Long-term anti-predator learning and memory differ across populations and sexes in an intertidal snail.

Anti-predator behaviours in response to predator cues can be innate, or they can be learned through prior experience and remembered over time. The duration and strength of continued anti-predator behaviour after predator cues are no longer present, and the potential for an enhanced response when re-exposed to predator cues later is less known but could account for the observed variation in anti-predator responses. We measured the carryover effects of past predation exposure and the potential for anti-predator learning and memory in the marine snail Nucella canaliculata from six populations distributed over 1000 km of coastline. We exposed lab-reared snails to cues associated with a common crab predator or seawater control in two serial experiments separated by over seven months. Responses were population- and sex-dependent, with some populations retaining anti-predator behaviours while others showed a capacity for learning and memory. Male snails showed a strong carryover of risk aversion, while females were able to return to normal feeding rates and grow more quickly. These behavioural differences culminated in strong impacts on feeding and growth rates, demonstrating that this variation has implications for the strength of trait-mediated indirect interactions, which can impact entire ecosystems.

Context-dependent multimodal behaviour in a coral reef fish.

Animals are expected to respond flexibly to changing circumstances, with multimodal signalling providing potential plasticity in social interactions. While numerous studies have documented context-dependent behavioural trade-offs in terrestrial species, far less work has considered such decision-making in fish, especially in natural conditions. Coral reef ecosystems host 25% of all known marine species, making them hotbeds of competition and predation. We conducted experiments with wild Ambon damselfish (Pomacentrus amboinensis) to investigate context-dependent responses to a conspecific intruder; specifically, how nest defence is influenced by an elevated predation risk. We found that nest-defending male Ambon damselfish responded aggressively to a conspecific intruder, spending less time sheltering and more time interacting, as well as signalling both visually and acoustically. In the presence of a model predator compared to a model herbivore, males spent less time interacting with the intruder, with a tendency towards reduced investment in visual displays compensated for by an increase in acoustic signalling instead. We therefore provide ecologically valid evidence that the context experienced by an individual can affect its behavioural responses and multimodal displays towards conspecific threats.

Seasonal activity patterns of a Kalahari mammal community: Trade-offs between environmental heat load and predation pressure.

Mammals in arid zones have to trade off thermal stress, predation pressure, and time spent foraging in a complex thermal landscape. We quantified the relationship between the environmental heat load and activity of a mammal community in the hot, arid Kalahari Desert. We deployed miniature black globe thermometers within the existing Snapshot Safari camera trap grid on Tswalu Kalahari Reserve, South Africa. Using the camera traps to record species' activity throughout the 24-h cycle, we quantified changes in the activity patterns of mammal species in relation to heat loads in their local environment. We compared the heat load during which species were active between two sites with differing predator guilds, one where lion (Panthera leo) biomass dominated the carnivore guild and the other where lions were absent. In the presence of lion, prey species were generally active under significantly higher heat loads, especially during the hot and dry spring. We suggest that increased foraging under high heat loads highlights the need to meet nutritional requirements while avoiding nocturnal activity when predatory pressures are high. Such a trade-off may become increasingly costly under the hotter and drier conditions predicted to become more prevalent as a result of climate change within the arid and semi-arid regions of southern Africa.

Noise constrains heterospecific eavesdropping more than conspecific reception of alarm calls.

Many vertebrates eavesdrop on alarm calls of other species, as well as responding to their own species' calls, but eavesdropping on heterospecific alarm calls might be harder than conspecific reception when environmental conditions make perception or recognition of calls difficult. This could occur because individuals lack hearing specializations for heterospecific calls, have less familiarity with them, or require more details of call structure to identify calls they have learned to recognize. We used a field playback experiment to provide a direct test of whether noise, as an environmental perceptual challenge, reduces response to heterospecific compared to conspecific alarm calls. We broadcast superb fairy-wren (Malurus cyaneus) and white-browed scrubwren (Sericornis frontalis) flee alarm calls to each species with or without simultaneous broadcast of ambient noise. Using two species allows isolation of the challenge of heterospecific eavesdropping independently of any effect of call structure on acoustic masking. As predicted, birds were less likely to flee to heterospecific than conspecific alarm calls during noise. We conclude that eavesdropping was harder in noise, which means that noise could disrupt information on danger in natural eavesdropping webs and so compromise survival. This is particularly significant in a world with increasing anthropogenic noise.

Continuous low-intensity predation by owls (Strix aluco) on bats (Nyctalus lasiopterus) in Spain and the potential effect on bat colony stability.

Owls prey on bats, but information on owl predation is scarce, its impact on bat mortality is unclear, and reports on behavioural responses, including roost-switching and fission-fusion behaviour, are equivocal. To study the link between owl predation and anti-predator behaviour in bats, we evaluated seven months of video recordings at roosts and the behaviour of 51 passive integrated transponder (PIT)-tagged bats and bats without tags in a geographically isolated colony of greater noctule bats (Nyctalus lasiopterus) in Spain. We found the tawny owl Strix aluco to almost continuously hunt N. lasiopterus, from perches and on the wing, well after the bats emerged at dusk and when they returned to their roosts. We recorded 39 unsuccessful and three successful attacks. Nonetheless, we found no evidence that owl predation modifies bat behaviour. While the bats constituted only a very small proportion of the owls' diet, owl predation accounted for an estimated 30-40% of bat mortality, which may have a significant impact on small, local or isolated bat populations, in particular, and thereby shape regional bat distributions. We hypothesize that low roost availability may also affect the bats' potential response to predation, which could lead to natural predation having an excessive impact on bat populations.

The effect of waterfowl signals and Pseudocorynosoma enrietti infection on the behaviour of the amphipod Hyalella patagonica.

In the present study, we sought to determine whether i) a waterfowl signal induces avoidance behaviour of the amphipod Hyalella patagonica, ii) infection by the acanthocephalan Pseudocorynosoma enrietti affects the behaviour of the amphipod, and iii) the parasite interferes with the amphipod response to waterfowl. We evaluated amphipod behaviour experimentally by measuring activity levels, phototaxis, geotaxis, and clinging behaviour. The main findings of this study indicate that uninfected amphipods show avoidance behaviour by reducing their activity in the presence of a predator signal. Secondly, infected amphipods show altered behaviour, such as swimming in bright areas near the water surface, which makes them more visible to predators in nature. Lastly, the presence of predatory cues causes infected amphipods to drop to the bottom, which increases their visibility to predators. The present research allows us to perceive the intricate interplay among predators, parasites, and their intermediate hosts and advance our understanding of these complex ecological dynamics.

Rain, recreation and risk: Human activity and ecological disturbance create seasonal risk landscapes for the prey of an ambush predator.

Predation risk and prey responses exhibit fluctuations in space and time. Seasonal ecological disturbances can alter landscape structure and permeability to influence predator activity and efficacy, creating predictable patterns of risk for prey (seasonal risk landscapes). This may create corresponding seasonal shifts in antipredator behaviour, mediated by species ecology and trade-offs between risk and resources. Yet, how human recreation interacts with seasonal risk landscapes and antipredator behaviour remains understudied. In South Florida, we investigated the impact of a seasonal ecological disturbance, specifically flooding, which is inversely related to human activity, on interactions between Florida panthers (Puma concolor coryi) and white-tailed deer (Odocoileus virginianus). We hypothesized that human activity and ecological disturbances would interact with panther-deer ecology, resulting in the emergence of two distinct seasonal landscapes of predation risk and the corresponding antipredator responses. We conducted camera trap surveys across southwestern Florida to collect detection data on humans, panthers and deer. We analysed the influence of human site use and flooding on deer and panther detection probability, co-occurrence and diel activity during the flooded and dry seasons. Flooding led to decreased panther detections and increased deer detections, resulting in reduced deer-panther co-occurrence during the flooded season. Panthers exhibited increased nocturnality and reduced diel activity overlap with deer in areas with higher human activity. Supporting our hypothesis, panthers' avoidance of human recreation and flooding created distinct risk schedules for deer, driving their antipredator behaviour. Deer utilized flooded areas to spatially offset predation risk during the flooded season while increasing diurnal activity in response to human recreation during the dry season. We highlight the importance of understanding how competing risks and ecological disturbances influence predator and prey behaviour, leading to the generation of seasonal risk landscapes and antipredator responses. We emphasize the role of cyclical ecological disturbances in shaping dynamic predator-prey interactions. Furthermore, we highlight how human recreation may function as a 'temporal human shield,' altering seasonal risk landscapes and antipredator responses to reduce encounter rates between predators and prey.

Guppies in large groups cooperate more frequently in an experimental test of the group size paradox.

The volunteer's dilemma, in which a single individual is required to produce a public good, predicts that individuals in larger groups will cooperate less frequently. Mechanistically, this could result from trade-offs between costs associated with volunteering and costs incurred if the public good is not produced (nobody volunteers). During predator inspection, one major contributor to the cost of volunteering is likely increased probability of predation; however, a predator also poses a risk to all individuals if nobody inspects. We tested the prediction that guppies in larger groups will inspect a predator less than those in smaller groups. We also predicted that individuals in larger groups would perceive less threat from the predator stimulus because of the protective benefits of larger groups (e.g. dilution). Contrary to prediction, we found that individuals in large groups inspected more frequently than those in smaller groups, but (as predicted) spent less time in refuges. There was evidence that individuals in intermediate-sized groups made fewest inspections and spent most time in refuges, suggesting that any link between group size, risk and cooperation is not driven by simple dilution. Extensions of theoretical models that capture these dynamics will likely be broadly applicable to risky cooperative behaviour.

Behavioural differences in predator aware and predator naïve Wellington tree weta, Hemideina crassidens.

Insects have evolved a wide range of behavioural traits to avoid predation, with anti-predator behaviours emerging as important adaptive responses to the specific strategies employed by predators. These responses may become ineffective, however, when a species is introduced to a novel predator type. When individuals cannot recognise an introduced predator for instance, they may respond in ways that mean they fail to avoid, escape, or neutralize a predator encounter. New Zealand's endemic insect fauna evolved in the absence of terrestrial mammalian predators for millions of years, resulting in the evolution of unique fauna like the large, flightless Orthopteran, the weta. Here we investigate how experience with introduced mammalian predators might influence anti-predator behaviours by comparing behaviours in a group of Wellington tree weta (Hemideina crassidens) living in an ecosanctuary, Zealandia, protected from non-native mammalian predators, and a group living in adjacent sites without mammalian predator control. We used behavioural phenotyping assays with both groups to examine rates of activity and defensive aggression shortly after capture, and again after a period of acclimation. We found that weta living in protected areas were more active shortly after capture than weta in non-protected habitats where mammalian predators were present. Male weta living in non-protected areas tended to be less aggressive than any other group. These results suggest that lifetime experience with differing predator arrays may influence the expression of antipredator behaviour in tree weta. Disentangling innate and experiential drivers of these behavioural responses further will have important implications for insect populations in rapidly changing environments.

Experimental test of the influence of light availability on the evolution of eye size and behaviour in Daphnia.

There exists extensive variation in eye size. Much work has provided a connection between light availability and differences in eye size across taxa. Experimental tests of the role of the light environment on the evolution of eye size are lacking. Here, we performed a selection experiment that examined the influence of light availability on shifts in eye size and the connection between eye size and phototactic (anti-predator) behaviour in Daphnia. We set-up replicate experimental populations of Daphnia, repeatedly evaluated phenotypic shifts in eye size during the ~50-day experiment, and performed a common garden experiment at the end of the experiment to test for evolutionary shifts in eye size and behaviour. Our phenotypic analyses showed that eye size rapidly diverged between the light treatments; relative eye size was consistently larger in the low versus high light treatments. Selection on eye size was also modified by variation in density as increases in Daphnia density favoured a larger eye. However, we did not observe differences in eye size between the light treatments following two generations of common garden rearing at the end of the experiment. We instead observed strong shifts in anti-predator behaviour. Daphnia from the low light treatment exhibited decreased phototactic responses to light. Our results show that decreased light relaxes selection on anti-predator behaviour. Such trends provide new insights into selection on eye size and behaviour.

Red deer Cervus elaphus blink more in larger groups.

Most animals need to spend time being vigilant for predators, at the expense of other activities such as foraging. Group-living animals can benefit from the shared vigilance effort of other group members, with individuals reducing personal vigilance effort as group size increases. Behaviors like active scanning or head lifting are usually used to quantify vigilance but may not be accurate measures of this. We suggest that measuring an animal's blinking rate gives a meaningful measure of vigilance: increased blinking implies reduced vigilance, as the animal cannot detect predators when its eyes are closed. We describe an observational study of a captive population of red deer, where we measured the blinking rates of individual deer from groups of differing sizes (where mean group size ranged between 1 and 42.7 individuals). We demonstrate that as group size increases in red deer, individuals increase their blink rate, confirming the prediction that vigilance should decrease. Blinking is a simple non-invasive measure and offers a useful metric for assessing the welfare of animals experiencing an increase in perceived predation risk or other stressors.

Latitudinal cline of death-feigning behaviour in a beetle (Tribolium castaneum).

Death-feigning behaviour is a phenomenon in which a prey is rendered motionless due to stimulation or threat from a predator. This anti-predator defence mechanism has been observed across numerous animal taxa and is considered adaptive in nature. However, longer durations of death feigning can result in decreased opportunities for feeding and reproduction, and therefore is often associated with fitness costs as compared to environments without predators. Differences have also been observed in the frequencies and durations of death feigning within populations, and these differences are thought to be influenced by the balance between survival and other fitness costs. Furthermore, this balance is predicted to vary in response to changes in environmental conditions. In this study, we examined the death feigning in 38 populations of the red flour beetle (Tribolium castaneum). Our results demonstrate that frequencies and durations of the death feigning in T. castaneum show geographical variations and a latitude cline, indicating that this behaviour is influenced by location as well as latitude. This study is the first to demonstrate the existence of a latitudinal cline in death feigning and suggests that death-feigning behaviour might have evolved in response to environmental factors that vary with latitude.

Beyond simple habituation: Anthropogenic habitats influence the escape behaviour of spur-winged lapwings in response to both human and non-human threats.

Habitat development may affect wildlife behaviour, favouring individuals or behaviours that cope better with perceived threats (predators). Bolder behaviours in human-dominated habitats (HDH; e.g. urban and rural settlements) may represent habituation specifically to humans, or a general reduction in predator-avoidance response. However, such carry-over effects across threat types (i.e. beyond humans) and phases of the escape sequence have not been well studied to date. Here we investigated escape behaviours of a locally common wader species, the spur-winged lapwing Vanellus spinosus. We assayed their flight initiation distance (FID) and subsequent escape behaviours in agricultural areas and in HDH. We found that lapwings in HDH were bolder, and that the difference was manifested in several phases of the predator-avoidance sequence (shorter FIDs, shorter distances fled, and a higher probability of escape by running vs. flying). When re-approached (by an observer) after landing, lapwings in HDH were also more repetitive in their FID than those in other habitats. To determine whether this apparent bolder behaviour in HDH areas is merely a consequence of habituation to humans or represents a broader behavioural change, we introduced an additional threat type-a remotely-operated taxidermic jackal ('Jack-Truck'). Finding bolder responses in the HDH to the human threat alone (and not to the Jack-Truck) could have supported the habituation hypothesis. In contrast, however, we found a bolder response in the HDH to both threat types, as well as a correlation between their FIDs across different sites. These bolder behaviours suggest that HDH impose a broader behavioural change on lapwings, rather than just simple habituation. Overall, our findings demonstrate how FID trials can reveal strong behavioural carry-over effects of HDH following human and non-human threats, including effects on the subsequent phases of escaping the predator. Further, FID assays may reveal consistent behavioural types when assessed under field conditions, and offer a direct way to differentiate among the various poorly understood and non-mutually exclusive mechanisms that lead to behavioural differences among organisms in HDH. The mechanistic perspective is essential for understanding how rapid urbanization impacts wildlife behaviour, populations, and the range of behaviours within them, even in species apparently resilient to such environmental changes.

Identifying the most effective behavioural assays and predator cues for quantifying anti-predator responses in mammals: a systematic review.

BACKGROUND: Mammals, globally, are facing population declines. Protecting and breeding threatened populations inside predator-free havens and translocating them back to the wild is commonly viewed as a solution. These approaches can expose predator-naïve animals to predators they have never encountered and as a result, many conservation projects have failed due to the predation of individuals that lacked appropriate anti-predator responses. Hence, robust ways to measure anti-predator responses are urgently needed to help identify naïve populations at risk, to select appropriate animals for translocation, and to monitor managed populations for changes in anti-predator traits. Here, we undertake a systematic review that collates existing behavioural assays of anti-predator responses and identifies assay types and predator cues that provoke the greatest behavioural responses. METHODS: We retrieved articles from academic bibliographic databases and grey literature sources (such as government and conservation management reports), using a Boolean search string. Each article was screened against eligibility criteria determined using the PICO (Population-Intervention-Comparator-Outcome) framework. Using data extracted from each article, we mapped all known behavioural assays for quantifying anti-predator responses in mammals and examined the context in which each assay has been implemented (e.g., species tested, predator cue characteristics). Finally, with mixed effects modelling, we determined which of these assays and predator cue types elicit the greatest behavioural responses based on standardised difference in response between treatment and control groups. REVIEW FINDINGS: We reviewed 5168 articles, 211 of which were eligible, constituting 1016 studies on 126 mammal species, a quarter of which are threatened by invasive species. We identified six major types of behavioural assays: behavioural focals, capture probability, feeding station, flight initiation distance, giving-up density, and stimulus presentations. Across studies, there were five primary behaviours measured: activity, escape, exploration, foraging, and vigilance. These behaviours yielded similar effect sizes across studies. With regard to study design, however, studies that used natural olfactory cues tended to report larger effect sizes than those that used artificial cues. Effect sizes were larger in studies that analysed sexes individually, rather than combining males and females. Studies that used 'blank' control treatments (the absence of a stimulus) rather than a treatment with a control stimulus had higher effect sizes. Although many studies involved repeat measures of known individuals, only 15.4% of these used their data to calculate measures of individual repeatability. CONCLUSIONS: Our review highlights important aspects of experimental design and reporting that should be considered. Where possible, studies of anti-predator behaviour should use appropriate control treatments, analyse males and females separately, and choose organic predator cues. Studies should also look to report the individual repeatability of behavioural traits, and to correctly identify measures of uncertainty (error bars). The review highlights robust methodology, reveals promising techniques on which to focus future assay development, and collates relevant information for conservation managers.

When aggressiveness could be too risky: linking personality traits and predator response in superb fairy-wrens.

Personality syndromes in animals may have adaptive benefits for survival. For example, while engaging in predator deterrence, reactive individuals tend to prioritise their own survival, while proactive individuals engage in riskier behaviours. Studies linking animal personality measured in captivity with individual fitness or behaviours in the wild are sparse, which is a gap in knowledge this study aims to address. We used playback experiments in superb fairy-wrens (Malurus cyaneus), a common Australian songbird with a cooperative breeding system, to assess whether three personality traits measured during short-term captivity correlated with behavioural responses in the wild to a perceived nest and adult predator, the grey currawong (Strepera versicolor). We used three standard measures of personality in birds: struggle responses to human handling (boldness), exploration during a novel environment test, and aggressiveness during a mirror presentation. Superb fairy-wrens showed a significantly stronger response to the predator playback than to the control (willie wagtail, Rhipidura leucophrys) playback, suggesting that they recognised the predator playback as a threat without any accompanying visual stimulus. Birds that attacked their mirror image during the mirror presentation and those that spent a moderate amount of time close to the mirror responded more strongly to predator playback (by approaching the speaker faster and closer, spending more time near the speaker, and being more likely to alarm call) compared to those with low aggressiveness or those that spent very short or long durations close to the mirror. Neither boldness nor exploration in the novel environment test predicted playback response. Our results align with a growing number of studies across species showing the importance of animal personalities as factors for fitness and survival.

Alarm cues and alarmed conspecifics: neural activity during social learning from different cues in Trinidadian guppies.

Learning to respond appropriately to novel dangers is often essential to survival and success, but carries risks. Learning about novel threats from others (social learning) can reduce these risks. Many species, including the Trinidadian guppy (Poecilia reticulata), respond defensively to both conspecific chemical alarm cues and conspecific anti-predator behaviours, and in other fish such social information can lead to a learned aversion to novel threats. However, relatively little is known about the neural substrates underlying social learning and the degree to which different forms of learning share similar neural mechanisms. Here, we explored the neural substrates mediating social learning of novel threats from two different conspecific cues (i.e. social cue-based threat learning). We first demonstrated that guppies rapidly learn about threats paired with either alarm cues or with conspecific threat responses (demonstration). Then, focusing on acquisition rather than recall, we discovered that phospho-S6 expression, a marker of neural activity, was elevated in guppies during learning from alarm cues in the putative homologue of the mammalian lateral septum and the preoptic area. Surprisingly, these changes in neural activity were not observed in fish learning from conspecific demonstration. Together, these results implicate forebrain areas in social learning about threat but raise the possibility that circuits contribute to such learning in a stimulus-specific manner.

Transcriptomic profiles of consistent risk-taking behaviour across time and contexts in European sea bass.

Bolder individuals have greater access to food sources and reproductive partners but are also at increased risk of predation. Boldness is believed to be consistent across time and contexts, but few studies have investigated the stability of this trait across variable environments, such as varying stress loads or long periods of time. Moreover, the underlying molecular components of boldness are poorly studied. Here, we report that boldness of 1154 European sea bass, evaluated using group risk-taking tests, is consistent over seven months and for individuals subjected to multiple environments, including a chronically stressful environment. Differences in risk-taking behaviour were further supported by differences observed in the responses to a novel environment test: shy individuals displayed more group dispersion, more thigmotaxic behaviour and lower activity levels. Transcriptomic analyses performed on extreme phenotypes revealed that bold individuals display greater expression for genes involved in social and exploration behaviours, and memory in the pituitary, and genes involved in immunity and responses to stimuli in the head kidney. This study demonstrates that personality traits come with an underpinning molecular signature, especially in organs involved in the endocrine and immune systems. As such, our results help to depict state-behaviour feedback mechanisms, previously proposed as key in shaping animal personality.

Sensory environment affects Icelandic threespine stickleback's anti-predator escape behaviour.

Human-induced changes in climate and habitats push populations to adapt to novel environments, including new sensory conditions, such as reduced visibility. We studied how colonizing newly formed glacial lakes with turbidity-induced low-visibility affects anti-predator behaviour in Icelandic threespine sticklebacks. We tested nearly 400 fish from 15 populations and four habitat types varying in visibility and colonization history in their reaction to two predator cues (mechano-visual versus olfactory) in high versus low-visibility light treatments. Fish reacted differently to the cues and were affected by lighting environment, confirming that cue modality and light levels are important for predator detection and evasion. Fish from spring-fed lakes, especially from the highlands (likely more diverged from marine fish than lowland fish), reacted fastest to mechano-visual cues and were generally most active. Highland glacial fish showed strong responses to olfactory cues and, counter to predictions from the flexible stem hypothesis, the greatest plasticity in response to light levels. This study, leveraging natural, repeated invasions of novel sensory habitats, (i) illustrates rapid changes in anti-predator behaviour that follow due to adaptation, early life experience, or both, and (ii) suggests an additional role for behavioural plasticity enabling population persistence in the face of frequent changes in environmental conditions.

The broken-wing display across birds and the conditions for its evolution.

The broken-wing display is a well-known and conspicuous deceptive signal used to protect birds' broods against diurnal terrestrial predators. Although commonly associated with shorebirds, it remains unknown how common the behaviour is across birds and what forces are associated with the evolution of the display. Here, we use the broken-wing display as a paradigmatic example to study the evolution of a behaviour across Aves. We show that the display is widespread: it has been described in 52 families spread throughout the phylogeny, suggesting that it independently evolved multiple times. Further, we evaluated the association with 16 ecological and life-history variables hypothesized to be related to the evolution of the broken-wing display. Eight variables were associated with the display. We found that species breeding farther from the equator, in more dense environments, with shorter incubation periods, and relatively little nest cover were more likely to perform the display, as were those in which only one parent incubates eggs, species that mob nest predators and species that are altricial or multi-brooded. Collectively, our comprehensive approach identified forces associated with the repeated evolution of this conspicuous display, thereby providing new insights into how deceptive behaviours evolve in the context of predator-prey interactions.

Beaked whales and state-dependent decision-making: how does body condition affect the trade-off between foraging and predator avoidance?

Body condition is central to how animals balance foraging with predator avoidance-a trade-off that fundamentally affects animal fitness. Animals in poor condition may accept greater predation risk to satisfy current foraging 'needs', while those in good condition may be more risk averse to protect future 'assets'. These state-dependent behavioural predictions can help interpret responses to human activities, but are little explored in marine animals. This study investigates the influence of body condition on how beaked whales trade-off foraging and predator avoidance. Body density (indicating lipid-energy stores) was estimated for 15 foraging northern bottlenose whales tagged near Jan Mayen, Norway. Composite indices of foraging (diving and echolocation clicks) and anti-predation (long ascents, non-foraging dives and silent periods reducing predator eavesdropping) were negatively related. Experimental sonar exposures led to decreased foraging and increased risk aversion, confirming a foraging/perceived safety trade-off. However, lower lipid stores were not related to a decrease in predator avoidance versus foraging, i.e. worse condition animals did not prioritize foraging. Individual differences (personalities) or reproductive context could offer alternative explanations for the observed state-behaviour relationships. This study provides evidence of foraging/predator-avoidance trade-offs in a marine top predator and demonstrates that animals in worse condition might not always take more risks.