the occurrence of an Allee effect. Our study provides reference points for the effective exploitation of pike and perch populations, and underscores the need for ecosystem-based harvesting management.

Recently developed theoretical models of stage-structured consumer–resource systems have shown that stage-specific biomass overcompensation can arise in response to increased mortality rates. We parameterized a stage-structured population model to simulate the effects of increased adult mortality caused by a pathogen outbreak in the perch (Perca fluviatilis) population of Windermere (UK) in 1976. The model predicts biomass overcompensation by juveniles in response to increased adult mortality due to a shift in food-dependent growth and reproduction rates. Considering cannibalism between life stages in the model reinforces this compensatory response due to the release from predation on juveniles at high mortality rates. These model predictions are matched by our analysis of a 60- year time series of scientific monitoring of Windermere perch, which shows that the pathogen outbreak induced a strong decrease in adult biomass and a corresponding increase in juvenile biomass. Age-specific adult fecundity and size at age were higher after than before the disease outbreak, suggesting that the pathogen-induced mortality released adult perch from competition, thereby increasing somatic and reproductive growth. Higher juvenile survival after the pathogen outbreak due to a release from cannibalism likely contributed to the observed biomass overcompensation. Our findings have general implications for predicting population- and community-level responses to increased size-selective mortality caused by exploitation or disease outbreaks.

Anthropogenic factors, including climate warming, are increasing the incidence and prevalence of infectious diseases worldwide. Infectious diseases caused by pathogenic parasites can have severe impacts on host survival, thereby altering the selection regime and inducing evolutionary responses in their hosts. Knowledge about such evolutionary consequences in natural populations is critical to mitigate potential ecological and economic effects. However, studies on pathogen-induced trait changes are scarce and the pace of evolutionary change is largely unknown, particularly in vertebrates. Here, we use a time series from long-term monitoring of perch to estimate temporal trends in the maturation schedule before and after a severe pathogen outbreak. We show that the disease induced a phenotypic change from a previously increasing to a decreasing size at maturation, the most important life-history transition in animals. Evolutionary rates imposed by the pathogen were high and comparable to those reported for populations exposed to intense human harvesting. Pathogens thus represent highly potent drivers of adaptive phenotypic evolution in vertebrates.

We use six decades of catch-at-age data for perch (Perca fluviatilis) and pike (Esox lucius) in Windermere (UK) to estimate age- and sex-specific population sizes, natural mortalities and catchabilities in both species. Population sizes are estimated by fitting age structured population models to the catch-at-age data using standard maximum likelihood methods. We validate our methods using data simulations, and use our estimates of vital rates (natural mortality, recruitment and catahability) to address important aspects of fisheries biology. Our model indicates that strong fishery selection against male perch apparently triggered a population collapse, highlighting that sex-selective fisheries can be harmful even at a reasonable exploitation rate (here ≤30%). Recruitment (R) increased with the abundance of spawners (S) in both species, but it also responded to both abiotic and other biotic factors. In particular, increased predator (pike) abundance induced a change from compensation to depensation in the prey (perch) SR relationship, thus favouring the occurrence of an Allee effect. Our study provides reference points for the effective exploitation of pike and perch populations, and underscores the need for ecosystem-based harvesting management.

Chronic social stress diverts energy away from growth, reproduction and immunity, and is thus a potential driver of population dynamics. However, the effects of social stress on demographic density dependence remain largely overlooked in ecological theory. Here we combine behavioural experiments, physiology and population modelling to show in a top predator (pike Esox lucius) that social stress alone may be a primary driver of demographic density dependence. Doubling pike density in experimental ponds under controlled prey availability did not significantly change prey intake by pike (i.e. did not significantly change interference or exploitative competition), but induced a neuroendocrine stress response reflecting a size-dependent dominance hierarchy, depressed pike energetic status and lowered pike body growth rate by 23 per cent. Assuming fixed size-dependent survival and fecundity functions parameterized for the Windermere (UK) pike population, stress-induced smaller body size shifts age-specific survival rates and lowers age-specific fecundity, which in Leslie matrices projects into reduced population rate of increase (λ) by 37–56%. Our models also predict that social stress flattens elasticity profiles of λ to age-specific survival and fecundity, thus making population persistence more dependent on old individuals. Our results suggest that accounting for non-consumptive social stress from competitors and predators is necessary to accurately understand, predict and manage food-web dynamics.

The form of Darwinian selection has important ecological and management implications. Negative effects of harvesting are often ascribed to size truncation (i.e. strictly directional selection against large individuals) and resultant decrease in trait variability, which depresses capacity to buffer environmental change, hinders evolutionary rebound and ultimately impairs population recovery. However, the exact form of harvest-induced selection is generally unknown and the effects of harvest on trait variability remain unexplored. Here we use unique data from the Windermere (UK) long-term ecological experiment to show in a top predator (pike, Esox lucius) that the fishery does not induce size truncation but disruptive (diversifying) selection, and does not decrease but rather increases variability in pike somatic growth rate and size at age. This result is supported by complementary modelling approaches removing the effects of catch selectivity, selection prior to the catch and environmental variation. Therefore, fishing most likely increased genetic variability for somatic growth in pike and presumably favoured an observed rapid evolutionary rebound after fishery relaxation. Inference about the mechanisms through which harvesting negatively affects population numbers and recovery should systematically be based on a measure of the exact form of selection. From a management perspective, disruptive harvesting necessitates combining a preservation of large individuals with moderate exploitation rates, and thus provides a comprehensive tool for sustainable exploitation of natural resources.

The form of Darwinian selection has important ecological and management implications. Negative effects of harvesting are often ascribed to size truncation (i.e. strictly directional selection against large individuals) and resultant decrease in trait variability, which depresses capacity to buffer environmental change, hinders evolutionary rebound and ultimately impairs population recovery. However, the exact form of harvest-induced selection is generally unknown and the effects of harvest on trait variability remain unexplored. Here we use unique data from the Windermere (UK) long-term ecological experiment to show in a top predator (pike, Esox lucius) that the fishery does not induce size truncation but disruptive (diversifying) selection, and does not decrease but rather increases variability in pike somatic growth rate and size at age. This result is supported by complementary modelling approaches removing the effects of catch selectivity, selection prior to the catch and environmental variation. Therefore, fishing most likely increased genetic variability for somatic growth in pike and presumably favoured an observed rapid evolutionary rebound after fishery relaxation. Inference about the mechanisms through which harvesting negatively affects population numbers and recovery should systematically be based on a measure of the exact form of selection. From a management perspective, disruptive harvesting necessitates combining a preservation of large individuals with moderate exploitation rates, and thus provides a comprehensive tool for sustainable exploitation of natural resources.

Selection can alter predator-prey interactions. However, whether and how complex food-webs respond to selection remain largely unknown. We show in the field that antagonistic selection from predators and pathogens on prey body-size can be a primary driver of food-web functioning. In Windermere, U.K., pike (Esox lucius, the predator) selected against small perch (Perca fluviatilis, the prey), while a perch-specific pathogen selected against large perch. The strongest selective force drove perch trait change and ultimately determined the structure of trophic interactions. Before 1976, the strength of pike-induced selection overrode the strength of pathogen-induced selection and drove a change to larger, faster growing perch. Predation-driven increase in the proportion of large, infection-vulnerable perch presumably favored the pathogen since a peak in the predation pressure in 1976 coincided with pathogen expansion and a massive perch kill. After 1976, the strength of pathogen-induced selection overrode the strength of predator-induced selection and drove a rapid change to smaller, slower growing perch. These changes made perch easier prey for pike and weaker competitors against juvenile pike, ultimately increasing juvenile pike survival and total pike numbers. Therefore, although predators and pathogens exploited the same prey in Windermere, they did not operate competitively but synergistically by driving rapid prey trait change in opposite directions. Our study empirically demonstrates that a consideration of the relative strengths and directions of multiple selective pressures is needed to fully understand community functioning in nature.

Selection can alter predator-prey interactions. However, whether and how complex food-webs respond to selection remain largely unknown. We show in the field that antagonistic selection from predators and pathogens on prey body-size can be a primary driver of food-web functioning. In Windermere, U.K., pike (Esox lucius, the predator) selected against small perch (Perca fluviatilis, the prey), while a perch-specific pathogen selected against large perch. The strongest selective force drove perch trait change and ultimately determined the structure of trophic interactions. Before 1976, the strength of pike-induced selection overrode the strength of pathogen-induced selection and drove a change to larger, faster growing perch. Predation-driven increase in the proportion of large, infection-vulnerable perch presumably favored the pathogen since a peak in the predation pressure in 1976 coincided with pathogen expansion and a massive perch kill. After 1976, the strength of pathogen-induced selection overrode the strength of predator-induced selection and drove a rapid change to smaller, slower growing perch. These changes made perch easier prey for pike and weaker competitors against juvenile pike, ultimately increasing juvenile pike survival and total pike numbers. Therefore, although predators and pathogens exploited the same prey in Windermere, they did not operate competitively but synergistically by driving rapid prey trait change in opposite directions. Our study empirically demonstrates that a consideration of the relative strengths and directions of multiple selective pressures is needed to fully understand community functioning in nature.

Selective harvest of large individuals should alter natural adaptive landscapes and drive evolution toward reduced somatic growth and increased reproductive investment. However, few studies have simultaneously considered the relative importance of artificial and natural selection in driving trait changes in wild populations. Using 50 years of individual-based data on Windermere pike (Esox lucius), we show that trait changes tracked the adaptive peak, which moved in the direction imposed by the dominating selective force. Individual lifetime somatic growth decreased at the start of the time series because harvest selection was strong and natural selection was too weak to override the strength of harvest selection. However, natural selection favoring fast somatic growth strengthened across the time series in parallel with the increase in pike abundance and, presumably, cannibalism. Harvest selection was overridden by natural selection when the fishing effort dwindled, triggering a rapid increase in pike somatic growth. The two selective forces appear to have acted in concert during only one short period of prey collapse that favored slow-growing pike. Moreover, increased somatic growth occurred concurrently with a reduction in reproductive investment in young and small female pike, indicating a tradeoff between growth and reproduction. The age-specific amplitude of this change paralleled the age-specific strength of harvest pressure, suggesting that reduced investment was also a response to increased life expectancy. This is the first study to demonstrate that a consideration of both natural selection and artificial selection is needed to fully explain time-varying trait dynamics in harvested populations.

Selective harvest of large individuals should alter natural adaptive landscapes and drive evolution toward reduced somatic growth and increased reproductive investment. However, few studies have simultaneously considered the relative importance of artificial and natural selection in driving trait changes in wild populations. Using 50 years of individual-based data on Windermere pike (Esox lucius), we show that trait changes tracked the adaptive peak, which moved in the direction imposed by the dominating selective force. Individual lifetime somatic growth decreased at the start of the time series because harvest selection was strong and natural selection was too weak to override the strength of harvest selection. However, natural selection favoring fast somatic growth strengthened across the time series in parallel with the increase in pike abundance and, presumably, cannibalism. Harvest selection was overridden by natural selection when the fishing effort dwindled, triggering a rapid increase in pike somatic growth. The two selective forces appear to have acted in concert during only one short period of prey collapse that favored slow-growing pike. Moreover, increased somatic growth occurred concurrently with a reduction in reproductive investment in young and small female pike, indicating a tradeoff between growth and reproduction. The age-specific amplitude of this change paralleled the age-specific strength of harvest pressure, suggesting that reduced investment was also a response to increased life expectancy. This is the first study to demonstrate that a consideration of both natural selection and artificial selection is needed to fully explain time-varying trait dynamics in harvested populations.