Exposure to the insecticide, imidacloprid, impairs predator-recognition learning in damselfly larvae.

Many aquatic organisms use chemosensory information to learn about local predation threats, but contaminants in their environment may impair such cognitive processes. Neonicotinoids are a class of water-soluble systemic insecticides that have become a major concern in aquatic systems. In this study, we explored how a 10-day exposure to various concentrations (0, 0.1, 1.0, or 10.0 µg/L) of the neonicotinoid imidacloprid affects the learned recognition of predator odour by non-target damselfly larvae (Lestes spp). Unexposed larvae and those exposed to the low concentration (0.1 µg/L) demonstrated an appropriate learned response to a novel predator odour following a conditioning with the odour paired with chemical alarm cues. However, such learning failed to occur for larvae that were exposed to imidacloprid concentrations of 1.0 and 10.0 µg/L. Thus, either the cognitive processing of the chemical information was impaired or the chemistry of one or both of the conditioning cues was altered, making them ineffective for learning. In a second experiment, we found evidence for this latter hypothesis. In the absence of background imidacloprid exposure, larvae did not show significant learned responses to the predator odour when the conditioning cues were mixed with imidacloprid (initial pulse solution of 3.0 µg/L) at the start of conditioning (reaching a final concentration of 0.01 µg/L). These findings indicate that even low levels of imidacloprid can have important implications for chemosensory cognition of non-target species in aquatic environments.

Acquired predator recognition via epidermal alarm cues but not dietary alarm cues by isolated pupfish.

We tested whether Shoshone pupfish Cyprinodon nevadensis shoshone and Amargosa River pupfish C. n. amargosae respond behaviourally to conspecific chemical alarm cues released when epidermal tissue is damaged by a predator. We found that both subspecies reduced activity and vertical position in the water column in response to alarm cues. We then tested if pupfish can use alarm cue to acquire recognition of a novel predator. We trained pupfish with (1) water + odour of largemouth bass fed a diet of earthworms, (2) alarm cues from skin extract (epidermal alarm cues) + odour of bass fed a diet of earthworms, or (3) water + odour of bass fed a diet of pupfish (dietary alarm cues). Pupfish responded to epidermal alarm cues but not to dietary alarm cues. Pupfish were retested with the odour of bass that were fed an earthworm diet. Pupfish that had previously received epidermal alarm cues reduced vertical position and activity relative to the other two treatments. This is the first demonstration of acquired recognition of a novel predator by a pupfish, the first report of partial predator naiveté, and opens the possibility of predator-recognition training as a tool for management and conservation of endangered desert fishes.

Uncertainty about old information results in differential predator memory in tadpoles.

As information ages, it may become less accurate, resulting in increased uncertainty for decision makers. For example, chemical alarm cues (AC) are a source of public information about a nearby predator attack, and these cues can become spatially inaccurate through time. These cues can also degrade quickly under natural conditions, and cue receivers are sensitive to such degradation. Although numerous studies have documented predator-recognition learning from fresh AC, no studies have explored learning from aged AC and whether the uncertainty associated with this older information contributes to shortening the retention of learned responses (i.e. the 'memory window'). Here, we found that wood frog tadpoles, Lithobates sylvaticus, learned to recognize a novel odour as a predator when paired with AC aged under natural conditions for up to 1 h. However, only tadpoles conditioned with fresh AC were found to retain this learned response when tested 9 days after conditioning. These results support the hypothesis that the memory window is shortened by the uncertainty associated with older information, preventing the long-term costs of a learned association that was based on potentially outdated information.

Social dynamics impact scolding behaviour in captive groups of common ravens (Corvus corax).

BACKGROUND: Predator avoidance can have immense impacts on fitness, yet individual variation in the expression of anti-predator behaviour remains largely unexplained. Existing research investigating learning of novel predators has focused either on individuals or groups, but not both. Testing in individual settings allows evaluations of learning or personality differences, while testing in group settings makes it impossible to distinguish any such individual differences from social dynamics. In this study, we investigate the effect of social dynamics on individual anti-predator behaviour. We trained 15 captive ravens to recognize and respond to a novel experimental predator and then exposed them to this predator in both group and isolation settings across 1.5 years to tease apart individual differences from social effects and evaluate two hypotheses: (1) weaker anti-predator responses of some individuals in the group occurred, because they failed to recognize the experimental predator as a threat, leading to weak responses when separated, or (2) some individuals had learned the new threat, but their scolding intensity was repressed in the group trials due to social dynamics (such as dominance rank), leading to increased scolding intensity when alone. RESULTS: We found that dominance significantly influences scolding behaviour in the group trials; top-ranked individuals scold more and earlier than lower ranking ones. However, in the separation trials scolding duration is no longer affected by rank. CONCLUSIONS: We speculate that, while top-ranked individuals use their anti-predator responses to signal status in the group, lower-ranking ravens may be suppressed from, or are less capable of, performing intense anti-predator behaviour while in the group. This suggests that, in addition to its recruitment or predator-deterrent effects, alarm calling may serve as a marker of individual quality to conspecifics.

Preliminary evidence for one-trial social learning of vervet monkey alarm calling.

How do non-human primates learn to use their alarm calls? Social learning is a promising candidate, but its role in the acquisition of meaning and call usage has not been studied systematically, neither during ontogeny nor in adulthood. To investigate the role of social learning in alarm call comprehension and use, we exposed groups of wild vervet monkeys to two unfamiliar animal models in the presence or absence of conspecific alarm calls. To assess the learning outcome of these experiences, we then presented the models for a second time to the same monkeys, but now without additional alarm call information. In subjects previously exposed in conjunction with alarm calls, we found heightened predator inspection compared to control subjects exposed without alarm calls, indicating one-trial social learning of 'meaning'. Moreover, some juveniles (but not adults) produced the same alarm calls they heard during the initial exposure whereas the authenticity of the models had an additional effect. Our experiment provides preliminary evidence that, in non-human primates, call meaning can be acquired by one-trail social learning but that subject age and core knowledge about predators additionally moderate the acquisition of novel call-referent associations.

Disturbance cues function as a background risk cue but not as an associative learning cue in tadpoles.

Chemical information has an important role in the sensory ecology of aquatic species. For aquatic prey, chemical cues are a vital source of information related to predator avoidance and risk assessment. For instance, alarm cues are released by prey that have been injured by predators. In addition to providing accurate information about current risk, repeated exposure to alarm cues can elicit a fear response to novel stimuli (neophobia) in prey. Another source of chemical information is disturbance cues, released by prey that have been disturbed or harassed (but not injured) by a predator. While disturbance cues have received much less attention than alarm cues, they appear to be useful as an early warning signal of predation risk and have the potential to be used as a priming cue for learning. In this study, we used wood frog (Lithobates sylvaticus) tadpoles to test whether repeated exposure to disturbance cues during the embryonic stage can induce neophobic behaviour. Three weeks following repeated exposure to disturbance cues, tadpoles showed reduced activity when exposed to a novel odour, but they no longer displayed an antipredator response to disturbance cues. In a second experiment, we found that tadpoles failed to learn that a novel odour was dangerous following a pairing with disturbance cues, whereas alarm cues facilitated such learning. Our results add to the growing body of information about disturbance cues and provide evidence of their function as an embryonic risk cue but not as an associative learning cue.

Behavioral patterns in the early-stage antipredator response change after tail autotomy in adult wall lizards.

Autotomy is a drastic antipredator defense consisting of the voluntary shedding of a body part to escape from the predators. The loss of a body part may impair locomotion, feeding or mating, so animals may face a higher predation risk shortly after autotomy. Thus, until regeneration is completed, prey may adjust their behavior to reduce predation risk, and this could involve secondary costs. We assessed the effect of tail loss on the antipredator behavior of wall lizards (Podarcis muralis), comparing the behavior of tailed and tailless individuals exposed to a predatory snake (Coronella austriaca) scent, under controlled experimental conditions. Tailless lizards spent significantly more time performing behaviors with antipredatory significance (e.g., moving slowly), whereas tailed individuals performed exploratory walking for significantly more time. Moreover, tailless lizards spent more time basking, which probably increases the effectiveness of their cryptic design and decreases detection by predators. Lizards intensified the tongue flick rates when exposed to a pungent control or snake scents, as compared to their response to a neutral control. Besides, both tailed and tailless lizards intensified some aspects of their antipredator behavior (walking slowly and avoiding refuge use) when exposed to snake scent, which indicates discrimination of the smell of predatory snakes. Lizards decreased refuge use when exposed to predator scents, probably because the refuges are evaluated as unsafe due to a high concentration of snake scents. To conclude, our experiments showed that, after losing their tails, wall lizards modify their behavior in a way that likely minimizes predation risk.

Cowards or clever guys: an alternative nest defence strategy employed by shrikes against magpies.

Red-backed shrikes (Lanius collurio) show a substantial variability in their nest defence behaviour, which usually follows the rules of optimal parental behaviour, vigorously attacking egg and chick predators and only passively guarding against harmless animals. Nevertheless, shrikes hesitate to attack the Eurasian magpie (Pica pica), which specializes in plundering passerine nests. Our previous studies have suggested that this behaviour may be the result of an alternative defence strategy, relying on nest crypsis. To test this hypothesis, at the shrike nests, we presented a magpie dummy associated with playbacks drawing the predators' attention to the presence of the nest. We predicted that the presentation of a magpie dummy associated with shrike alarm calls moves the parents to action, causing them to chase the magpie away from the nest. We showed that the presence of a magpie dummy associated with shrike alarm calls elicits a significantly more active response in shrike parents compared to a magpie dummy associated with neutral song. Parents actively moved around the dummy and produced alarm calls; nevertheless, most of the tested pairs hesitated to attack the dummy. We may conclude that the low nest defence activity of shrike parents towards magpie dummy was partly the result of an alternative strategy, which may be cancelled out by alerting the predator to the location of the nest; nevertheless, shrikes seem to be afraid of the magpie and hesitate to attack it physically.

Untrained birds' ability to recognise predators with changed body size and colouration in a field experiment.

BACKGROUND: During recognition process, multiple parameters of the encountered stimulus may play a role. Previous studies with wild birds identified the importance of several salient features (e.g., eyes, beak, prominent elements of colouration) which birds use to recognise other bird species, such as predators or nest parasites. In the present study, we observed the responses of passerines visiting winter feeders to stimuli in the form of dummies of Eurasian sparrowhawk which were modified in body size and/or colouration but always carried the salient features of raptors (hooked beak, talons) and one species-specific feature of the sparrowhawk (yellow eyes). In the vicinity of a feeder, we placed a dummy of an unmodified sparrowhawk, life-sized sparrowhawk with pigeon, great tit, or robin colouration, a small, great tit-sized sparrowhawk dummy with unmodified or pigeon colouration, or an unmodified pigeon dummy, which functioned as a harmless control. Then we measured how it affected the number of visits. RESULTS: We found that birds were less afraid of small dummies regardless of their colouration than they were of life-sized raptor dummies or even the pigeon dummy. This contrasts with the results of a previous laboratory experiment where great tits' reaction to small dummies was comparably fearful to their response to life-size dummies. In our experiment, birds were also not afraid of life-sized dummies with modified colouration except for a robin-coloured dummy, which caused an equally significant fear reaction as an unmodified sparrowhawk dummy. It is likely that this dummy resembled the colouration of a male sparrowhawk closely enough to cause this effect. CONCLUSIONS: Based on our observations, we conclude that birds use contextual features to evaluate the size of other birds. Distance and familiar reference points seem to play an important part in this process.

Tone deaf: Association of an auditory stimulus with predation risk by zebrafish Danio rerio does not generalize to another auditory stimulus.

Predator recognition by small fishes can be acquired when chemical alarm cues released from damaged skin (by a predator attack) are paired with a novel stimulus, such as the appearance or odor of a predator. Once learned, fish can extend recognition of risk by generalizing to associate risk with additional stimuli that are similar to the conditioned novel stimulus. Here, we trained zebrafish to associate a novel auditory stimulus with predation risk, and then tested to see if they generalize risk to all sound stimuli or whether the conditioned response is limited to the sound frequency of the conditioning stimulus. We found that zebrafish Danio rerio readily associated risk of predation with Tone 1 (285 Hz), as evidenced by reduction in activity, increased time spent near the substratum and increased shelter use, but fish conditioned to fear Tone 1 completely ignored presentation of a second tone of 762 Hz. These data suggest that generalization does not occur as easily for auditory cues as they do for olfactory and visual cues, perhaps due to differences in the properties of sensory biology or the cognitive mechanisms that process information in different sensory modalities.

--Effect of multimodal cues from a predatory fish on refuge use and foraging on an amphidromous shrimp.

BACKGROUND: Prey can alter their behavior when detecting predator cues. Little is known about which sensory channel, number of channels, or the interaction among channels that shrimp species use to evaluate the threat from predators. The amphidromous shrimp Xiphocaris elongata has an induced defense, an elongated rostrum, where predatory fishes are present. We sought to test if kairomones or visual cues when presented singly from fish either eating flakes or shrimp, had more effect on altering the temporal feeding and refuge use patterns of long-rostrum (LR) X. elongata. We were also interested in elucidating potential interactions among cues when presented simultaneously in different combinations (kairomones + visual + mechanosensory, kairomones + alarm + visual, kairomones + alarm, kairomones + visual) on the same response variables. We expected that when presented alone kairomones will significantly increase refuge use and decrease foraging, particularly late at night, in comparison to visual cues alone, and that multiple cues when presented simultaneously will further increase refuge use and decrease foraging at night. METHODS: We exposed shrimp to individual or multiple cues from the predatory fish mountain mullet, Augonostomus monticola. We examined shrimp behavior with respect to refuge use and foraging activity during four time periods (after sunset, nighttime, sunrise, and sunset) in a 24-hour period. RESULTS: Shrimp presented fish visual and chemical cues singly did not differ from one another but differed from control shrimp (no cues) with respect to refuge use or foraging. The number of shrimp using refuge in the treatment with most cues (KVM: kairomones+ visual + mechanosensory) was higher than in all the treatments with less cues. A significant decline in foraging was observed when multiple cues were presented simultaneously. The highest number of shrimp foraged one hour after sunset and at nighttime. A significant interaction was observed between cue treatments and time periods, with shrimp in the KVM treatment foraging less and using more refuge late at night and at sunrise than shrimp in other treatments or time periods. CONCLUSIONS: The observation that fish chemical and visual cues when presented singly produced similar refuge use and foraging patterns was contrary to expectation and suggests that visual and chemical cues, when presented alone, provide redundant information to X. elongata with regards to predation threat. The significant increase in refuge use and reduction in foraging observed in the KVM treatment suggest multimodal signal enhancement in the perception of threat. This makes evolutionary sense in "noisy" environments, such as streams, where detection, localization, and intention of predators is much improved when cues are received through multiple sensory channels.

The specificity of sperm-mediated paternal effects in threespine sticklebacks.

Parental effects may help offspring respond to challenging environments, but whether parental exposure to different environmental challenges induces similar responses in offspring is largely unknown. We compared the offspring of threespine stickleback (Gasterosteus aculeatus) fathers who had been exposed to a potentially threatening stimulus (net), a native predator (sculpin), or who had been left unexposed (control). Relative to offspring of control fathers, offspring of sculpin-exposed fathers were more responsive (greater change in activity) to a simulated sculpin predator attack, while offspring of net-exposed fathers were less responsive (fewer antipredator behaviors) and showed altered stress responses compared to the control. To evaluate whether parental exposure primes offspring to respond to specific stimuli (e.g., offspring of net-exposed fathers respond most strongly to a net), we then exposed offspring of each paternal treatment to nets, native sculpin models, or non-native trout models. Paternal treatment did not influence offspring response to different stimuli; instead, offspring were generally more responsive to the native sculpin predator compared to nets or non-native trout predator, suggesting that sticklebacks have innate predator recognition of native predators. Collectively, these results underscore that, while parental exposure to non-ecologically relevant stressors elicits effects in intergenerational studies, these findings may not mirror those produced when parents encounter ecologically relevant stressors. Knowing that parental effects can be predator-specific furthers our understanding of the ways in which parental effects may evolve to be adaptive and suggests the potential for transgenerational plasticity to affect how animals respond to human induced environmental change, including non-native predators.

Relaxed predation theory: size, sex and brains matter.

Australia's wildlife is being considerably impacted by introduced mammalian predators such as cats (Felis catus), dogs (Canis lupus familiaris), and foxes (Vulpes vulpes). This is often attributed to native wildlife being naïve to these introduced predators. A systematic review of the literature reveals that native metatherians (body mass range 0.02-25 kg) do not recognise, and show relaxed antipredator behaviours towards, native and some introduced mammalian predators. Native eutherians (all with body mass < 2 kg), however, do appear to recognise and exhibit antipredator behaviours towards both native and introduced predators. Based on our findings, we propose a novel theory, the 'Relaxed Predation Theory'. Our new theory is based on the absence of large mammalian predators leading to reduced predation pressure in Australia during the past 40000-50000 years, and on three key differences between Australian metatherians and eutherians: size, sex, and brains. In light of this Relaxed Predation Theory, we make a number of recommendations for the conservation of Australian wildlife: (i) predator avoidance training of suitable species; (ii) exclusion fencing to exclude some, but not all, predators to facilitate the development of antipredator behaviours; (iii) captive breeding programs to prevent the extinction of some species; and (iv) reintroduction of Australia's larger predators, potentially to compete with and displace introduced predators. A more detailed understanding of the responses of Australian mammals to predators will hopefully contribute to the improved conservation of susceptible species.

Visual Cues Have a More Extensive Effect on Topmouth Gudgeon (Pseudorasbora parva) than Chemosensory Cues in Identifying Novel Predators.

Through population expansion and accidental or deliberate introduction, prey commonly encounter novel predators they had never seen before. Several studies have shown that animals can generalize their learned recognition of a familiar predator to novel ones according to predators' identical or similar features. This process in fish mainly depends on the visual and chemosensory cues they receive. However, there is a lack of understanding of the different effects of these two cues. Topmouth gudgeons (Pseudorasbora parva) that had never seen turtles were captured and used as the subjects, and three freshwater turtles of different genera were used as predators. Before and after using one turtle for predator training treatment of topmouth gudgeons, fish responses to visual and chemosensory cues of each turtle were tested and recorded, and it was found that predator training promoted topmouth gudgeons' recognition of the risks represented by visual cues of all three turtles and by chemosensory cues of the turtle that were used in training. These results further verify the generalization of predator recognition in fish and indicate that visual cues have a more extensive effect on fish than chemosensory cues in identifying novel predators, especially predators that are distantly related to the familiar threats.

Rapid Learning and Long-Term Memory for Dangerous Humans in Ravens (Corvus corax).

Like many predatory species, humans have pronounced individual differences in their interactions with potential prey: some humans pose a lethal threat while others may provide valuable resources. Recognizing individual humans would thus allow prey species to maximize potential rewards while ensuring survival. Previous studies on corvids showed they can recognize and remember individual humans. For instance, wild American crows produced alarm calls toward specifically masked humans up to 2.7 years after those humans had caught and ringed them while wearing that mask. However, individual behavior of the crows or the impact of social features on their responses, was hardly examined. Here, we studied predator learning and social effects on responses, using a similar method, in captive common ravens (Corvus corax). We investigated learning and the impact of key social components on individual reactions to artificial predators. Human experimenters wore two types of masks while walking past two raven aviaries. In four training trials, the "dangerous" mask was presented while carrying a dead raven, whereas the "neutral" mask was presented empty-handed. Between every training trial and in all following trials, we presented both masks without dead ravens. We assessed the subjects' (i) learning speed, (ii) selective long-term response, and (iii) potential effects of social dynamics on individual alarm calling frequency. Ravens learned quickly (often based on the first trial), and some individuals distinguished the dangerous from the neutral mask for the next 4 years. Despite having received the same amount and quality of exposure to the dangerous mask, we found pronounced individual differences in alarm calling that were fairly consistent across test trials in socially stable situations: dominance, but not sex explained individual differences in alarm responses, indicating the potential use of alarm calls as "status symbols." These findings fit to those in wild bird populations and dominant individuals signaling their quality. Changes in the individuals' participation and intensity of alarm calling coincided with changes in group composition and pair formation, further supporting the role of social context on ravens' alarm calling.

Parents know best: transgenerational predator recognition through parental effects.

In highly biodiverse systems, such as coral reefs, prey species are faced with predatory threats from numerous species. Recognition of predators can be innate, or learned, and can help increase the chance of survival. Research suggests that parental exposure to increased predatory threats can affect the development, behaviour, and ultimately, success of their offspring. Breeding pairs of damselfish (Acanthochromis polyacanthus) were subjected to one of three olfactory and visual treatments (predator, herbivore, or control), and their developing embryos were subsequently exposed to five different chemosensory cues. Offspring of parents assigned to the predator treatment exhibited a mean increase in heart rate two times greater than that of offspring from parents in herbivore or control treatments. This increased reaction to a parentally known predator odour suggests that predator-treated parents passed down relevant threat information to their offspring, via parental effects. This is the first time transgenerational recognition of a specific predator has been confirmed in any species. This phenomenon could influence predator-induced mortality rates and enable populations to adaptively respond to fluctuations in predator composition and environmental changes.

Zooming in on mechanistic predator-prey ecology: Integrating camera traps with experimental methods to reveal the drivers of ecological interactions.

Camera trap technology has galvanized the study of predator-prey ecology in wild animal communities by expanding the scale and diversity of predator-prey interactions that can be analysed. While observational data from systematic camera arrays have informed inferences on the spatiotemporal outcomes of predator-prey interactions, the capacity for observational studies to identify mechanistic drivers of species interactions is limited. Experimental study designs that utilize camera traps uniquely allow for testing hypothesized mechanisms that drive predator and prey behaviour, incorporating environmental realism not possible in the laboratory while benefiting from the distinct capacity of camera traps to generate large datasets from multiple species with minimal observer interference. However, such pairings of camera traps with experimental methods remain underutilized. We review recent advances in the experimental application of camera traps to investigate fundamental mechanisms underlying predator-prey ecology and present a conceptual guide for designing experimental camera trap studies. Only 9% of camera trap studies on predator-prey ecology in our review use experimental methods, but the application of experimental approaches is increasing. To illustrate the utility of camera trap-based experiments using a case study, we propose a study design that integrates observational and experimental techniques to test a perennial question in predator-prey ecology: how prey balance foraging and safety, as formalized by the risk allocation hypothesis. We discuss applications of camera trap-based experiments to evaluate the diversity of anthropogenic influences on wildlife communities globally. Finally, we review challenges to conducting experimental camera trap studies. Experimental camera trap studies have already begun to play an important role in understanding the predator-prey ecology of free-living animals, and such methods will become increasingly critical to quantifying drivers of community interactions in a rapidly changing world. We recommend increased application of experimental methods in the study of predator and prey responses to humans, synanthropic and invasive species, and other anthropogenic disturbances.

Brown Mouse Lemurs (Microcebus rufus) May Lack Opportunities to Learn about Predator Calls.

Diurnal primates commonly modify their behaviour in response to predator vocalizations, but little is known about how nocturnal primates perceive predator calls. Mouse lemurs do not innately perceive predator calls as dangerous but may learn to do so in the wild. I hypothesized that wild brown mouse lemurs (Microcebus rufus) in Ranomafana National Park, Madagascar, differentiate between predator and non-predator vocalizations and exposed 29 adult subjects to calls of 3 sympatric predators (goshawk, fossa and owl), 3 sympatric non-predators and 4 controls. Brown mouse lemurs did not differentiate among stimuli, and there was no evidence that any recording was associated with danger, as neither activity nor spatial preferences changed in response to a stimulus, nor did subjects exhibit obvious behaviours indicative of danger such as flight or position on the branches. The failure to perceive predator calls as dangerous is common among small nocturnal solitary prey species and could be explained by a lack of learning opportunities. In social species acquired behaviours can spread quickly by social learning. Solitary species like mouse lemurs have to individually learn to associate predator calls with danger which is dependent on encounter rates. Unlike diurnal prey mouse lemurs lack the opportunity to observe diurnal raptors and associate their calls with the bird, and owls are proficient predators, which are difficult to observe in a rain forest where they hunt cryptically from perches. While sometimes nocturnal, the fossa is solitary and largely silent, and is also particularly rare at my study site. Cryptic, rare or non-vocal predators provide prey with few opportunities to associate their calls with danger. Mouse lemurs are subject to high predation rates by a wide range of predators and need to balance the costs and benefits of antipredation behaviours. Their cryptic lifestyle and preference for dense vegetation lowers the likelihood of being detected by predators, and they utilize visual and olfactory predator cues to make decisions about risk. Furthermore, their high reproductive output compensates for the high predation rates.

Alarm cue-mediated response and learning in zebrafish larvae.

We investigated the behavioural and learning response of zebrafish larvae to chemicals released by injured conspecifics (the alarm cue). Many aquatic vertebrates and invertebrates exhibit an innate antipredator response to alarm cues because in nature, they reliably indicate the presence of predators. Likewise, when an individual simultaneously perceives a novel odour and alarm cue, it learns to recognise the novel odour as a predator odour. Alarm cue-mediated behavioural response and learning have been reported in some fish and amphibians during early ontogeny, but in zebrafish, they have been described only for adults. In this study, we demonstrated that zebrafish at 12 and 24 days post fertilization exhibited reduced activity when exposed to alarm cue obtained by homogenised larvae of the same age, with this response being greater for the older zebrafish. In addition, we showed that 24-dpf zebrafish conditioned with alarm cue plus a novel odour learned to recognise the novel odour as a threat and responded to it with antipredator behaviour. The innate behavioural response and the learned response after conditioning may be used to develop paradigms with which to study anxiety, fear, stress, learning and memory in zebrafish larvae.

Predator-induced changes of male and female mating preferences: innate and learned components.

While many mating preferences have a genetic basis, the question remains as to whether and how learning/experience can modify individual mate choice decisions. We used wild-caught (predator-experienced) and F1 laboratory-reared (predator-naïve) invasive Western mosquitofish Gambusia affinis from China to test whether mating preferences (assessed in a first mate choice test) would change under immediate predation threat. The same individuals were tested in a second mate choice test during which 1 of 3 types of animated predators was presented: 1) a co-occurring predator, 2) a co-evolved but not currently co-occurring predator, and 3) a non-piscivorous species as control. We compared preference scores derived from both mate choice tests to separate innate from experiential effects of predation. We also asked whether predator-induced changes in mating preferences would differ between sexes or depend on the choosing individual's personality type and/or body size. Wild-caught fish altered their mate choice decisions most when exposed to the co-occurring predator whereas laboratory-reared individuals responded most to the co-evolved predator, suggesting that both innate mechanisms and learning effects are involved. This behavior likely reduces individuals' risk of falling victim to predation by temporarily moving away from high-quality (i.e., conspicuous) mating partners. Accordingly, effects were stronger in bolder than shyer, large- compared with small-bodied, and female compared with male focal individuals, likely because those phenotypes face an increased predation risk overall. Our study adds to the growing body of literature appreciating the complexity of the mate choice process, where an array of intrinsic and extrinsic factors interacts during decision-making.