Energy allocation strategy modifies growth–survival trade-offs in juvenile fish across ecological and environmental gradients

In young temperate zone fishes, conflicting energy demands lead to variability in growing season and winter survival. Growing season survival is driven by size-dependent predation risk whereas winter survival is constrained by autumn body size, energy storage and winter duration. We developed a model of the seasonality of energetics coupled to empirical measures of resource availability, size-dependent predation and temperature seasonality for rainbow trout (Oncorhynchus mykiss) in two sets of lakes in British Columbia, Canada, representing endpoints of a gradient of temperature, growing season duration and winter duration. This model was used to determine the energy allocation strategy which maximized first-year survival across these gradients. Survival was sensitive to the timing of the switch from somatic to storage strategies in cold, short growing season, low resource environments. A broader range of energy allocation strategies were viable in warmer, longer growing season and higher resource lakes. We used empirical observations of autumn energy storage and our modeled values for size-dependent minimal lipid levels needed to survive winter in each system to estimate winter survival for juvenile rainbow trout. Winter survival estimates were 6% in cold lakes with low resources, 82% in warm, lakes with low resources and 100% in warm lakes with high resources. Fish in warm lakes with ample resources allocated substantially more to storage than the minimum required to survive winter generated from our model, suggesting additional selection pressures for increased storage when there was ample surplus energy. We concluded that growth–survival trade-offs, modified by seasonality of the environment, influenced the growing season energy allocation strategies for young-of-the-year fish, and suggested this may be important for understanding population viability across environmental gradients.     

Growth hormone-induced effects on mortality, energy status and growth: a field study on Brown Trout (Salmo trutta)

1. Growth hormone (GH) treatment increases the growth rate and competitive ability of salmonids under laboratory conditions. Since fast growth should increase fitness, why is endogenous secretion of GH not higher in wild fish? To address this question, three hypotheses were suggested. H₁: high GH levels reduce antipredator responses and may therefore increase mortality from predation. H₂: high GH levels reduce long-term (e.g. over winter) survival by reducing allocation to critical energy reserves. H₃: GH is not beneficial for growth under natural conditions.
2. To test these hypotheses, the performance of GH-treated juvenile Brown Trout ( Salmo trutta ) and control (placebo) trout was compared in an enclosed stream section subjected to natural predation. Four experiments were conducted during winter, spring, summer and autumn, respectively.
3. Mortality rates were not significantly different between GH-treated and control trout in any of the four experiments so H₁ was not supported. Energy reserves were generally lower in GH-treated fish, which is consistent with H₂, whereas growth rates in mass were higher in GH-treated fish than in controls so H₃ was not supported. This suggests that GH promotes growth at the expense of investment in maintenance.
4. Judging from growth and mortality rates, the fitness of GH-treated and control trout appeared similar. Thus, escaped GH-manipulated fish may compete successfully with wild fish.
5. Hatchery-raised trout with higher initial condition index suffered higher mortality rates than more slender fish. This novel finding may be explained by reduced escape ability related to body morphology, reduced behavioural responses towards predators by high-condition trout, or predator preferences for high-condition fish.     

Rainbow trout in seasonal environments: phenotypic trade‐offs across a gradient in winter duration

Survival through periods of resource scarcity depends on the balance between metabolic demands and energy storage. The opposing effects of predation and starvation mortality are predicted to result in trade‐offs between traits that optimize fitness during periods of resource plenty (e.g., during the growing season) and those that optimize fitness during periods of resource scarcity (e.g., during the winter). We conducted a common environment experiment with two genetically distinct strains of rainbow trout to investigate trade‐offs due to (1) the balance of growth and predation risk related to foraging rate during the growing season and (2) the allocation of energy to body size prior to the winter. Fry (age 0) from both strains were stocked into replicate natural lakes at low and high elevation that differed in winter duration (i.e., ice cover) by 59 days. Overwinter survival was lowest in the high‐elevation lakes for both strains. Activity rate and growth rate were highest at high elevation, but growing season survival did not differ between strains or between environments. Hence, we did not observe a trade‐off between growth and predation risk related to foraging rate. Growth rate also differed significantly between the strains across both environments, which suggests that growth rate is involved in local adaptation. There was not, however, a difference between strains or between environments in energy storage. Hence, we did not observe a trade‐off between growth and storage. Our findings suggest that intrinsic metabolic rate, which affects a trade‐off between growth rate and overwinter survival, may influence local adaptation in organisms that experience particularly harsh winter conditions (e.g., extended periods trapped beneath the ice in high‐elevation lakes) in some parts of their range.     

Size-structured Interactions between Native and Introduced Species: Can Intraguild Predation Facilitate Invasion by Stream Salmonids?

Dynamics of biological invasions may be complicated in size-structured animal populations. Differences in timing of life history events such as juvenile emergence create complex interaction webs where different life stages of native and non-native species act as predators, competitors, and prey. Stream salmonids are an ideal group for studying these phenomena because they display competition and predation in size-structured populations and have been introduced worldwide. For example, introduced rainbow trout (Oncorhynchus mykiss) are invading streams of Hokkaido Island, Japan and have caused declines in native masu salmon (O. masou) populations. However, age-0 rainbow trout emerge later than age-0 masu salmon and are smaller, which raises the question of why they are able to recruit and therefore invade in the face of a larger competitor. We conducted experiments in laboratory stream channels to test effects of increasing density of age-0 and age-1 rainbow trout on age-0 masu salmon. Age-1 rainbow trout dominated age-0 masu salmon by aggressive interference, relegating them to less favorable foraging positions downstream and reducing their foraging frequency and growth. The age-1 trout also reduced masu salmon survival by predation of about 40% of the individuals overall. In contrast, age-0 rainbow trout had little effect on age-0 masu salmon. Instead, the salmon dominated the age-0 trout by interference competition and reduced their survival by predation of 60% of the individuals. In each case, biotic interactions by the larger species on the smaller were strongly negative due to a combination of interspecific competition and intraguild predation. We predict that together these produce a positive indirect effect in the interaction chain that will allow the recruitment of rainbow trout in the face of competition and predation from age-0 masu salmon, and thereby facilitate their invasion in northern Japan.     

Biphasic growth in fish II: empirical assessment

In [Quince, et al., 2008. Biphasic growth in fish I: Theoretical foundations. J. Theor. Biol., doi:10.1016/j.jtbi.2008.05.029], we developed a set of biphasic somatic growth models, where maturation is accompanied by a deceleration of growth due to allocation of energy to reproduction. Here, we use growth data from both hatchery-raised and wild populations of a large freshwater fish (lake trout, Salvelinus namaycush) to test these models. We show that a generic biphasic model provides a better fit to these data than the von Bertalanffy model. We show that the observed deceleration of somatic growth in females varies directly with gonad weight at spawning, with observed egg volumes roughly 50% of the egg volumes predicted under the unrealistic assumption of perfectly efficient energy transfer from somatic lipids to egg lipids. We develop a Bayesian procedure to jointly fit a biphasic model to observed growth and maturity data. We show that two variants of the generic biphasic model, both of which assume that annual allocation to reproduction is adjusted to maximise lifetime reproductive output, provide complementary fits to wild population data: maturation time and early adult growth are best described by a model with no constraints on annual reproductive investment, while the growth of older fish is best described by a model that is constrained so that the ratio of gonad size to somatic weight (g) is fixed. This behaviour is consistent with the additional observation that g increases with size and age among younger, smaller breeding females but reaches a plateau among older, larger females. We then fit both of these optimal models to growth and maturation data from nineteen wild populations to generate population-specific estimates of ‘adapted mortality’ rate: the adult mortality consistent with observed growth and maturation schedules, given that both schedules are adapted to maximise lifetime reproductive output. We show that these estimates are strongly correlated with independent estimates of the adult mortality experienced by these populations.     

Predators select against high growth rates and risk-taking behaviour in domestic trout populations

Domesticated (farm) salmonid fishes display an increased willingness to accept risk while foraging, and achieve high growth rates not observed in nature. Theory predicts that elevated growth rates in domestic salmonids will result in greater risk-taking to access abundant food, but low survival in the presence of predators. In replicated whole-lake experiments, we observed that domestic trout (selected for high growth rates) took greater risks while foraging and grew faster than a wild strain. However, survival consequences for greater growth rates depended upon the predation environment. Domestic trout experienced greater survival when risk was low, but lower survival when risk was high. This suggests that animals with high intrinsic growth rates are selected against in populations with abundant predators, explaining the absence of such phenotypes in nature. This is, to our knowledge, the first large-scale field experiment to directly test this theory and simultaneously quantify the initial invasibility of domestic salmonid strains that escape into the wild from aquaculture operations, and the ecological conditions affecting their survival.     

Cost of predation avoidance in young-of-year lake trout (Salvelinus namaycush): growth differential in sub-optimal habitats

Selection of habitat to avoid predation may affect the diet of young-of-year (YOY) lake trout (Salvelinus namaycush). YOY lake trout may use inshore habitat to avoid predation; this habitat may be sub-optimal for growth. To test this, YOY lake trout were penned in nearshore and offshore pelagic areas of two arctic lakes. Toolik Lake had a lake trout population, the other lake, S6, did not. YOY lake trout in Toolik Lake lost weight, but those offshore lost less weight. The YOY lake trout in Lake S6 gained weight and those offshore gained more weight. The primary diet item of the YOY lake trout in both lakes during this experiment was the zooplankter Diaptomis probilofensis; it was also one of the most abundant species. However, its density inshore in Lake S6 was similar to inshore and offshore densities in Toolik Lake. The increased availability of alternative zooplankton prey in Lake S6 may account for the growth differential of YOY lake trout in Lake S6 relative to Toolik Lake. Bioenergetic modeling of YOY lake trout suggests that growth similar to that in the offshore of Lake S6 would be necessary for successful recruitment. If the reduced zooplankton availability in Toolik Lake leads to the reduced growth of YOY in the inshore and offshore pelagic areas, then these fish will be more susceptable to winter predation/starvation. For YOY lake trout to survive in Toolik Lake they most likely shift to feeding on benthic prey before the end of their first summer. Dept. of Chemical Engineering     

Does Temporal Variation in Predation Risk Influence the Intensity of Antipredator Responses? A Test of the Risk Allocation Hypothesis

Temporal variation in predation risk may fundamentally influence antipredator responses of prey animals. To maximize lifetime fitness, prey must be able to optimize energy gain and minimize predation risk, and responses to current levels of risk may be influenced by background levels of risk. A ‘risk allocation’ model has recently been proposed to predict the intensity of antipredator responses that should occur as predation risk varies over time. Prey animals from high‐risk environments should respond to predators with relatively low intensities of antipredator behaviour because long periods of antipredator behaviour may result in unacceptable decreases in levels of foraging activity. Moreover, animals that are under frequent risk should devote more energy to foraging during brief pulses of safety compared with animals under infrequent attack. In this study, we experimentally tested the risk allocation hypothesis. We exposed juvenile rainbow trout, Oncorhynchus mykiss, to three levels of risk (high, moderate and low) crossed with two levels of temporal variation (exposed to risk three times a day and once a day). In accordance with the model, we found that trout exposed to risky situations more frequently responded with significantly less intense antipredator behaviour than trout exposed to risk infrequently. The intensity of response of trout exposed to moderate risk three times a day decreased to levels similar to situations of no risk. However, in contrast to the second prediction of the model, animals under frequent risk were not more active during periods of safety compared with animals under infrequent risk. Although behaviour in the face of predation risk was dependent on the broader temporal context in which risk varied, the specific predictions of the risk allocation model were only partly supported.     

Biology of the minnow Phoxinus phoxinus and its interactions with brown trout Salmo trutta in Øvre Heimdalsvatn, Norway

Minnows have recently been introduced into the mountain lakes of southern Norway. In Øvre Heimdalsvatn, an intensively studied lake, the minnow population has increased rapidly during the last decade. Studies were performed on feeding habits, habitat preferences, age, growth, energy content, densities, parasites, predation and spawning activities. Several aspects were found to be different from other studies, e.g. higher age of sexual maturation, lower growth rate, no feeding on plants, and low infection of parasites. The interaction with the only other fish, the brown trout, is discussed in relation to the minnow population. In 1978 the minnows constituted little more than 10% of production and food consumption of the trout population, and only limited food competition and predation was documented. Increasing competition is likely with a greater minnow population.     

Diel feeding periodicity, daily ration and prey selectivity in juvenile brown trout in a subarctic river

Feeding of age-1 brown trout Salmo trutta in a third-order river in northern Finland was usually highest in the twilight hours and lowest around midday. Diel periodicity in food intake was less distinct and rarely significant for age-0 trout. Daily rations declined seasonally, being lowest in October, and highest in June (age-1 trout) or early August (age-0 trout). Prey selection did not differ between day and night, but differences between age classes and sampling dates were distinct. Age-0 trout preferred Ephemerella nymphs in summer and Micrasema larvae later in the season. Age-1 trout fed selectively on caddis larvae on all sample dates. Aerial insects and Baetis nymphs were avoided by both age classes on most occasions. These patterns of preference are probably explained by a trend towards epibenthic feeding, which may be a predominant foraging mode in some trout populations. Nevertheless, repuscular peaks in feeding periodicity suggest that trout were able to capitalize on the increased availability of drifting prey in the twilight, especially in the early summer months.     

Risk-taking behaviour in foraging young-of-the-year perch varies with population size structure

I investigated if risk-taking behaviour of young-of-the-year (YOY) perch Perca fluviatilis was connected with population-specific predation patterns in four lakes in northern Sweden. The lakes differ in perch size distribution, according to earlier fishing surveys. Thus, the most intense predation pressure by cannibals is assumed to occur at different prey-size windows in the four lakes. In an aquarium study, I observed groups of perch, and registered time spent foraging in an open habitat and number of prey attacks in the presence of a predator. Perch from Ängersjön, with the highest proportion of large fish in the population, spent more time in the open area than those from Fisksjön that has a dense population of mainly small perch. The Ängersjön perch also made more prey attacks than did perch from Fisksjön and Bjännsjön. Relative differences in predation risk in the four lakes were estimated as cannibalistic attack rates, on a range of sizes of YOY perch, calculated from population size distributions. Principal component analysis on predation risk patterns resulted in two components, of which PC1 explained 79.1% of the variation. High scores of PC1 indicated low cannibalistic attack rates on smaller perch (10–20 mm) and high rates on larger fish (≥30 mm), while low scores indicated the opposite. The level of risk-taking behaviour in the aquarium study positively correlated with lake-specific PC1 scores. The perch with the most cautious behaviour in the aquaria originated from the population with the highest predation pressure on early stages. The boldest perch came from the lake with low predation on the smallest, but with higher predation on larger YOY perch. Thus, the influence of predation risk on behaviour patterns in perch may depend on the timing of the highest exposure to predators.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Antipredatory behaviour in lizards: interactions between group size and predation risk

Group size effects on antipredatory behaviour are well documented in numerous animals, but little is known about how the level of predation risk influences this process. We tested the hypothesis that group size and level of risk interact to affect the levels of antipredatory behaviour in the group-living sun skink, Lampropholis delicata. We controlled the size of lizard groups (N=1, 2, 4, 8 or 12 females) and altered predation risk by providing either a basking tile covered with chemical cues from a predator (high risk) or one without scent (low risk). The time allocated to individual antipredatory behaviour decreased significantly with increasing group size. The relation between group size and time allocated to individual antipredatory behaviour was nonlinear and asymptotic, and did not change under low and high risks of predation. However, group size and predation risk interacted to affect significantly the time that lizards allocated to antipredatory behaviour. When the overall risk from predators was high, individual responsiveness decreased strongly as group size became larger. In contrast, when the overall risk from predators was low, individual responsiveness decreased weakly as group size became larger. Consequently, the time that lizards allocated to antipredatory behaviour under different risks of predation converged as group size increased.     

Is the midsummer decline of Daphnia really induced by age-0 fish predation? Comparison of fish consumption and MDaphnia mortality and life history parameters in a biomanipulated reservoir

We analysed the temporal pattern of a Daphnia galeata populationand the development of the age-0 Ash community in a long-termbiomanipulated lake (Bautzen reservoir, Germany) during springand early summer of two successive years. In Bautzen reservoir,the age-0 Rsh-Daphnia interaction is a key process within thefood web due to the low abundance of adult zooplanktivorousfish. Daily consumption of daphnids by age-0 fish, as estimatedby a bioenergetics model, was compared to daily mortality ratesof daphnids. In addition, we estimated life history parametersof Daphnia that may indicate predation impact by fish. A midsummerdecline of daphnids occurred only in July of 1996, whereas in1995 the Daphnia biomass remained >4 mg wet weight 1^–1for the entire summer. The percentage of total Daphnia mortalitydue to fish predation before the onset of the midsummer declinewas –2% day^–1. Temporal patterns of individual size,clutch size and size at maturity also indicated that the 1996midsummer decline of daphnids was not the exclusive consequenceof age-0 fish predation. Instead, low reproductive capacityof daphnids also contributed significantly to the decline. Consequently,year-to-year variation of the Daphnia dynamics may be determinedby a fine-tuned ‘timing’ between the period of reducedfecundity and the time of the strongest predation impact byage-0 fish.     

Evolutionary models of metabolism, behaviour and personality

I explore the relationship between metabolism and personality by establishing how selection acts on metabolic rate and risk-taking in the context of a trade-off between energy and predation. Using a simple time budget model, I show that a high resting metabolic rate is not necessarily associated with a high daily energy expenditure. The metabolic rate that minimizes the time spent foraging does not maximize the net gain rate while foraging, and it is not always advantageous for animals to have a higher metabolic rate when food availability is high. A model based on minimizing the ratio of mortality rate to net gain rate is used to determine how a willingness to take risks should be correlated with metabolic rate. My results establish that it is not always advantageous for animals to take greater risks when metabolic rate is high. When foraging intensity and metabolic rate coevolve, I show that in a particular case different combinations of foraging intensity and metabolic rate can have equal fitness.     

Spatial and Temporal Trade-Offs by Bluegills in Floodplain River Ecosystems

Ecological trade-offs by organisms to minimize mortality and maximize growth is a foundational theme in ecology. Yet, these trade-offs are rarely examined within spatially complex, temporally variable ecosystems, such as floodplain rivers. Here, we evaluate ecological trade-offs across space and time for the bluegill (Lepomis macrochirus) in two unregulated river ecosystems in southeastern USA. Life-history differences among spatially segregated main channel and floodplain lake populations were used to assess effects of habitat type on bluegill fitness. Growth, condition, and gonadal somatic index were all significantly enhanced in floodplain lakes relative to the main channel. Furthermore, stomach fullness was significantly higher, and predator densities significantly lower in floodplain lakes thereby providing an ecological explanation for the life-history plasticity observed across the riverscape. However, historical observations suggested that although floodplain lakes are highly productive for bluegills, they are also prone to complete desiccation by drought approximately every 5 years, revealing the ultimate value of channel habitat, which does not dry, as desiccation refugia. Bluegills are faced with a balancing act associated with variation in foraging opportunities, and risks to predation and desiccation, that change in both the temporal and the spatial dimensions of floodplain rivers. The differential responses to these opportunities and risks help to explain why both habitats remain actively populated by bluegills, as well as many other organisms, in these and many other natural rivers.     

Intraspecific variation in behaviour: effects of evolutionary history, ontogenetic experience and sex

Geographical variation in behaviour within species is common. However, how behavioural plasticity varies between and within locally adapted populations is less studied. Here, we studied behavioural plasticity induced by perceived predation risk and food availability in pond (low predation - high competition) vs. coastal marine (high predation - low competition) nine-spined sticklebacks (Pungitius pungitius) reared in a common garden experiment. Pond sticklebacks were more active feeders, more risk-taking, aggressive and explorative than marine sticklebacks. Perceived predation risk decreased aggression and risk-taking of all fish. Food restriction increased feeding activity and risk-taking. Pond sticklebacks became more risk-taking than marine sticklebacks under food shortage, whereas well-fed fish behaved similarly. Among poorly fed fish, males showed higher drive to feed, whereas among well-fed fish, females did. Apart from showing how evolutionary history, ontogenetic experience and sex influence behaviour, the results provide evidence for habitat-dependent expression of adaptive phenotypic plasticity.     

Impacts of introduced brown and rainbow trout on benthic invertebrate communities in shallow New Zealand lakes

1. Brown and rainbow trout have been introduced to many inland waters in New Zealand, but research on the impacts on native communities has focused mainly on streams. The purpose of this study was to compare the benthic communities of trout and troutless lakes. Based on previous studies in North America and Europe, we predicted that the benthic biomass, and especially the abundance of large invertebrates, would be lower in lakes with trout as compared to those without. We surveyed the invertebrate fauna of 43 shallow, high‐elevation lakes (26 with and 17 without trout) in four geographic clusters on the central South Island and then conducted a detailed quantitative study of invertebrate biomass and community structure in 12 of these lakes. 2. Benthic community composition and diversity of lakes with and without trout were nearly identical and biomass was as high or higher in the lakes with as without trout. There was no evidence that trout have caused local extinctions of benthic invertebrates. Although the proportional abundance of large‐bodied aquatic was slightly lower in lakes with than without trout, the abundance of several groups of large‐bodied benthic taxa (dragonflies, caddisflies and water bugs) did not differ. 3. Our findings are in contrast to those in North American and Europe where trout introductions into previously troutless lakes have led to declines in the abundance of benthic invertebrates, especially large‐bodied taxa. We propose that the modest effects of trout in New Zealand could be explained by (i) the high areal extent of submergent vegetation that acts as a benthic refuge, (ii) low intensity of trout predation on benthic communities and/or (iii) characteristics of the benthic invertebrates that make them relatively invulnerable to fish predation. 4. Regardless of the relative importance of these hypotheses, our results emphasise that the same invertebrates occurred in all of the lakes, regardless of size, elevation and presence of trout, suggesting habitat generalists dominate the benthic fauna in shallow New Zealand lakes.     

Influence of sex, body size, and reproduction on overwinter lipid depletion in brook trout

Expressed as percentages of total fresh body weight, lipids of brook trout Salvelinus fontinalis declined between October and April: reproductive males from 2·89 to 1·22%, reproductive females from 3·19 to 1·84%, and non-reproductive males and females from 2·75 to 2·08%. The absolute and proportional overwinter reduction in lipids among reproductive trout was more than twice that of non-reproductive trout, with reproductive males losing significantly more lipids than reproductive females. Larger reproductive individuals lost more lipids during winter, relative to body size, than smaller individuals, although such an effect was not evident among non-reproductive trout. The average overwinter reduction in lipids for reproductive males (58%), females (42%), and non-reproductive trout (24%) was negatively associated with mark-recapture estimates of overwinter survival probabilities of 0·27, 0·36, and 0·58, respectively, providing support for the hypothesis that energy is allocated to reproduction to the detriment of post-reproductive survival. Our emergent hypothesis that reproductive costs differ between sexes, and the life history consequences thereof, merit further study.     

The role of food quality in clonal succession in Daphnia: an experimental test

A high genetic variation and recurrent changes in the genetic structure have been found in many pelagic populations. However, evidence that directly links these changes to differences in the ecological performance of particular genotypes is scarce. We hypothesized that within Daphnia, the specialization of clones occurring in a particular season to the food quality specific for that time of the year is responsible for the observed changes in the genetic structure of a population. This hypothesis was tested by comparing the fitness of spring and summer clones of the Daphnia longispina group, given food of biochemical quality relevant to these seasons. We identified significant intraspecific differences between clones of Daphnia that are specific for particular seasons, but there was no evidence that clones are adapted to the food quality available at the respective times of year. Summer clones reproduce at smaller size, and have a lower juvenile specific growth rate as compared to spring clones, irrespective of food quality. Spring clones invest more energy in somatic growth at the cost of reproduction, whereas summer clones invest more energy in reproduction at the cost of somatic growth. On the basis of the observed differences between spring and summer clones in their patterns of energy allocation, we suggest that other factors, most likely predation, are the major forces driving phenotypic and genetic diversity in the investigated Daphnia population of a large lake.