Antipredator Responses Research Articles

Running away or running to? Do prey make decisions solely based on the landscape of fear or do they also include stimuli from a landscape of safety?

Predator-prey interactions are a key part of ecosystem function, and non-consumptive effects fall under the landscape of fear theory. Under the landscape of fear, the antipredator responses of prey are based on the spatial and temporal distribution of predatory cues in the environment. However, the aversive stimuli (fear) are not the only stimuli prey can utilize when making behavioral decisions. Prey might also be using attractive stimuli that represent safety to guide decision making. Using a novel, orthogonal design, we were able to spatially separate aversive and attractive stimuli to determine whether prey are utilizing safety cues to navigate their environment. Crayfish Faxonius rusticus were placed in the center of a behavioral arena. Aversive stimuli of either predatory bass Micropterus salmoides cues or conspecific alarm cues increased along the x-axis of the behavioral arena. Safety cues (shelters) increased along the y-axis by decreasing the number of shelter openings in this direction. Crayfish were allowed two phases to explore the arena: one without the fearful stimuli and one with the stimuli. Linear mixed models were conducted to determine whether movement behaviors and habitat utilization were affected by the phase of the trial and the type of aversive stimuli. Crayfish responded more strongly to alarm cues than to fear cues, with only alarm cues significantly impacting habitat utilization. When responding to alarm cues, crayfish used safety cues as well as fear cues to relocate themselves within the arena. Based on these results, we argue that crayfish utilize a landscape of safety in conjunction with a landscape of fear when navigating their environment.

Visual recognition and coevolutionary history drive responses of amphibians to an invasive predator

Abstract During biotic invasions, native prey are abruptly exposed to novel predators and are faced with unprecedented predatory pressures. Under these circumstances, the lack of common evolutionary history may hamper predator recognition by native prey, undermining the expression of effective antipredator responses. Nonetheless, mechanisms allowing prey to overcome evolutionary naïveté exist. For instance, in naïve prey, history of coevolution with similar native predators or detection of general traits characterizing predators can favor the recognition of stimuli released by invasive predators. However, few studies have assessed how these mechanisms shape prey response at the community level. Here, we evaluated behavioral responses in naïve larvae of 13 amphibian species to chemical and visual cues associated with an invasive predator, the American red swamp crayfish (Procambarus clarkii). Moreover, we investigated how variation among species responses was related to their coexistence with similar native crayfish predators. Amphibian larvae altered their behavior in presence of visual stimuli of the alien crayfish, while chemical cues elicited feeble and contrasting behavioral shifts. Activity reduction was the most common and stronger response, whereas some species exhibited more heterogeneous strategies also involving distancing and rapid escape response. Interestingly, species sharing coevolutionary history with the native crayfish were able to finely tune their response to the invasive one, performing bursts to escape. These results suggest native prey can respond to invasive predators through recognition of generic risk cues (e.g., approaching large shapes), still the capability of modulating antipredator strategies may also depend on their coevolutionary history with similar native predators.

Plasticity and flexibility in the anti-predator responses of treefrog tadpoles

Tadpoles can respond to perceived predation risk by adjusting their life history, morphology, and behavior in an adaptive way. Adaptive phenotypic plasticity can evolve by natural selection only if there is variation in reaction norms and if this variation is, at least in part, heritable. To provide insights into the evolution of adaptive phenotypic plasticity, we analyzed the environmental and parental components of variation in predator-induced life history (age and size at metamorphosis), morphology (tail depth), and behavior of Italian treefrog tadpoles (Hyla intermedia). Using an incomplete factorial design, we raised tadpoles either with or without caged predators (dragonfly larvae, gen. Aeshna) and, successively, we tested them in experimental arenas either with or without caged predators. Results provided strong evidence for an environmental effect on all three sets of characters. Tadpoles raised with caged predators (dragonfly larvae, gen. Aeshna) metamorphosed earlier (but at a similar body size) and developed deeper tails than their fullsib siblings raised without predators. In the experimental arenas, all tadpoles, independent of their experience, flexibly changed their activity and position, depending on whether the cage was empty or contained the predator. Tadpoles of the two experimental groups, however, showed different responses: those raised with predators were always less active than their predator-naive siblings and differences slightly increased in the presence of predators. Besides this strong environmental component of phenotypic variation, results provided evidence also for parental and parental-by-environment effects, which were strong on life-history, but weak on morphology and behavior. Interestingly, additive parental effects were explained mainly by dams. This supports the hypothesis that phenotypic plasticity might mainly depend on maternal effects and that it might be the expression of condition-dependent mechanisms.Significance statementAnimals, by plastically adjusting their phenotypes to the local environments, can often sensibly improve their chances of survival, suggesting the hypothesis that phenotypic plasticity evolved by natural selection. We test this hypothesis in the Italian treefrog tadpoles, by investigating the heritable variation in the plastic response to predators (dragonfly larvae). Using an incomplete factorial common-garden experiment, we showed that tadpoles raised with predators metamorphosed earlier (but at similar body size), developed deeper tails, and were less active than their siblings raised without predators. The plastic response varied among families, but variation showed a stronger maternal than paternal component. This suggests that plasticity might largely depend on epigenetic factors and be the expression of condition-dependent mechanisms.

Noisy waters can influence young-of-year lobsters’ substrate choice and their antipredatory responses

Offshore human activities lead to increasing amounts of underwater noise in coastal and shelf environments, which may affect commercially-important benthic invertebrate groups like the re-stocked Helgoland European lobster (Homarus gammarus) in the German Bight (North Sea). It is crucial to understand the impact tonal low-frequency noises, like maritime transport and offshore energy operations, may have on substrate choice and lobsters' behavior to assess potential benefits or bottlenecks of new hard-substrate artificial offshore environments that become available. In this study, we investigated the full factorial effect of a tonal low-frequency noise and predator presence on young-of-year (YOY) European lobsters' in a diurnal and nocturnal experiment. Rocks and European oyster shells (Ostrea edulis) were offered as substrate to YOY lobsters for 3 h. Video recordings (n = 134) allowed the identification of lobsters' initial substrate choice, diel activity and key behaviors (peeking, shelter construction, exploration and hiding). To ensure independence, YOY lobsters in the intermolt stage were randomly selected and assigned to the experimental tanks and used only once. We provide the first evidence that stressors alone, and in combination, constrain YOY lobsters' initial substrate choice towards rocks. During nighttime, the joint effect of exposure to a constant low-frequency noise and predator presence decreased antipredator behavior (i.e., hiding) and increased exploration behavior. Noise may thus interfere with YOY lobsters' attention and decision-making processes. This outcome pinpoints that added tonal low-frequency noise in the environment have the potential to influence the behavior of early-life stages of European lobsters under predator pressure and highlights the importance of including key benthic invertebrates' community relationships in anthropogenic noise risk assessments. Among others, effects of noise must be taken into consideration in plans involving the multi-use of any offshore area for decapods’ stock enhancement, aquaculture, and temporary no-take zones.

Effects of a group-living experience on the antipredator responses of individual tadpoles

The tendency to aggregate during the larval stage is widespread and highly variable among anuran species. Several studies have highlighted the link between tadpole group density and their activity level, confirming that, usually, living in groups brings several antipredator benefits. However, nearly all studies have focused on the average behavioural responses of tadpoles tested in groups. In this study, we explored the effects of living in groups of three different sizes (1, 5 and 25 individuals per group) on the antipredator behaviour of individual green toad, Bufotes balearicus, tadpoles. We first assessed their basal activity and then examined changes in mobility rate and total distance after exposure to the chemical cues of predatory dragonfly, Aeshna cyanea, larvae. For both the pre- and poststimulus activity levels, we also tested the effects of the presence of conspecifics' chemical cues in the experimental tub. Our results showed that (1) a previous brief (8 days) experience of group living is sufficient to affect the basal level of activity of individual tadpoles, which increased with group size; (2) tadpoles that were reared alone did not lower their activity further when exposed to predators' odour; (3) the antipredator response of high-density-reared tadpoles decreased in the presence of conspecifics’ cues, supporting the so-called dilution effect, which, anyway, may need a minimum group size to be apparent. We conclude that both previous group-rearing experience and current perception of the surrounding environment may affect antipredator behaviour in individual tadpoles.

Reactive anti-predator behavioral strategy shaped by predator characteristics.

Large mammalian herbivores use a diverse array of strategies to survive predator encounters including flight, grouping, vigilance, warning signals, and fitness indicators. While anti-predator strategies appear to be driven by specific predator traits, no prior studies have rigorously evaluated whether predator hunting characteristics predict reactive anti-predator responses. We experimentally investigated behavioral decisions made by free-ranging impala, wildebeest, and zebra during encounters with model predators with different functional traits. We hypothesized that the choice of response would be driven by a predator’s hunting style (i.e., ambush vs. coursing) while the intensity at which the behavior was performed would correlate with predator traits that contribute to the prey’s relative risk (i.e., each predator’s prey preference, prey-specific capture success, and local predator density). We found that the choice and intensity of anti-predator behaviors were both shaped by hunting style and relative risk factors. All prey species directed longer periods of vigilance towards predators with higher capture success. The decision to flee was the only behavior choice driven by predator characteristics (capture success and hunting style) while intensity of vigilance, frequency of alarm-calling, and flight latency were modulated based on predator hunting strategy and relative risk level. Impala regulated only the intensity of their behaviors, while zebra and wildebeest changed both type and intensity of response based on predator traits. Zebra and impala reacted to multiple components of predation threat, while wildebeest responded solely to capture success. Overall, our findings suggest that certain behaviors potentially facilitate survival under specific contexts and that prey responses may reflect the perceived level of predation risk, suggesting that adaptive functions to reactive anti-predator behaviors may reflect potential trade-offs to their use. The strong influence of prey species identity and social and environmental context suggest that these factors may interact with predator traits to determine the optimal response to immediate predation threat.

The \u201cright\u201d side of sleeping: laterality in resting behaviour of Aldabra giant tortoises (Aldabrachelys gigantea)

Although some studies investigated lateralization in reptiles, little research has been done on chelonians, focusing only on few behaviours such as righting response and escape preference. The aim of this study was to investigate lateralization in Aldabra giant tortoises (Aldabrachelys gigantea), focusing on asymmetrical positioning of the limbs and the head during resting behaviour, called sleep-like behaviour, involving both wild tortoises and individuals under human care. Subjects of the study were 67 adult Aldabra tortoises (54 free ranging on Curieuse, 13 under human care in Mahè Botanical Garden). For each tortoise observed during sleep-like behaviour, we recorded the position of the head (on the left, on the right or in line with the body midline) and we collected which forelimb and hindlimb were kept forward. Moreover, the number of subjects in which limbs were in a symmetrical position during the sleep-like behaviour was recorded. Based on our results, the number of tortoises with asymmetrical position of head and limb was higher (head: 63%; forelimbs: 88%; hindlimbs: 70%) than the number of tortoises with symmetrical position of the head and the limb. Regarding the head, throughout the subjects found with the asymmetrical position of the head during sleep-like behaviour, tortoises positioning the head on the right (42%) were more than those sleeping with the head on the left (21%). We found a relationship between the position of the forelimbs and hindlimbs during sleep-like behaviour. We reported no differences between Mahè (under human care) and Curieuse (wild) tortoises. Findings of this preliminary study underlined traces of group-level lateralization in head positioning during the sleep-like behaviour, possibly due to a left-eye/right-hemisphere involvement in anti-predatory responses and threatening stimuli as reported in reptiles and other vertebrates. This study aims at adding data on brain lateralization, often linked to lateralized behaviours, in reptiles, especially in chelonians.

Raised by aliens: constant exposure to an invasive predator triggers morphological but not behavioural plasticity in a threatened species tadpoles

During biotic invasions, native communities are abruptly exposed to novel and often severe selective pressures. The lack of common evolutionary history with invasive predators can hamper the expression of effective anti-predator responses in native prey, potentially accelerating population declines. Nonetheless, rapid adaptation and phenotypic plasticity may allow native species to cope with the new ecological pressures. We tested the hypothesis that phenotypic plasticity is fostered when facing invasive species and evaluated whether plasticity offers a pool of variability that might help the fixation of adaptive phenotypes. We assessed behavioural and morphological trait variation in tadpoles of the Italian agile frog (Rana latastei) in response to the invasive crayfish predator, Procambarus clarkii, by rearing tadpoles under different predation-risk regimes: non-lethal crayfish presence and crayfish absence. After two-month rearing, crayfish-exposed tadpoles showed a plastic shift in their body shape and increased tail muscle size, while behavioural tests showed no effect of crayfish exposure on tadpole behaviour. Furthermore, multivariate analyses revealed weak divergence in morphology between invaded and uninvaded populations, while plasticity levels were similar between invaded and uninvaded populations. Even if tadpoles displayed multiple plastic responses to the novel predator, none of these shifts underwent fixation after crayfish arrival (10–15 years). Overall, these findings highlight that native prey can finely tune their responses to invasive predators through plasticity, but the adaptive value of these responses in whitstanding the novel selective pressures, and the long-term consequences they can entail remain to be ascertained.

Habituation in anuran tadpoles and the role of risk uncertainty.

The ability to learn in the context of predation allows prey to respond to threats by adjusting their behavior based on specific information acquired from their current environment. Habituation is a process that allows animals to adapt to environmental changes. Very little is known about habituation in wild animals in general and there are no studies on habituation in anuran tadpoles in particular. Here, we performed three experiments to investigate the behavioral response of predator naïve Pleurodema thaul tadpoles to repeated stimulation with two predation risk cues (injured conspecific and predator fed cues) which a priori provide different information regarding risk. Experiment 1 showed that P. thaul tadpoles habituate the antipredator response when undergo predation risk chemical cues from injured conspecific and that response is long term. Experiment 2 showed that P. thaul tadpoles did not habituate their antipredator response when exposed to cues derived from an event of nymph odonate preying on P. thaul tadpoles (predator fed cues). Experiment 3 specifically evaluated the risk imposed by each of the risk cues used in Experiment 1 and Experiment 2 and showed that the degree of perceived risk in tadpoles appear to be similar in a single experience with any risk stimuli. We suggest that the behavioral habituation of tadpoles in the context of predation could be modulated by the level of uncertainty associated with risk stimuli.

Food availability and long-term predation risk interactively affect antipredator response.

Food availability and temporal variation in predation risk are both important determinants of the magnitude of antipredator responses, but their effects have rarely been examined simultaneously, particularly in wild prey. Here, we determine how food availability and long-term predation risk affect antipredator responses to acute predation risk by monitoring the foraging response of free-ranging snowshoe hares (Lepus americanus) to an encounter with a Canada lynx (Lynx canadensis) in Yukon, Canada, over four winters (2015-2016 to 2018-2019). We examined how this response was influenced by natural variation in long-term predation risk (2-month mortality rate of hares) while providing some individuals with supplemental food. On average, snowshoe hares reduced foraging time up to 10h after coming into close proximity (≤75m) with lynx, and reduced foraging time an average of 15.28±7.08min per lynx encounter. Hares tended to respond more strongly when the distance to lynx was shorter. More importantly, the magnitude of hares' antipredator response to a lynx encounter was affected by the interaction between food-supplementation and long-term predation risk. Food-supplemented hares reduced foraging time more than control hares after a lynx encounter under low long-term risk, but decreased the magnitude of the response as long-term risk increased. In contrast, control hares increased the magnitude of their response as long-term risk increased. Our findings show that food availability and long-term predation risk interactively drive the magnitude of reactive antipredator response to acute predation risk. Determining the factors driving the magnitude of antipredator responses would contribute to a better understanding of the indirect effects of predators on prey populations.

Qingbo, a common cyprinid fish, responds diversely in behavior and locomotion to predators with different hunting modes.

Almost all prey live in habitats with predators with different hunting modes; however, most studies on predation have investigated the effects of only one predator at a time. In this study, we aimed to investigate whether qingbo (Spinibarbus sinensis), a common cyprinid fish, responds differently to active hunting and ambush predators and how qingbo responds when both types of predators coexist. Juvenile qingbo were subjected to catfish (Clarias fuscus, active hunter) exposure, snakehead fish (Channa argus, ambush hunter) exposure, or mixed predator exposure (catfish and snakehead coexistence) for a duration of 60days. Then, their growth, behaviors, swimming performance, and metabolism were measured. Qingbo subjected to active hunting predator exposure exhibited decreased activity and predator inspection and improved fast-start escape performance compared to those in the control group. However, none of the parameters of the fish subjected to ambush predator exposure changed significantly. Fish subjected to mixed predator exposure exhibited improved fast-start escape performance but increased maintenance energy expenditure, whereas no changes were observed in any of the behavioral variables. Qingbo showed a stronger anti-predator response to active hunting predators than to ambush predators, suggesting that the fish exhibit a stronger anti-predator response to a current direct threat than to a potential threat (a predator exists nearby but seldom presents in attack behavior). Additionally, the response of prey fish to multiple predators was quite complex, and the coexistence and interaction of multiple predator species with different hunting modes may lead to serious stress responses and confound the prey's behavioral responses to each predator.

Acoustic and visual stimuli combined promote stronger responses to aerial predation in fish.

Bird predation poses a strong selection pressure on fish. Since birds must enter the water to catch fish, a combination of visual and mechano-acoustic cues (multimodal) characterize an immediate attack, while single cues (unimodal) may represent less dangerous disturbances. We investigated whether fish could use this information to distinguish between non-threatening and dangerous events and adjust their antipredator response to the perceived level of risk. To do so, we investigated the antipredator behavior of the sulphur molly (Poecilia sulphuraria), a small freshwater fish which is almost exclusively preyed on by piscivorous birds in its endemic sulfide spring habitat. In a field survey, we confirmed that these fish frequently have to distinguish between disturbances stemming from attacking birds (multimodal) and those which pose no (immediate) threat such as bird overflights (unimodal). In a laboratory experiment, we then exposed fish to artificial visual and/or acoustic stimuli presented separately or combined. Sensitivity was high regardless of stimulus type and number (more than 96% of fish initiated diving), but fish dove deeper, faster, and for longer when both stimuli were available simultaneously. Based on the system’s high rates of bird activity, we argue that such an unselective dive initiation with subsequent fine-tuning of diving parameters in accordance to cue modality represents an optimal strategy for these fish to save energy necessary to respond to future attacks. Ultimately, our study shows that fish anticipate the imminent risk posed by disturbances linked to bird predation through integrating information from both visual and acoustic cues.

When ancestry haunts – can evolutionary links to ancestors affect vulnerability of Australian prey to introduced predators? A preliminary study

The high extinction risk of Australian marsupials has been attributed to their failure to recognise novel predators, the application of inappropriate antipredator responses, and advanced hunting strategies of novel predators. This study is a preliminary attempt to explore whether the Lumholtz’ tree-kangaroo (Dendrolagus lumholtzi) (a) is able to recognise odour cues from different predators as threats, and (b) possesses predator-archetype specific antipredator responses. A small number of available captive tree-kangaroos were exposed to faecal odours from two extant predators of different archetypes (python, dingo), a regionally extinct predator which closely matches past terrestrial predators (Tasmanian devil), and a novel predator (domestic dog). Lavender oil was used as non-predator novel odour and water as control. Results suggest that all subjects associated the presented odours with a threat, albeit to different degrees, but did not display predator-archetype specific responses. It appears that this species applies an ancestral antipredator response of flight-on-the ground when encountering predators, including novel predators. Although the results need to be confirmed with more animals, further studies on the vulnerability of Australian prey to novel predators should take the ancestral history of Australian prey species into account.

Ecophysiological Aspects and sxt Genes Expression Underlying Induced Chemical Defense in STX-Producing Raphidiopsis raciborskii (Cyanobacteria) against the Zooplankter Daphnia gessneri.

Cyanobacteria stand out among phytoplankton when they form massive blooms and produce toxins. Because cyanotoxin genes date to the origin of metazoans, the hypothesis that cyanotoxins function as a defense against herbivory is still debated. Although their primary cellular function might vary, these metabolites could have evolved as an anti-predator response. Here we evaluated the physiological and molecular responses of a saxitoxin-producing Raphidiopsis raciborskii to infochemicals released by the grazer Daphnia gessneri. Induced chemical defenses were evidenced in R. raciborskii as a significant increase in the transcription level of sxt genes, followed by an increase in saxitoxin content when exposed to predator cues. Moreover, cyanobacterial growth decreased, and no significant effects on photosynthesis or morphology were observed. Overall, the induced defense response was accompanied by a trade-off between toxin production and growth. These results shed light on the mechanisms underlying zooplankton–cyanobacteria interactions in aquatic food webs. The widespread occurrence of the cyanobacterium R. raciborskii in freshwater bodies has been attributed to its phenotypic plasticity. Assessing the potential of this species to thrive over interaction filters such as zooplankton grazing pressure can enhance our understanding of its adaptive success.

Pre-release training, predator interactions and evidence for persistence of anti-predator behavior in reintroduced `alalā, Hawaiian crow

Animal translocations commonly fail due to predation after release, especially if animals are reared in human care, away from natural predation pressure. Anti-predator training can be a useful tool for combating the predator naivety of released animals, but its effective implementation requires attention to numerous details. We present the step-by-step development of an anti-predator training regime, tailored to the critically endangered `alalā (Corvus hawaiiensis). `Alalā are the last remaining corvid species of the Hawaiian islands, and historically the level of predation they experienced from the Hawaiian hawk, `io (Buteo solitarius) is unclear. However, one factor thought to compromise outcomes of earlier attempts to reintroduce `alalā in the 1990′s was predation by `io, motivating the development of anti-predator training for recent `alalā releases. We documented evidence for what appears to be competent anti-predator behavior in conservation-bred `alalā, with a range of behavioral strategies for coping with `io observed during a series of controlled presentations of `io and `io-related stimuli. These behavioral responses included vigilance, alarm calling, and mobbing during experimental trials conducted in release training aviaries. Our results did not conclusively establish that anti-predator learning occurred as a result of the training, or that the training produced enhanced survival. However, following release, `io-`alalā encounters were observed on more than 35 occasions, and `alalā responses mirrored those observed pre-release, including the first ever recorded mobbing interactions by juvenile flocks of `alalā in the wild. While it is unclear the extent to which training encouraged these species-appropriate anti-predator responses, their occurrence suggests that the `alalā has retained and can express defensive behavioral strategies. By documenting the design process, training execution and behavioral outcomes in `alalā, we highlight details that other reintroduction programs may need to consider when preparing animals for reintroduction alongside natural predators.

Antipredator responses toward cat fur in wild brown rats tested in a semi-natural environment

Abstract Sensitivity to predator-related cues and performance of antipredator behaviors are universal among prey species. Rodents exhibit a diverse suite of antipredator behaviors that have been examined in both field and laboratory studies. However, the results from the laboratory have not always translated to the field. While laboratory studies consistently indicate strong fear-inducing effects of cat fur/skin odors, it is unclear whether this occurs in the field with wild rats. To address this issue, we tested the antipredator responses of wild brown rats (Rattus norvegicus) to predatory (domestic cat fur) and nonpredatory (common brushtail possum fur) odor cues in a semi-natural experimental paradigm. Rats were housed in open air enclosures containing two feeding stations. Following several nights of acclimatization, the feeding stations were paired with cat fur, possum fur, or no fur. Rats spent less time at a feeding station that was paired with cat fur. Duration of time spent at feeding stations increased across consecutive test days and across hours within individual test nights, although the rate of increase within nights was lower for cat fur paired stations. This overall increase might reflect habituation of antipredator behaviors, increasing hunger, or loss of cue potency over time. We suggest that wild brown rats recognize and respond to cat fur odor cues, but their behavioral response is highly adaptable and finely tuned to the trade-off between predation risk and starvation that occurs across short temporal scales.

Long-term Exposure to Predation Threat Can Modulate the Expression of Sexually Selected Traits in Male Guppy (Poecilia reticulata)

The expression of sexual traits can be affected by different environmental factors among which predation may be particularly important life-history traits. This study investigated the effects of predation risk on courtship behavior, growth and sexual color patterns in male guppy (Poecilia reticulata). In the study, juvenile male guppies were randomly assigned to two treatments, namely T1 (predation threat) and T2 (no predation threat). The courtship performances were visually observed and recorded, while color patterns and morphological traits were measured by using the ImageJ software from the captured photos. The courtship behavioral trials revealed that predation threatened males performed significantly lower number of sigmoid displays than those of no-predation group. Further, the predation scared group had significantly shorter standard length, body area and a reduced number and area of orange spots than their counter group. However, the predation threat did not affect significantly the gonopodial thrusts, and black and iridescent color components (spot number and area). The reduction of costly traits (e.g. behavior, color patterns and body size) is common anti-predator response which presumably reduces predation risks. Male guppies are probably using this form of defence in response to the increased predation risk. The overall result suggests that growth and sexually selected trait expression are sensitive to predation risk and thereby aid in our understanding to predict the evolution of phenotypic variation in natural systems.

Disentangling Natural and Anthropogenic Forms of Mortality and Serious Injury in a Poorly Studied Pelagic Dolphin

Killer whale (Orcinus orca) populations specialize in both prey and prey acquisition tactics around the world and may be a primary evolutionary driver of the habits of small cetaceans. Entanglement in fishing gear is the most significant anthropogenic threat to the survival of cetaceans worldwide. Distinguishing between natural and human-caused sources of mortality and injury is a key task in marine mammal conservation and management. In British Columbia (BC), Canada, mammal-eating killer whales co-occur with Pacific white-sided dolphins (Lagenorhynchus obliquidens). Bycatch mortality rates are unknown here due to lack of systematic fisheries observer coverage. Drawing from more than three decades of first-hand observations of killer whale attacks on Pacific white-sided dolphins, we identify common themes with respect to predatory behavior of killer whales and anti-predatory responses of dolphins. With input from veterinary pathologists, we outline clues to distinguish killer whale rake marks from scars and wounds likely to be caused by fishery interactions. We examined photographs of 415 well-marked Pacific white-side dolphins for evidence of injuries and scars consistent with either killer whale attacks or fishery interactions. In this case study, healed scars from interactions with killer whale predators were ∼8× more common than scars from fishery interactions (3.9 vs. 0.5%), suggesting that predation is a much bigger threat to Pacific white-sided dolphins in the study area than anthropogenic impacts, or that dolphins are much less likely to survive a fishery interaction than a predation attempt. To advance our knowledge on poorly studied species, multiple lines of evidence will be needed.

Severity Scoring of Behavioral Responses of Sperm Whales (Physeter macrocephalus) to Novel Continuous versus Conventional Pulsed Active Sonar

Controlled exposure experiments (CEEs) have demonstrated that naval pulsed active sonar (PAS) can induce costly behavioral responses in cetaceans similar to antipredator responses. New generation continuous active sonars (CAS) emit lower amplitude levels but more continuous signals. We conducted CEEs with PAS, CAS and no-sonar control on free-ranging sperm whales in Norway. Two panels blind to experimental conditions concurrently inspected acoustic-and-movement-tag data and visual observations of tagged whales and used an established severity scale (0–9) to assign scores to putative responses. Only half of the exposures elicited a response, indicating overall low responsiveness in sperm whales. Responding whales (10 of 12) showed more, and more severe responses to sonar compared to no-sonar. Moreover, the probability of response increased when whales were previously exposed to presence of predatory and/or competing killer or long-finned pilot whales. Various behavioral change types occurred over a broad range of severities (1–6) during CAS and PAS. When combining all behavioral types, the proportion of responses to CAS was significantly higher than no-sonar but not different from PAS. Responses potentially impacting vital rates i.e., with severity ≥4, were initiated at received cumulative sound exposure levels (dB re 1 μPa2 s) of 137–177 during CAS and 143–181 during PAS.

Evolution and Adaptation of Anti-predation Response of Prey in a Two-Patchy Environment.

When perceiving a risk from predators, a prey may respond by reducing its reproduction and decreasing or increasing (depending on the species) its mobility. We formulate a patch model to investigate the aforementioned fear effect which is indirect, in contrast to the predation as a direct effect, of the predator on the prey population. We consider not only cost but also benefit of anti-predation response of the prey, and explore their trade-offs together as well as the impact of the fear effect mediated dispersals of the prey. In the case of constant response level, if there is no dispersal and for some given response functions, the model indicates the existence of an evolutionary stable strategy which is also a convergence stable strategy for the response level; and if there is dispersal, the analysis of the model shows that it will enhance the co-persistence of the prey on both patches. Considering the trait as another variable, we continue to study the evolution of anti-predation strategy for the model with dispersal, which leads to a three-dimensional system of ordinary differential equations. We perform some numerical simulations, which demonstrate global convergence to a positive equilibrium with the response level evolving towards a positive constant level, implying the existence of an optimal anti-predation response level.