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Phenotypic plasticity is common among animal taxa. While there are clearly limits and likely costs to plasticity, these costs are unknown for most organisms. Further, as plasticity is partially genetically determined, the potential magnitude of exhibited plasticity may vary among individuals. In addition to phenotypic plasticity, various animal taxa also display sexual size dimorphism, a feature ultimately thought to arise due to differential size-dependent fitness costs and benefits between sexes. We hypothesized that differential selection acting on males and females can indirectly select for unequal genetically defined plasticity potential between the sexes. We evaluate this possibility for Eurasian perch (Perca fluviatilis), a species that displays modest sexual size dimorphism and habitat-related morphological plasticity. Using 500-year simulations of an ecogenetic agent-based model, we demonstrate that genetically determined morphological plasticity potential may evolve differently for males and females, leading to greater realized morphological variation between habitats for one sex over the other. Genetically determined potential for plasticity evolved differently between sexes across (a) various sex-specific life-history differences and (b) a variety of assumed costs of plasticity acting on both growth and survival. Morphological analyses of Eurasian perch collected in situ were consistent with model predictions: realized morphological variation between habitats was greater for females than males. We suggest that due to sex-specific selective pressures, differences in male and female genetically defined potential for plasticity may be a common feature across organisms.

Metabolic rates are one of many measures that are used to explain species' response to environmental change. Static respirometry is used to calculate the standard metabolic rate (SMR) of fish, and when combined with exhaustive chase protocols it can be used to measure maximum metabolic rate (MMR) and aerobic scope (AS) as well. While these methods have been tested in comparison to swim tunnels and chambers with circular currents, they have not been tested in comparison with a no‐chase control. We used a repeated‐measures design to compare estimates of SMR, MMR and AS in European perch Perca fluviatilis following three protocols: (a) a no‐chase control; (b) a 3‐min exhaustive chase; and (c) a 3‐min exhaustive chase followed by 1‐min air exposure. We found that, contrary to expectations, exhaustive chase protocols underestimate MMR and AS at 18°C, compared to the no‐chase control. This suggests that metabolic rates of other species with similar locomotorty modes or lifestyles could be similarly underestimated using chase protocols. These underestimates have implications for studies examining metabolic performance and responses to climate change scenarios. To prevent underestimates, future experiments measuring metabolic rates should include a pilot with a no‐chase control or, when appropriate, an adjusted methodology in which trials end with the exhaustive chase instead of beginning with it.

Stable isotopes represent a unique approach to provide insights into the ecology of organisms. ��13C and ��15N have specifically been used to obtain information on the trophic ecology and food-web interactions. Trophic discrimination factors (TDF, ��13C and ��15N) describe the isotopic fractionation occurring from diet to consumer tissue, and these factors are critical for obtaining precise estimates within any application of ��13C and ��15N values. It is widely acknowledged that metabolism influences TDF, being responsible for different TDF between tissues of variable metabolic activity (e.g., liver vs. muscle tissue) or species body size (small vs. large). However, the connection between the variation of metabolism occurring within a single species during its ontogeny and TDF has rarely been considered. Here, we conducted a 9-month feeding experiment to report ��13C and ��15N of muscle and liver tissues for several weight classes of Eurasian perch (Perca fluviatilis), a widespread teleost often studied using stable isotopes, but without established TDF for feeding on a natural diet. In addition, we assessed the relationship between the standard metabolic rate (SMR) and TDF by measuring the oxygen consumption of the individuals. Our results showed a significant negative relationship of SMR with ��13C, and a significant positive relationship of SMR with ��15N of muscle tissue, but not with TDF of liver tissue. SMR varies inversely with size, which translated into a significantly different TDF of muscle tissue between size classes. In summary, our results emphasize the role of metabolism in shaping-specific TDF (i.e., ��13C and ��15N of muscle tissue) and especially highlight the substantial differences between individuals of different ontogenetic stages within a species. Our findings thus have direct implications for the use of stable isotope data and the applications of stable isotopes in food-web studies., Zweitver��ffentlichungen der Universit��t Potsdam : Mathematisch-Naturwissenschaftliche Reihe; 1179

Fatty acids are widely used to study trophic interactions in food web assemblages. Generally, it is assumed that there is a very small modification of fatty acids from one trophic step to another, making them suitable as trophic biomarkers. However, recent literature provides evidence that many fishes possess genes encoding enzymes with a role in bioconversion, thus the capability for bioconversion might be more widespread than previously assumed. Nonetheless, empirical evidence for biosynthesis occurring in natural populations remains scarce. In this study, we investigated different feeding types of perch (Perca fluviatilis) that are specialized on specific resources with different levels of highly unsaturated fatty acids (HUFAs), and analyzed the change between HUFA proportions in perch muscle tissue compared to their resources. Perch showed matching levels to their resources for EPA, but ARA and especially DHA were accumulated. Compound-specific stable isotope analyses helped us to identify the origin of HUFA carbon. Our results suggest that perch obtain a substantial amount of DHA via bioconversion when feeding on DHA-poor benthic resources. Thus, our data indicate the capability of bioconversion of HUFAs in a natural freshwater fish population. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04913-y.

Individual diet and habitat specialisation are widespread in animal taxa and often related to levels of predation and competition. Mobile consumers such as predatory fish can stabilise lake food webs by ranging over a larger area than their prey, thereby switching between habitats. Although, this switching assumes that the predator has equal preference for the available prey, individual diet specialisation and morphological adaptations to different habitats could potentially prevent individuals from switching between habitats. In this study, we assessed the niche width and individual specialisation in Eurasian perch Perca fluviatilis in response to a shift in habitat use by manipulating the ability for this top predator to couple habitats. We ran an eight weeks pond experiment, to test the effect of habitat switching on diet and morphological specialisations. We show that habitat coupling influenced individual diet specialisation and niche use in expected directions where specialisation increased with decreasing habitat switching. In contrast to expectations, the morphological variation decreased with increasing diet specialisation. Our results expand on previous work and suggest that individual specialisation and niche width can impact the ability of mobile predators to couple habitats. Furthermore, it shows the importance of individual specialisations in relation to habitat coupling.

Fatty acids (FAs) are key nutrients for fitness which take part in multiple physiological processes over the ontogeny of organisms. Yet, we lack evidence on how FA nutrition mediates life-history trade-offs and ontogenetic niche shifts in natural populations. In a field study, we analyzed ontogenetic changes in the FAs of Eurasian perch (Perca fluviatilis L.), a widespread fish that goes through ontogenetic niche shifts and can have high individual niche specialization. Diet explained most of the variation in the FA composition of perch dorsal muscle over early ontogeny (28%), while the total length explained 23%, suggesting that perch significantly regulated FA composition over early ontogeny. Condition explained 1% of the remaining variation. 18:3n-3 (ALA) and 18:4n-3 (SDA) indicated planktivory; 18:1n-7, benthivory; and 22:6n-3 (DHA), piscivory in perch diet. Conversely, perch regulated long-chained polyunsaturated fatty acids (PUFAs), such as 20:5n-3 (EPA), 20:4n-6 (ARA) and 22:6n-3 (DHA) over ontogeny, emphasizing the role of such FAs in early growth and sexual maturation. Adult perch increasingly retained 16:1n-7 and 18:1n-9 suggesting higher energy storage in older perch. Furthermore, differences in DHA availability in diet correlated with intra-cohort differences in perch growth, potentially hindering the overall use of benthic resources and promoting earlier shifts to piscivory in littoral habitats. Overall, this study indicates that in addition to diet, internal regulation may be more important for FA composition than previously thought. Differences between FA needs and FA availability may lead to life-history trade-offs that affect the ecology of consumers, including their niche. Electronic supplementary material The online version of this article (10.1007/s00442-020-04668-y) contains supplementary material, which is available to authorized users.

Among-individual diet variation is common in natural populations and may occur at any trophic level within a food web. Yet, little is known about its variation among trophic levels and how such variation could affect phenotypic divergence within populations. In this study we investigate the relationships between trophic position (the population’s range and average) and among-individual diet variation. We test for diet variation among individuals and across size classes of Eurasian perch (Perca fluviatilis), a widespread predatory freshwater fish that undergoes ontogenetic niche shifts. Second, we investigate among-individual diet variation within fish and invertebrate populations in two different lake communities using stable isotopes. Third, we test potential evolutionary implications of population trophic position by assessing the relationship between the proportion of piscivorous perch (populations of higher trophic position) and the degree of phenotypic divergence between littoral and pelagic perch sub-populations. We show that among-individual diet variation is highest at intermediate trophic positions, and that this high degree of among-individual variation likely causes an increase in the range of trophic positions among individuals. We also found that phenotypic divergence was negatively related to trophic position in a population. This study thus shows that trophic position is related to and may be important for among-individual diet variation as well as to phenotypic divergence within populations. Electronic supplementary material The online version of this article (doi:10.1007/s00442-014-3203-4) contains supplementary material, which is available to authorized users.

Individuals are constantly in competition with one another and, on both ecological and evolutionary timescales, processes act to reduce this competition and promote the gain of fitness advantages via diversification. Here we have investigated the genetic (AFLP) and morphological (geometric morphometrics) aspects of the littoral–pelagic axis, a commonly observed resource polymorphism in freshwater fishes of postglacial lakes. We found a large degree of variation in the genetic and morphological divergence between littoral and pelagic perch and roach across Swedish lakes. Although there was evidence of assortative mating (elevated kinship values) in both species, we could not find any significant coupling of morphology and genetic divergence. Instead, there was evidence that the extent of resource polymorphism may be largely caused by phenotypic plasticity. These results suggest that assortative mating, which can lead to genetically determined adaptive divergence, does occur in these species, particularly perch, but not according to genetically fixed morphological traits. The behavioural mechanisms facilitating associative mating need to be investigated to explore the interaction between phenotypic plasticity and adaptive genetic divergence and their roles in diversification. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114, 929–940.

Trophic polymorphism has found to be common in many taxa and is a suggested mechanism of ecological speciation. To characterize the trophic linkages of specific morphotypes of organisms as well as a time-integrated niche use, several methods are available. In this study, we present data of multiple techniques to investigate the trophic divergence of Eurasian perch (Perca fluviatilis) that displays well-studied trophic polymorphism associated with littoral and pelagic habitats in lakes. We combined bulk stable isotope and fatty acid analyses on the muscle tissue of perch from three different lakes in Sweden with analyses of stomach content. By comparing the three methods, we aimed at providing a broad and highly resolved picture on the trophic divergence in freshwater fish. The degree in morphological divergence varied between perch caught in the three different lakes. Generally, perch caught in the pelagic zone were more streamlined compared to the ones caught in the littoral zone that had a deeper body, as shown by geometric morphometrics. The three diet assessment methods revealed different levels of information. Data on stomach content showed some preferences for specific dietary items in littoral and pelagic perch, but general trophic specialization could not be concluded due to the small sample size. Analyses of delta C-13 and delta N-15, however, confirmed these results as a long-term pattern connected to specific habitat use in two of the three lakes. Fatty acid signatures of perch reflected partly those of the prey items of the specific habitats. Although the proportions of the essential fatty acid 22:6n-3 were lower in littoral resources, the proportions in littoral fish were similar to the ones caught in the pelagic zone. We concluded that although a fundamental contribution from littoral resources exists in littoral phenotypes, a minor reliance on pelagic prey items is obviously needed to provide essential compounds. Thus, by combining the methods to characterize direct resource use (i.e., stomach analyses) with others that utilize trophic biomarkers (i.e., analyses of stable isotopes and fatty acids), we were able to illustrate the degree of variation in trophic divergence of perch but also shed some light on potential trade-offs that are related to resource specialization in freshwater fish.

Predation is a major driving force in evolution. Predation has been shown to select for size, morphology, and camouflage. Many animals use camouflage to reduce predation risk. In some cases, individuals can adjust their pigmentation, enabling them a higher survival in a heterogeneous environment. Here, we show that the difference in pigmentation between juvenile perch individuals (Perca fluvuiatilis) occupying different environments (open water and vegetated habitats of lakes) is likely a consequence of predator selection. Lightly pigmented individuals have a higher chance of survival in open water whereas darker pigmented individuals survive better in vegetation. As a response to predators, individuals forced into the vegetation by predators developed darker skin whereas the skin of individuals forced into open water became lighter. In a common garden experiment, in the absence of predation, we found that pigmentation in juvenile perch is only due to plasticity and not to genetic variation. However, contrary to predictions, individuals raised in open water developed darker skin compared to individuals raised in vegetation. This may be a response to UV-stress. Overall, our results suggest that predation can be a strong selective agent on pigmentation differences among conspecifics occupying different habitats.

The turnover and distribution of energy and nutrients in food webs is influenced by consumer stoichiometry. Although the stoichiometry of heterotrophs is generally considered to vary only little, t ...

Due to altered biogeochemical processes related to climate change, highly colored dissolved organic carbon (DOC) from terrestrial sources will lead to a water "brownification" in many freshwater systems of the Northern Hemisphere. This will create deteriorated visual conditions that have been found to affect habitat-specific morphological variations in Eurasian perch (Perca fluviatilis) in a previous study. So far, potential drivers and ultimate causes of these findings have not been identified. We conducted a field study to investigate the connection between morphological divergence and polyunsaturated fatty acid (PUFA) composition of perch from six lakes across a gradient of DOC concentration. We expected a decrease in the prevalence of PUFAs, which are important for perch growth and divergence with increasing DOC concentrations, due to the restructuring effects of DOC on aquatic food webs. In general, rate of morphological divergence in perch decreased with increasing DOC concentrations. Proportions of specific PUFAs (22:6n-3, 18:3n-3, 20:5n-3, and 20:4n-6) identified to primarily contribute to overall differences between perch caught in clear and brown-water lakes tended to be connected to overall decline of morphological divergence. However, no overall significant relationship was found, indicating no severe limitation of essential fatty acids for perch inhabiting brown water lakes. We further broaden our approach by conducting a laboratory experiment on foraging efficiency of perch. Therefore, we induced pelagic and littoral phenotypes by differences in habitat-structure and feeding mode and recorded attack rate in a feeding experiment. Generally, fish were less efficient in foraging on littoral prey (Ephemeroptera) when visual conditions were degraded by brown water color. We concluded that browning water may have a strong effect on the forager's ability to find particular food resources, resulting in the reduced development of evolutionary traits, such as habitat- specific morphological divergence.

It has been hypothesized that inter-specific competition will reduce species niche utilization and drive morphological evolution in character displacement. In the absence of a competitor, intra-specific competition may favor an expansion of the species niche and drive morphological evolution in character release. Despite of this theoretical framework, we sometimes find potential competitor species using the same niche range without any partitioning in niche. We used a database on test fishing in Sweden to evaluate the factors (inter- and intraspecific competition, predation, and abiotic factors) that could influence habitat choice of two competitor species. The pattern from the database shows that the occurrence of perch and roach occupying both littoral and pelagic habitats of lakes in Sweden is a general phenomenon. Furthermore, the results from the database suggest that this pattern is due to intra-specific competition rather than inter-specific competition or predation. In a field study, we estimated the morphological variation in perch and roach and found that, individuals of both species caught in the littoral zone were more deeper bodied compared to individuals caught in the pelagic zone. Pelagic perch fed more on zooplankton compared to littoral perch, independent of size, whereas the littoral perch had more macroinvertebrates and fish in their diet. Pelagic roach fed more on zooplankton compared to littoral roach, whereas littoral individuals fed more on plant material. Furthermore, we sampled littoral and pelagic fish from another lake to evaluate the generality of our first results and found the same habitat associated morphology in both perch and roach. The results show a consistent multi-species morphological separation in the littoral and pelagic habitats. This study suggests that intra-specific competition is possibly more important than inter-specific competition for the morphological pattern in the perch-roach system.

The morphological response of two 1 year-old prey species, perch Perca fluviatilis and roach Rutilus rutilus to the presence of predatory pike Esox lucius, was tested in a 6 week aquarium experiment. The growth of perch was higher than that of roach but there was no difference in growth of prey between predator and control treatments. The presence of pike caused morphological changes in both perch and roach. Perch showed an increase in body depth during the experiment whereas the morphological response of roach to pike was more related to a displacement of the dorsal and pelvic fins and width of the anal fin. Differences in morphological responses of perch and roach to pike can be interpreted by their overall differences in antipredator strategies.

Microplastic's triple threat The billions of tons of plastics that we release into the environment for the most part do not biodegrade. But they do degrade, breaking into ever smaller particles that end up in the oceans. Lönnstedt et al. show that the impacts of these microplastics are multifold (see the Perspective by Rochman). Eurasian perch larvae exposed to microplastics were less active, less responsive to predator cues, more likely to be eaten, and less likely to thrive—preferring to eat plastic rather than their natural prey. Science , this issue p. 1213 ; see also p. 1172

Resource competition has been hypothesized to be important in driving divergence by natural selection. The effect of competition on morphological divergence and plasticity has however rarely been investigated. Since low growth rates might constrain morphological modulation and individual growth rates usually are negatively related to the intensity of competition, there might be a connection between competition, growth rate and morphological divergence. We performed an aquarium experiment with young-of-the-year Eurasian perch (Perca fluviatilis L.) to investigate how individual growth rate affected morphological plasticity induced by contrasting habitat treatments. Furthermore, in a field study of 10 lakes we also related the degree of morphological differentiation between habitats to the intraspecific competitior biomass. In the aquarium experiment we found that morphological plasticity was growth rate dependent in that morphological differentiation between the habitat treatments was confined to high individual growth rates. In the field study we found that morphological differentiation between habitats decreased with increasing intraspecific competitior biomass. Since plasticity is hypothesized to be important in divergence and intraspecific biomass could serve as a proxy for the level of competition, we suggest that our results indicate that morphological divergence might be constrained during periods of intense intraspecific competition due to low growth rates. A possible scenario is that at low growth rates all energy available is used for metabolic maintenance and no surplus energy is therefore available for morphological modulation.

The work in this thesis deals with the ecology and evolution of adaptive individual variation. Ecologists have long used niche theory to describe the ecology of a species as a whole, treating conspecific individuals as ecological equivalent. During recent years, research about individual variation in diet and morphology has gained interest in adaptive radiations and ecological speciation. Such variation among individual niche use may have important conservation implications as well as ecological and evolutionary implications. However, up to date we know very little about the extension of this phenomenon in natural populations and the mechanisms behind it.The results in this thesis show that the extension of individual diet specialization is widely spread throughout the animal kingdom. The variation in diet is mainly correlated to morphological variation but not always. Furthermore, this variation in diet and morphology among individuals could be both genetically determined and environmentally induced and it mainly comes from trade-offs in foraging efficiency between different prey types. The results from a number of studies of perch also show that individual perch differ in morphology and diet depending on habitat, where littoral perch has a deeper body compared to pelagic perch. This difference in morphology corresponds to functional expectations and is related to foraging efficiency trade-offs between foraging in the littoral and pelagic zone of a lake. The variation in morphology in perch is mainly due to phenotypic plasticity but there are also small genetic differences between the littoral and pelagic perch. Two separate studies show that both predation and competition may be important mechanism for the variation in morphology and diet in perch.In conclusion, the results in this thesis show that individual variation in diet and habitat choice is a common phenomenon with lots of ecological and evolutionary implications. However, there are many mechanisms involved in this phenomenon on which we are just about to start learning more about, and only further research in this area will give us the full insight.

Studies on resource polymorphism have mainly been considered at the end stage of ontogeny, whereas many species undergo diet changes as they grow. We conducted a field survey to analyze the role of adaptive variation during ontogeny in Eurasian perch (Perca fluviatilis). We caught perch from the littoral and pelagic zones of a lake to investigate whether perch differ in morphology and diet between these habitats. We also investigated whether there were any differences in morphological trajectories during the ontogeny of perch collected from the two habitats. We found that perch caught in the littoral habitat, independently of size, had a deeper body, larger head and mouth and longer fins than perch caught in the pelagic zone. Macroinvertebrates and fish dominated the diet of littoral perch, whereas the diet of the pelagic perch consisted mainly of zooplankton and to some extent fish. Independently of size, the more streamlined individuals had a larger proportion of zooplankton and a smaller proportion of macroinvertebrates in their diet than the deeper-bodied individuals, indicating a relation between diet and morphology. Some morphological characters followed different ontogenetic trajectories in the two habitats; e.g. the changes to a deeper body and a larger head were faster in the littoral than in the pelagic perch. The relationship between the length of perch and the size of the mouth and fins also differed between perch from the two habitats, where the increase in the length of the pelvic fin and the area of the mouth increased faster with size in the littoral perch. Our findings show that variation in morphology between habitats differs during ontogeny in a way that corresponds to functional expectations for fish species that occupy these habitats.

The combined effects of predators on prey may substantially differ from thatof each predator species alone because of alterations in prey behavior. Using enclosures within a pond, we examined experimentally the effects of two piscivorous predators on prey mortality and prey resource levels in two habitats. The two predators use two different foraging modes, which also allowed us to examine the behaviorally induced indirect effects of prey on predator growth and prey food resources. Both perch (Perca fluviatilis) and pike (Esox lucius) caused significant mortality of roach (Rutilus rutilus), and the combined predator mortality was higher than predicted from a multiplicative prey consumption model. Growth rates of perch were similar when enclosures contained only perch and when they contained perch combined with pike. The growth rate of pike was higher when they were together with perch compared to when alone. Growth of roach was similar among treatments. The invertebrate food resources of roach increased by a factor 10 in the open water but remained at similar levels throughout the experiment in the vegetation. Biomass of Daphnia longispina, the dominant zooplankton species in the open water, was strongly correlated with mortality of roach, indicating a density-mediated indirect effect of predators on prey resources. There was no indirect effect on D. longispina in the vegetation caused by habitat restriction of roach and only a weak relationship in the open water. There was a strong indirect effect of pike predation on macroinvertebrates induced by a habitat shift of roach. Our results suggest that there was facilitation between predators caused by conflicting antipredator behavior of roach, which resulted in density-mediated indirect effects on prey resources. The behavioral response of roach to the two predators also induced indirect effects on invertebrate prey.

Structurally complex environments strongly affect the behaviours and foraging efficiencies of predators and prey. I studied temporal variation in the habitat distribution of perch (Perca fluviatilis) and pike (Esox lucius) in relation to habitat complexity and prey abundance in a lake. The study involved quantitative estimates of different habitat types, estimates of macroinvertebrate prey availability, and distribution and movement patterns of the fish. The numbers of 80-110 mm perch in the littoral zone decreased rapidly in spring, which was a result of either perch moving to the pelagic zone or predation mortality. Predation mortality is the most plausible explanation because piscivorous perch and pike >160 mm aggregated close to these high abundances of 80-110 mm perch, and 80-110 mm perch used only vegetated habitats as a possible protection against predators. Both the biomass and diversity of macroinvertebrates increased with vegetation density, whereas perch abundance was highest in an intermediate vegetation density. Pike size was inversely related to vegetation density as a result of potential cannibalism from the largest pike individuals, which preferred the tree structure habitat. Perch group size decreased with increasing vegetation density, and perch 160 mm. In contrast, perch >160 mm occurred at decreasing numbers with increasing group size and mainly stayed solitary or in pairs. Perch >160 mm showed no tendencies for homing behaviour and moved actively around the whole lake, whereas pike showed a strong homing behaviour. My study suggests that the structural complexity in the littoral zone can both qualitatively and quantitatively change the interaction between piscivorous predators and their prey.

To study the effects of different predators on the behavior and dynamics of their prey, we compared the performance of perch (Perca fluviatilis) and the abundance of their prey resource in four lakes with two different types of piscivory. In two of the lakes, perch were the only fish species present and piscivory was restricted to cannibalism. In the other two lakes, perch co-occurred with piscivorous pike (Esox lucius). Pike grow to larger sizes and can capture larger prey than piscivorous perch. Therefore, perch reach a size refuge earlier in the lakes without pike. Perch in the lakes lacking pike used both the epilimnion and hypolimnion habitats of the lakes, and a high proportion of the perch occurred below the thermocline. In contrast, perch in the lakes with pike were mainly captured along the littoral zone and were never captured below the thermocline. This difference in habitat use was reflected in different diets of perch in the different lakes. Perch in the lakes with pike were also less abundant than those in the lakes lacking pike and the growth rates of individual perch and the densities of predator-sensitive prey (pelagic and littoral) were higher, indicating that the presence of pike resulted in decreased intraspecific competition in perch. Evidence for a behavioral response to predation risk was found in 1+ (1 yr old) perch, which was suggested to be due to predation risk from both larger perch and pike. It is hypothesized that the use of the cold-water hypolimnion habitat by perch in the lakes lacking pike was a response of perch to reduce metabolic demands under conditions of severe resource limitation. Differences in gape limitation between predators species are suggested to have major effects on size-dependent mortalities and growth rates in prey and thereby on overall community dynamics. Evidence for this latter effect was observed in differences in zoo- plankton size structure between the lakes with and without pike.

When prey encounter predators, they use different cues to indicate how to respond to minimize the predation risk. How such proximate cues in the environment correspond to the ultimate behaviour of the prey are important for understanding the outcome of predator–prey interactions. The precision of the anti-predator response of juvenile perch,Perca fluviatilis, and roach,Rutilus rutilus, when subjected to predation by piscivorous perch and pike,Esox lucius, and to different types of structure was tested in a wading pool experiment. The predation risk was varied between two habitats (one open water and one structured) by confining the predators to one of the two habitats. The prey were free to choose between habitats. Both perch and pike attacked both prey species but they were only successful in capturing roach. Roach swam faster than juvenile perch in the presence of perch, whereas juvenile perch swam faster than roach in the presence of pike. Juvenile perch inspected the predators more and showed a more flexible inspection behaviour than did roach. Juvenile perch decreased the number of switches between habitats and stayed in the predator-free part of the pool in the presence of perch. In contrast, roach increased their habitat switch frequency in the presence of perch and stayed in the vegetation structure even when there were perch there. Both prey species preferred to stay in the vegetation structure when pike were there. Juvenile perch used both the vegetation and pipe structure as refuges whereas roach used only the vegetation structure as a refuge. The results suggest that juvenile perch use a different cue when assessing predation risk and display a more flexible behaviour compared with roach which simply move into vegetation under the threat of predation irrespective of predator location.

The outcome of predator-prey interactions depends on the characteristics of predators and prey as well as the structure of the environment. In a replicated field enclosure experiment, we tested the effects of quantity and quality of different prey refuges (no structure, structure forming a partial refuge, and structure forming a complete refuge) on the interaction between piscivorous perch (Perca fluviatilis) and juvenile perch and roach (Rutilus rutilus). We quantified the behaviour of the predators and the prey and predator-induced prey mortality. The piscivores stayed in or close to the prey refuge and were more dispersed in the presence than in the absence of prey refuges. Survival of juvenile perch and roach decreased in the presence of predators and was higher for juvenile roach than for juvenile perch. In addition, juvenile perch survival increased with refuge efficiency. Roach formed schools which were denser in the presence of predators, had a higher swimming speed (both in the open water and in the refuge) and used a larger area than juvenile perch. Both prey species decreased their distance to the prey refuge and increased the proportion of their time spent in the refuge in the presence of predators. The number of switches between the open-water habitat and the prey refuge was higher for juvenile roach than for juvenile perch. Juvenile perch used different parts of the prey refuge in a flexible way depending both on presence of predators and refuge type whereas juvenile roach used the different parts of the prey refuge in fixed proportions over all refuge treatments. Our results suggest that juvenile roach had a overall higher capacity to avoid predation than juvenile perch. However, in the presence of qualitatively different prey refuges juvenile perch responded to predators with more flexible refuge use than juvenile roach. The differences in antipredator capacities of juvenile perch and roach when subjected to piscivorous perch predation may depend on differences in life history patterns of the two species.

We investigated experimentally how presence or absence of different piscivores influenced habitat use, diet, and individual growth of two size classes of juvenile perch (Perca fluviatilis) and abundances of juvenile perch resources in different habitats. Pond enclosures encompassed 3 X 6 m of dense vegetation and 9 X 6 m of open habitat, and were stocked with 75 young-of-year and 30 1-yr-old perch and with either three piscivorous perch, three northern pike (Esox lucius), or no piscivores. Northern pike were both larger and possessed a larger gape than piscivorous perch. To isolate behavioral responses of juvenile perch to piscivores, we replaced consumed young-of-year perch. Prey fish mortality was lowest without piscivores and highest with northern pike. Young-of-year perch increased their use of vegetation in the presence of both piscivores, whereas 1-yr-old perch increased their use of vegetation only with northern pike. For both age classes of prey fish, increased use of the vegetation led to reduced individual growth, owing to two complementary mechanisms. First, the physical complexity of submerged macrophytes likely interfered with the benthic feeding of perch, Second, increased use of the (relatively small) vegetated habitat increased the mean density experienced by prey fish. Piscivore-induced changes of prey fish densities in the two habitats had substantial effects on the biomass of prey fish resources in the open habitat, but only minor effects in the vegetation. Sialis lutaria, the major predatory macroinvertebrate (approximate to 50% of total macroinvertebrate biomass in the open habitat), and total predatory macroinvertebrates were positively affected by piscivores in the open habitat, but not in the vegetation. Chironomids (

In size-structured populations, interactions are strongly dependent on size- specific foraging and anti-predator capacities of the organism. Conflicting size-specific selection pressures over the ontogeny often have different effects on different species leading to asymmetries in competitive and predator-prey interactions. Habitat complexity is likely to affect such asymmetric interactions due to species/size-specific competitive abilities in different habitats and due to the fact that habitat structural complexity may act differently as a prey refuge for different species. We experimentally analyzed the impact of a piscivorous predator (adult perch, Perca fluviatilis) on performance of juvenile perch and roach (Rutilus rutilus) at different levels of structural complexity (no structure, structure forming a partial refuge, and structure forming a complete refuge) in enclosures in an experimental pond. We measured predator diet and growth, prey fish habitat use, survival, diet and growth, and prey resource levels in different habitats. Prey fish (perch and roach) were found in the diet of piscivorous perch in no refuge and partial refuge treatments. Growth rate of the piscivorous perch decreased with increased refuge efficiency. Juvenile perch increased their proportional use of the structurally complex refuges in the presence of piscivorous perch and the survival increased with increased refuge efficiency (from partial to complete refuge). The diet of juvenile perch changed from predominantly cyclopoid copepods in the absence of predators to predominantly macroin- vertebrates in the presence of predators. There was no effect of predator-induced habitat restriction on growth of juvenile perch. Roach survival also increased with increased refuge efficiency in the presence of predators, and roach survival in the refuge treatments did not differ from each other or from the treatments with predators absent. Predator-induced habitat restriction in roach was associated with a decreased growth of roach. Our results suggest that, compared to juvenile roach, juvenile perch may compensate more for lost foraging opportunity in the open water via increased exploitation of structure- associated prey in refuges. As a result, predator-induced habitat shifts by juvenile perch and roach may alter competitive interactions between the species. On the other hand, structural complexity may form an almost complete refuge for juvenile roach from predators and thereby affect the predator-prey relationships between piscivorous perch and juvenile perch and roach to the advantage of juvenile roach. The demonstrated effects of structural complexity on competitive and predator-prey interactions between perch and roach can be related to the two species' distributions in lakes with different degrees of structural com- plexity.

While phenotypic responses to direct species interactions are well studied, we know little about the consequences of indirect interactions for phenotypic divergence. In this study we used lakes with and without the zebra mussel to investigate effects of indirect trophic interactions on phenotypic divergence between littoral and pelagic perch. We found a greater phenotypic divergence between littoral and pelagic individuals in lakes with zebra mussels and propose a mussel-mediated increase in pelagic and benthic resource availability as a major factor underlying this divergence. Lakes with zebra mussels contained higher densities of large plankton taxa and large invertebrates. We suggest that this augmented resource availability improved perch foraging opportunities in both the littoral and pelagic zones. Perch in both habitats could hence express a more specialized foraging morphology, leading to an increased divergence of perch forms in lakes with zebra mussels. As perch do not prey on mussels directly, we conclude that the increased divergence results from indirect interactions with the mussels. Our results hence suggest that species at lower food web levels can indirectly affect phenotypic divergence in species at the top of the food chain.

Trait combinations that lead to a higher efficiency in resource utilization are important drivers of divergent natural selection and adaptive radiation. However, variation in environmental features might constrain foraging in complex ways and therefore impede the exploitation of critical resources. We tested the effect of water transparency on intra-population divergence in morphology of Eurasian perch (Perca fluviatilis) across seven lakes in central Sweden. Morphological divergence between near-shore littoral and open-water pelagic perch substantially increased with increasing water transparency. Reliance on littoral resources increased strongly with increasing water transparency in littoral populations, whereas littoral reliance was not affected by water transparency in pelagic populations. Despite the similar reliance on pelagic resources in pelagic populations along the water transparency gradient, the utilization of particular pelagic prey items differed with variation in water transparency in pelagic populations. Pelagic perch utilized cladocerans in lakes with high water transparency and copepods in lakes with low water transparency. We suggest that under impaired visual conditions low utilization of littoral resources by littoral perch and utilization of evasive copepods by pelagic perch may lead to changes in morphology. Our findings indicate that visual conditions can affect population divergence in predator populations through their effects on resource utilization.

A predator's foraging strategy is generally described as either a sit-and-wait or an actively searching strategy. Two different behaviour patterns, often associated with these strategies, are territorial behaviour for a sit-and-wait predator, and group foraging behaviour for an actively searching predator. In a pond experiment, the effect of predator density on the efficiency of a sit-and-wait predator, pike, and an actively searching predator, perch, was tested. The foraging efficiencies of these two predators were predicted to be density-dependent since perch is a social species with its efficiency enhanced by mutualistic interactions, while pike is a territorial species with its efficiency reduced by competitive interactions. The predators were stocked at two densities (one or five individuals) in enclosures containing the same prey densities (75 individuals of juvenile perch). The behaviour of predators and prey, prey mortality and predator growth were studied. As expected, swimming speed and move frequency were significantly higher for perch than for pike. In the five-pike treatment, predator activity (swimming speed and move frequency) was highly size-dependent with the large pike both swimming faster and moving more frequently than the small pike. The activity levels of the pike were also reflected in the area used, which was larger for the large pike. Perch at both densities and the large pike used similar areas. As hypothesized, the growth rate of the predators corresponded to their foraging strategies. The growth rate of group foraging perch was higher than that of single perch and pike in both density treatments. In the single predator treatments, pike grew better than perch. In contrast to perch, the single pike grew more than the five pike together.

Spatial and temporal heterogeneity in the environment is a common feature affecting many natural populations. For example, both the resource levels and optimal habitat choices of individuals likely change over time. One way for organisms to cope with environmental variation is to display adaptive plasticity in traits such as behavior and morphology. Since trait plasticity is hypothesized to be a prerequisite for character divergence, studies of mechanisms behind such plasticity are warranted. In this study, we looked at the interaction of two potentially important environmental variables on behavioral and morphological plasticity in Eurasian perch (Perca fluviatilis L.). More specifically, the plastic responses in activity and morphology of perch exposed to different resource levels and simulated habitat types were studied in an aquarium experiment. The resource level experienced had a large influence on plasticity in both activity and morphology. Behavioral adaptations have been thought to mediate morphological transitions, and we suggest that the morphological response to the resource level was mediated by differences in activity and growth rates. The habitat type also affected morphological plasticity but to a lesser extent, and there was no effect on activity from habitat type. Based on these results, we suggest that it is essential to include several environmental factors acting in concert when studying mechanisms behind trait plasticity. We also propose that variation in resource levels might play a key role in fostering trait plasticity in at least fish populations, while other environmental variables such as divergent habitat complexities and prey types might be less influential. Dynamics in resource levels and optimal habitat choices might thus be important factors influencing character divergence in natural populations.

Predators can cause a shift in both density and frequency of a prey phenotype that may lead to phenotypic divergence through natural selection. What is less investigated is that predators have a variety of indirect effects on prey that could potentially have large evolutionary responses. We conducted a pond experiment to test whether differences in predation risk in different habitats caused shifts in behavior of prey that, in turn, would affect their morphology. We also tested whether the experimental data could explain the morphological variation of perch in the natural environment. In the experiment, predators caused the prey fish to shift to the habitat with the lower predation risk. The prey specialized on habitat-specific resources, and there was a strong correlation between diet of the prey fish and morphological variation, suggesting that resource specialization ultimately affected the morphology. The lack of differences in competition and mortality suggest that the morphological variation among prey was induced by differences in predation risk among habitats. The field study demonstrated that there are differences in growth related to morphology of perch in two different habitats. Thus, a trade-off between foraging and predator avoidance could be responsible for adaptive morphological variation of young perch.

In predator-prey interactions, the efficiency of the predator is dependent on characteristics of both the predator and the prey, as well as the structure of the environment. In a field enclosure experiment, we tested the effects of a prey refuge on predator search mode, predator efficiency and prey behaviour. Replicated enclosures containing young of the year (0+) and 1-year-old (1+) perch were stocked with 3 differentially sized individuals of either of 2 piscivorous species, perch (Perca fluviatilis), pike (Esox lucius) or no piscivorous predators. Each enclosure contained an open predator area with three small vegetation patches, and a vegetated absolute refuge for the prey. We quantified the behaviour of the predators and the prey simultaneously, and at the end of the experiment the growth of the predators and the mortality and habitat use of the prey were estimated. The activity mode of both predator species was stationary. Perch stayed in pairs in the vegetation patches whereas pike remained solitary and occupied the corners of the enclosure. The largest pike individuals stayed closest to the prey refuge whereas the smallest individuals stayed farthest away from the prey refuge, indicating size-dependent interference among pike. Both size classes of prey showed stronger behavioural responses to pike than to perch with respect to refuge use, distance from refuge and distance to the nearest predator. Prey mortality was higher in the presence of pike than in the presence of perch. Predators decreased in body mass in all treatments, and perch showed a relatively stronger decrease in body mass than pike during the experiment. Growth differences of perch and pike, and mortality differences of prey caused by predation, can be explained by predator morphology, predator attack efficiency and social versus interference behaviour of the predators. These considerations suggest that pike are more efficient piscivores around prey refuges such as the littoral zones of lakes, whereas perch have previously been observed to be more efficient in open areas, such as in the pelagic zones of lakes.

PREDATORY EFFICIENCY AND PREY SELECTION - INTERACTIONS BETWEEN PIKE ESOX-LUCIUS, LUCIUS, PERCH PERCA-FLUVIATILIS AND RUDD SCARDINUS-ERYTHROPHTHALMUS