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Abstract: Individual variation in the behavior of prey can influence predation risk in complex ways. We ran individual roach (Rutilus rutilus), a common freshwater fish, through a standard refuge emergence protocol to characterize their boldness, a key animal personality trait. We then paired a bold and a shy roach and exposed the pair to one of two predator species that have contrasting hunting modes to ascertain how personality traits shaped their survival during predator encounters. When a paired bold and shy prey fish interacted with a perch predator (active foraging mode), bold and shy prey were consumed in almost equal numbers. However, pike predators (ambush foraging mode) selectively consumed more shy prey, and prey body size and boldness score both contributed significantly to which prey fish was eaten. Our findings support the idea that multiple predators with different foraging modes, and hence differential selection on prey personality, could contribute to maintaining variation in personality in prey populations. Furthermore, for social species, including shoaling fish, the ultimate consequences of an individual’s personality may depend upon the personality of its nearby conspecifics.Significance statement: Animals of the same species often look similar, but individuals show differences in their behavior that can have important consequences, for instance when these individuals interact with predators. The common roach is a freshwater fish that shows inter-individual variation in its propensity to take risks, a key personality trait often termed boldness. Variation in boldness may affect the outcome when roach interact with predators, i.e., if they get eaten or survive. However, we found the impact of roachs’ personality type depends on what species of predatory fish they face. When we put a shy and a bold roach together with predatory perch, the roachs’ personality did not significantly affect which individual was eaten. But when the predator was a pike, the predators selectively ate more shy roach, and the likelihood an individual would be eaten depended on their body size. [ABSTRACT FROM AUTHOR]

Organisms display an impressive array of defence strategies in nature. Inducible defences (changes in morphology and/or behaviour within a prey's lifetime) allow prey to decrease vulnerability to predators and avoid unnecessary costs of expression. Many studies report considerable interindividual variation in the degree to which inducible defences are expressed, yet what underlies this variation is poorly understood. Here, we show that individuals differing in a key personality trait also differ in the magnitude of morphological defence expression. Crucian carp showing risky behaviours (bold individuals) expressed a significantly greater morphological defence response when exposed to a natural enemy when compared with shy individuals. Furthermore, we show that fish of different personality types differ in their behavioural plasticity, with shy fish exhibiting greater absolute plasticity than bold fish. Our data suggest that individuals with bold personalities may be able to compensate for their risk-prone behavioural type by expressing enhanced morphological defences. [ABSTRACT FROM AUTHOR]

Animal migration is one of the most spectacular and visible behavioural phenomena in nature with profound implications for a range of ecological and evolutionary processes. Successful migration hinges on the ability to exploit temporary resources (e.g. food) and evade threats (e.g. predators) as they arise, and thus the timing of migration is often regarded as a dominant predictor of individual migratory success.However, with the exception of intensively studied taxa (mainly birds), relatively few studies have investigated inter‐individual annual and seasonal variation in migratory timing and performance, or tested predictions on how migration across high and low predation‐risk habitats may exert selection on migratory timing. In particular, studies that assess the survival consequences of variation in migratory timing remain rare, which is most likely due to the logistical challenges associated with monitoring survival success and population‐level characteristics simultaneously.Here, we address the above‐mentioned questions using roach Rutilus rutilus, a fish that migrates from lakes characterised by high predation risk into low‐risk streams during winter. Specifically, we used individual‐based tracking of roach in two European lake systems over multiple migration periods (9 and 7 years respectively), to obtain highly detailed (year‐round scheduling, repeat journeys and the fate of individuals) data on the variability/synchrony of migratory timing in spring and autumn respectively.We report seasonal differences in the variability of migratory timing, with lower variance and higher migration synchrony in spring lake arrival timing as compared to autumn lake departure timing. Furthermore, the timing of autumn migration is more variable across years than the timing of spring migration. Second, we find that later arrival to the lake habitat is positively associated with apparent survival from 1 year to the next, whereas we found no effect of lake departure timing on survival probability.These findings represent rare evidence showing how intraspecific variation in timing in migratory fish differs across years and seasons, and how variation in timing can translate into survival consequences for prey in systems characterised by high predation risk. [ABSTRACT FROM AUTHOR]

Migration is a widespread phenomenon, with powerful ecological and evolutionary consequences. Morphological adaptations to reduce the energetic costs associated with migratory transport are commonly documented for migratory species. However, few studies have investigated whether variation in body morphology can be explained by variation in migratory strategy within a species., We address this question in roach Rutilus rutilus, a partially migratory freshwater fish that migrates from lakes into streams during winter. We both compare body shape between populations that differ in migratory opportunity (open vs. closed lakes), and between individuals from a single population that vary in migratory propensity (migrants and residents from a partially migratory population). Following hydrodynamic theory, we posit that migrants should have a more shallow body depth, to reduce the costs associated with migrating into streams with higher flow conditions than the lakes the residents occupy all year round., We find evidence both across and within populations to support our prediction, with individuals from open lakes and migrants from the partially migratory population having a more slender, shallow-bodied morphology than fish from closed lakes and all-year residents., Our data suggest that a shallow body morphology is beneficial to migratory individuals and our study is one of the first to link migratory strategy and intraspecific variation in body shape. [ABSTRACT FROM AUTHOR]

In an experimental study we assessed if benthic bioturbating invertebrates affect the recruitment (hatching) of zooplankton from the sediment, and if this effect persists as differences in the zooplankton community in the water column, that is, if bioturbation quantitatively stimulates benthic–pelagic coupling. We investigated the effects of four different benthic invertebrates ( Asellus aquaticus, Chironomus plumosus, Tubifex tubifex in the presence or absence of the predator Sialis lutaria). In total, 45 zooplankton taxa hatched from the sediment and the hatching success of some of these was dependent on the species identity of the bioturbating invertebrate. The predator Sialis reduced the abundance of all three invertebrate species, but tended to positively influence the zooplankton recruitment rates, possibly through increasing the activity of the bioturbating invertebrates. The most striking effect of bioturbation on the hatching and pelagic zooplankton community properties was that, on average, 11% more species hatched in the Asellus treatment than in any other treatment. This was also mirrored in the zooplankton water column community where, on average, 7% more species established a viable population in treatments with Asellus as bioturbator. In a complementary field survey, Asellus was more common in littoral than in profundal sediments. Because Asellus strongly affected recruitment of zooplankton in our experiment, we argue that bioturbation may partly explain why recruitment of resting stages of both phyto- and zooplankton is generally higher in littoral than in profundal areas. [ABSTRACT FROM AUTHOR]

Migration is a widespread phenomenon in many ecosystems. Most often, studies on migration have focused on how migration strategies are dependent on ecological parameters, but little attention has been paid to the top-down effect of migration on ecosystem processes. Cyprinid fish in many European lakes undergo partial migration, where a part of the population leaves the lake and enters streams for the winter. In this study, we model the effect of partial migration by fish on lower trophic levels in a lake ecosystem. Our results suggest that spring phyto- and zooplankton dynamics, including occurrences of clear-water phases, can be related to the timing and magnitude of partial migration of planktivorous fish. From our results we conclude that partial migration can influence the dynamics of lower trophic levels in the ecosystem. Furthermore, we hypothesize that partial migration may affect the stability of alternative stable states and transitions between them. [ABSTRACT FROM AUTHOR]

Intraspecific interactions may increase or decrease foraging rates of individual consumers, and such facilitation or interference interactions should affect individual foraging economies as well as predator-prey processes at the population level. To mechanistically predict individual foraging performance, we need to investigate the effects of positive and negative interactions on separate foraging-cycle stages. We illustrate the importance and viability of examining the effects of facilitation and interference on different foraging-cycle stages using three piscivore species as a model system. We studied individual foraging behaviour when alone or in the presence of conspecifics, and show that northern pike foraging in the presence of conspecifics decrease attack frequencies and consumption rates, although no explicit agonistic behaviours were recorded. Pikeperch increase consumption rates in conspecific groups, possibly through a prey-mediated increase in capture success, as pikeperch showed no direct behavioural interactions. The actively cooperating Eurasian perch increase capture success and consumption rates in groups. The results demonstrate the need to combine behavioural studies of positive and negative effects of intraspecific interactions on foraging-cycle stages with quantifications of overall consumption rates. Pure behavioural observations may result in misinterpretations of the effects of interactions on foraging, while studies on consumption rates only would lack the mechanistic base of the obtained results. We also suggest that effects of intraspecific interactions during the foraging cycle should be incorporated in mechanistic models of facilitation and interference to elucidate the link between individual behaviours and higher-order processes. [ABSTRACT FROM AUTHOR]

We aimed to assess whether prey organisms with limited large-scale dispersal abilities are locally adapted to prevailing predator regimes by studying how chemical cues from predatory fish affected the behavior of Gammarus pulex (Amphipoda) from ponds with and without fish. We also examined, in the laboratory, the Fl generation from each pond by incubating them with or without cues from predatory fish. The potential benefits of a behavioral avoidance response were also assessed in an experiment in which G. pulex from the different ponds and incubations were exposed to fish predation. G. pulex from fish ponds increased their refuge use when exposed to fish cues, whereas populations from fishless ponds reduced their refuge use. The Fl generation responded similarly to their parents. Only the F1 generation from fish pond populations responded with more pronounced antipredatory behavior when raised in fish water. Moreover, both the original and the Fl generation of fish pond G. pulex survived longer when exposed to fish predation than those from fishless ponds, independent of whether they were raised in fish water or not (Fl). Our results suggest that the behavioral response to predator cues in G. pulex is an inherited trait, i.e., a local adaptation to prevailing predator regimes. [ABSTRACT FROM AUTHOR]

The propensity to kill and consume conspecifics (cannibalism) varies greatly between and within species, but the underlying mechanisms behind this variation remain poorly understood. A rich literature has documented that consistent behavioural variation is ubiquitous across the animal kingdom. Such inter-individual behavioural differences, sometimes referred to as personality traits, may have far-reaching ecological consequences. However, the link between predator personality traits and the propensity to engage in cannibalistic interactions remains understudied. Here, we first quantified personality in Eurasian perch (Perca fluviatilis), measured as activity (time spent moving) and sociability (time spent near conspecifics). We then gave perch of contrasting behavioural types the option to consume either conspecific or heterospecific (roach, Rutilus rutilus) prey. Individual perch characterized by a social-active behavioural phenotype (n = 5) selected roach before being cannibalistic, while asocial-inactive perch (n = 17) consumed conspecific and heterospecific prey evenly. Thus, asocial-inactive perch expressed significantly higher rates of cannibalism as compared to social-active individuals. Individual variation in cannibalism, linked to behavioural type, adds important mechanistic understanding to complex population and community dynamics, and also provides insight into the diversity and maintenance of animal personality. [ABSTRACT FROM AUTHOR]

Predation risk is often invoked to explain variation in stress responses. Yet, the answers to several key questions remain elusive, including the following: (1) how predation risk influences the evolution of stress phenotypes, (2) the relative importance of environmental versus genetic factors in stress reactivity and (3) sexual dimorphism in stress physiology. To address these questions, we explored variation in stress reactivity (ventilation frequency) in a post‐Pleistocene radiation of live‐bearing fish, where Bahamas mosquitofish (Gambusia hubbsi) inhabit isolated blue holes that differ in predation risk. Individuals of populations coexisting with predators exhibited similar, relatively low stress reactivity as compared to low‐predation populations. We suggest that this dampened stress reactivity has evolved to reduce energy expenditure in environments with frequent and intense stressors, such as piscivorous fish. Importantly, the magnitude of stress responses exhibited by fish from high‐predation sites in the wild changed very little after two generations of laboratory rearing in the absence of predators. By comparison, low‐predation populations exhibited greater among‐population variation and larger changes subsequent to laboratory rearing. These low‐predation populations appear to have evolved more dampened stress responses in blue holes with lower food availability. Moreover, females showed a lower ventilation frequency, and this sexual dimorphism was stronger in high‐predation populations. This may reflect a greater premium placed on energy efficiency in live‐bearing females, especially under high‐predation risk where females show higher fecundities. Altogether, by demonstrating parallel adaptive divergence in stress reactivity, we highlight how energetic trade‐offs may mould the evolution of the vertebrate stress response under varying predation risk and resource availability. [ABSTRACT FROM AUTHOR]

Patterns of animal migration and the ecological forces that shape them have been studied for centuries. Yet ecological impacts caused by the migration, such as altered predator–prey interactions and effects on community structure, remain poorly understood. This is to a large extent due to the scarcity of naturally replicated migration systems with negative controls, that is, ecosystems without migration. In this study, we tested whether partial migration of certain species within the overall prey community affects foraging ecology of top predators and thereby alters energy pathways in food webs. We carried out the study in independent replicated freshwater lake systems, four with and four without opportunity for prey migration. Specifically, we compared predator foraging mode in lakes where cyprinid prey fish perform seasonal partial migrations into connected streams with lakes lacking migratory opportunities for prey fish. We found clear seasonal bottom-up effects of prey migration on predators, including changes in size structure and total biomass of ingested prey, size-specific changes in littoral versus pelagic origin of diet, and a higher degree of feast-and-famine for predators in systems with migratory prey. Our analyses further showed that partially migratory prey species constitute a larger part of the prey community in systems that allow migration. Hence, prey migrations have important implications for predator foraging ecology and may cause seasonal shifts in the importance of their supporting energy pathways. We suggest that such bottom-up effects of partial migration may be a widespread phenomenon both in aquatic and in terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

Removal of cyprinid fish is a widely used biomanipulation tool to transform turbid shallow eutrophic lakes in north temperate regions into a clear water state. We here evaluate the removal of cyprinids from streams as a supplement to lake fishing. Since cyprinids often aggregate in high densities in lake inlet/outlet streams during winter migration, removal of fish in this space-confined habitat may be cost-efficient as compared to fish removal in the lake habitat. In two consecutive years, we annually removed up to 35% of the dominant cyprinids from an inlet stream to a lake and argue that this could easily be increased with a more targeted fishing effort. Concurrently, we monitored species- and length-specific variation in migration propensity, to explore how this relates to efficient fish removal. Smaller planktivores generally had a much higher migratory propensity than larger benthivores. Hence, stream fishing specifically targets species and size groups that are less efficiently controlled with traditional lake fishing methods. As a rule of thumb, stream fishing is most efficient when water temperature is 2-6°C. Prior to implementing fish removals from streams, the potential evolutionary consequences of the targeted removal of migratory phenotypes should be considered. [ABSTRACT FROM AUTHOR]

Whereas many studies have addressed the mechanisms driving partial migration, few have focused on the consequences of partial migration on trophic dynamics, and integrated studies combining the two approaches are virtually nonexistent. Here we show that temperature affects seasonal partial migration of cyprinid fish from lakes to predation refuges in streams during winter and that this migration in combination with temperature affects the characteristics and phenology of lower trophic levels in the lake ecosystem. Specifically, our six-year study showed that the proportion of fish migrating was positively related to lake temperature during the pre-migration growth period, i.e. during summer. Migration from the lake occurred later when autumn water temperatures were high, and timing of return migration to the lake occurred earlier at higher spring water temperatures. Moreover, the winter mean size of zooplankton in the lake increased with the proportion of fish being away from the lake, likely as a consequence of decreased predation pressure. Peak biomass of phytoplankton in spring occurred earlier at higher spring water temperatures and with less fish being away from the lake. Accordingly, peak zooplankton biomass occurred earlier at higher spring water temperature, but relatively later if less fish were away from the lake. Hence, the time between phyto- and zooplankton peaks depended only on the amount of fish being away from the lake, and not on temperature. The intensity of fish migration thereby had a major effect on plankton spring dynamics. These results significantly contribute to our understanding of the interplay between partial migration and trophic dynamics, and suggest that ongoing climate change may significantly affect such dynamics. [ABSTRACT FROM AUTHOR]

Partial migration, in which a fraction of a population migrate and the rest remain resident, occurs in an extensive range of species and can have powerful ecological consequences. The question of what drives differences in individual migratory tendency is a contentious one. It has been shown that the timing of partial migration is based upon a trade-off between seasonal fluctuations in predation risk and growth potential. Phenotypic variation in either individual predation risk or growth potential should thus mediate the strength of the trade-off and ultimately predict patterns of partial migration at the individual level (i.e. which individuals migrate and which remain resident). We provide cross-population empirical support for the importance of one component of this model--individual predation risk--in predicting partial migration in wild populations of bream Abramis brama, a freshwater fish. Smaller, high-risk individuals migrate with a higher probability than larger, low-risk individuals, and we suggest that predation risk maintains size-dependent partial migration in this system. [ABSTRACT FROM AUTHOR]

Animal movement is a multifaceted process that occurs for multiple reasons with powerful consequences for food web and ecosystem dynamics. New paradigms and technical innovations have recently pervaded the field, providing increasingly powerful means to deliver fine-scale movement data, attracting renewed interest. Specifically in the aquatic environment, tracking with acoustic telemetry now provides integral spatiotemporal information to follow individual movements in the wild. Yet, this technology also holds great promise for experimental studies, enhancing our ability to truly establish cause-and-effect relationships. Here, we argue that ponds with well-defined borders (i.e. "islands in a sea of land") are particularly well suited for this purpose. To support our argument, we also discuss recent experiences from studies conducted in an innovative experimental infrastructure, composed of replicated ponds equipped with modern aquatic telemetry systems that allow for unparalleled insights into the movement patterns of individual animals., (© 2023. BioMed Central Ltd., part of Springer Nature.)

Inducible defences allow prey to increase survival chances when predators are present while avoiding unnecessary costs in their absence. Many studies report considerable inter-individual variation in inducible defence expression, yet what underlies this variation is poorly understood. A classic vertebrate example of a predator-induced morphological defence is the increased body depth in crucian carp (Carassius carassius), which reduces the risk of predation from gape-size limited predators. Here, we report that among-individual variation in morphological defence expression can be linked to sex. We documented sexual dimorphism in lakes in which crucian carp coexisted with predators, where females showed shallower relative body depths than males, but not in a predator-free lake. When exposing crucian carp from a population without predators to perceived predation risk in a laboratory environment (presence/absence of pike, Esox lucius), we found that males expressed significantly greater morphological defence than females, causing sexual dimorphism only in the presence of predators. We uncovered a correlative link between the sex-specific inducible phenotypic response and gene expression patterns in major stress-related genes (POMC, MC3R, and MC4R). Together, our results highlight that sex-specific responses may be an important, yet underappreciated, component underlying inter-individual differences in the expression of inducible defences, even in species without pronounced sexual dimorphism., (© The Author(s) 2022. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE).)

Many species up- or downregulate their resting metabolic rate (RMR) when they encounter favourable or unfavourable feeding conditions, respectively. This is thought to promote faster growth when food is abundant and conserve energy reserves when food is scarce. The time it takes to express metabolic plasticity remain little studied. Here, we develop a conceptual model showing how rapid or slow metabolic plasticity alter growth trajectories in response to changes in food supply. We test predictions from the model in a food manipulation experiment with young-of-the-year northern pike, Esox lucius , a species that experience drastic changes in food supply in nature. We find that metabolic plasticity is expressed gradually over several weeks in this species. Rapid changes in food supply thus caused apparent trait-environment mismatches that persisted for at least five weeks. Contrary to predictions, pike grew faster at high food levels when they had previously experienced low food levels and downregulated their RMR. This was not owing to increases in food intake but probably reflected that low RMRs increased the energetic scope for growth when feeding conditions improved. This highlights the important but complex effects of metabolic plasticity on growth dynamics under variable resource levels on ecologically relevant time scales.

Due to their potential for affecting the modulation of behaviour, effects of selective serotonin reuptake inhibitors (SSRIs) in the environment are particularly interesting regarding interspecies interactions and non-consumptive effects (NCEs) induced by predator cues in prey organisms. We evaluated the effects of sertraline (0.4, 40 ng/L, 40 µg/L) over 8 days on activity and habitat choice in the freshwater snail Radix balthica, on snails' boldness in response to mechanical stimulation (simulating predator attack), and their activity/habitat choice in response to chemical cues from predatory fish. We hypothesised that sertraline exposure would detrimentally impact NCEs elicited by predator cues, increasing predation risk. Although there were no effects of sertraline on NCEs, there were observed effects of chemical cue from predatory fish on snail behaviour independent of sertraline exposure. Snails reduced their activity in which the percentage of active snails decreased by almost 50% after exposure to fish cue. Additionally, snails changed their habitat use by moving away from open (exposed) areas. The general lack of effects of sertraline on snails' activity and other behaviours in this study is interesting considering that other SSRIs have been shown to induce changes in gastropod behaviour. This raises questions on the modes of action of various SSRIs in gastropods, as well as the potential for a trophic "mismatch" of effects between fish predators and snail prey in aquatic systems.

Many prey organisms show induced morphological responses to predators including changes in protective spine length, such as in rotifers, although previous studies have mainly focused on how prey become larger than the predator gape-size optimum. Here we show that a large-sized predator makes prey rotifers escape below the gape-size optimum of the predator by reducing spine length. In experiments and field studies we show that during part of their ontogeny fish larvae feed intensively on the common rotifer Keratella cochlearis, and that larval fish predation reduces rotifer spine length both through induction of shorter spines and selective predation on long-spined individuals. We also describe a global scale pattern in spine length of K. cochlearis, showing an increasing variance in spine length with latitude. This pattern may be explained by differences in fish reproduction from once per year at high latitudes to several times per year at lower latitudes. That spine length is adaptively adjusted to the ontogeny of a dominant predator taxa provides a novel view on our understanding of factors affecting temporal and spatial variations in prey defense morphology., (© 2017 by the Ecological Society of America.)

Organisms display an impressive array of defence strategies in nature. Inducible defences (changes in morphology and/or behaviour within a prey's lifetime) allow prey to decrease vulnerability to predators and avoid unnecessary costs of expression. Many studies report considerable interindividual variation in the degree to which inducible defences are expressed, yet what underlies this variation is poorly understood. Here, we show that individuals differing in a key personality trait also differ in the magnitude of morphological defence expression. Crucian carp showing risky behaviours (bold individuals) expressed a significantly greater morphological defence response when exposed to a natural enemy when compared with shy individuals. Furthermore, we show that fish of different personality types differ in their behavioural plasticity, with shy fish exhibiting greater absolute plasticity than bold fish. Our data suggest that individuals with bold personalities may be able to compensate for their risk-prone behavioural type by expressing enhanced morphological defences.

Partial migration is a common phenomenon among many animals and occurs in many types of ecosystems. Understanding the mechanisms behind partial migration is of major importance for the understanding of population dynamics and, eventually, ecosystem processes. We studied the effects of food availability on the seasonal partial migration of cyprinid fish from a lake to connected streams during winter by the use of passive telemetry. Fish with increased access to food were found to migrate in higher proportion, earlier in the season, and to reside in the streams for a longer period compared to fish with decreased access to food. Furthermore, fewer unfed migrants returned to the lake, indicating higher overwinter mortality. Our results suggest that individual fish trade off safety from predation and access to food differently depending on their body condition, which results in a condition-dependent partial migration. Hence, our main conclusion is that individual decision-making is based on assessment of own condition which offers a mechanistic explanation to partial migration. Moreover, this may be of high importance for understanding population responses to environmental variation as well as ecosystem dynamics and stability.

Fishes show a remarkable diversity of shapes which have been associated with their swimming abilities and anti-predator adaptations. The crucian carp (Carassius carassius) provides an extreme example of phenotypic plasticity in body shape which makes it a unique model organism for evaluating the relationship between body form and function in fishes. In crucian carp, a deep body is induced by the presence of pike (Esox lucius), and this results in lower vulnerability to gape-limited predators, such as pike itself. Here, we demonstrate that deep-bodied crucian carp attain higher speed, acceleration and turning rate during anti-predator responses than shallow-bodied crucian carp. Therefore, a predator-induced morphology in crucian carp enhances their escape locomotor performance. The deep-bodied carp also show higher percentage of muscle mass. Therefore, their superior performance in escape swimming may be due to a combination of higher muscle power and higher thrust.

One of the most fundamental components of predator-prey models is encounter rate, modelled as the product of prey density and search efficiency. Encounter rates have, however, rarely been measured in empirical studies. In this study, we used a video system approach to estimate how encounter rates between piscivorous fish that use a sit-and-wait foraging strategy and their prey depend on prey density and environmental factors such as turbidity. We first manipulated prey density in a controlled pool and field enclosure experiments where environmental factors were held constant. In a correlative study of 15 freshwater lakes we then estimated encounter rates in natural habitats and related the results to both prey fish density and environmental factors. We found the expected positive dependence of individual encounter rates on prey density in our pool and enclosure experiments, whereas the relation between school encounter rate and prey density was less clear. In the field survey, encounter rates did not correlate with prey density but instead correlated positively with water transparency. Water transparency decreases with increasing prey density along the productivity gradient and will reduce prey detection distance and thus predator search efficiency. Therefore, visual predator-prey encounter rates do not increase, and may even decrease, with increasing productivity despite increasing prey densities.