Antipredator Responses Research Articles

Olfactory sensitivity and threat-sensitive responses to alarm cue in an invasive fish

Establishing how invasive animals perceive and manage the threat of predation improves our understanding of why certain species rapidly invade unfamiliar environs where others do not. The threat-sensitive response hypothesis predicts that prey that modulate their antipredator behavior in response to the perceived intensity of the threat will have a selective advantage. We investigated the behavioral and olfactory antipredator responses of migrating invasive sea lamprey to varying concentrations of a conspecific alarm cue in the laboratory and field. We hypothesized the odor indicates the presence and location of predation risk on river shorelines, allowing sea lamprey to either alter their migratory path and/or movement timing to reduce exposure to the predator. Migratory sea lampreys exhibited threat-sensitivity in space-use both in the laboratory and nature by increasingly avoiding areas as the concentration of alarm cue increased when safe space was available (areas without the alarm cue). The intensity of the avoidance at higher concentrations was greater in the field. Electrophysiological recordings from the olfactory organ confirmed a concomitant increase in signal output to the olfactory bulb of the brain when exposed to increased alarm cue concentrations. However, increasing the alarm cue concentration did not alter the animal’s motivation to move upstream in either avoidable risk or unavoidable risk scenarios. Exposure to alarm cue affected decisions regarding how to move, but not whether to move. Knowledge of how the animal uses the cue in nature reveals ways to use the cue to manipulate movement and achieve conservation goals.

Anti-predator responses to chemical, visual, and combined cues by an estuarine hermit crab from sandy tidal flats

Anti-predator responses to chemical, visual, and combined cues by an estuarine hermit crab from sandy tidal flats

Smelling danger: Lady beetle odors affect aphid population abundance and feeding, but not movement between plants

Predator-prey interactions are complex ecological interactions that influence community structure and function. Predators affect prey density directly by consuming and removing prey from the environment. Additionally, predators can elicit non-consumptive effects in prey which influence prey survival by altering prey behavior and physiology. Key to understanding the consequences of non-consumptive effects for prey populations is determining mechanisms of detection of predation risk. With insects, olfactory cues are commonly used to convey information. It has been shown that insect prey may eavesdrop on predator odor cues as a means to determine predation risk. Herein, we assessed whether the volatile odors of the predatory insect Harmonia axyridis (multicolored Asian lady beetle) affect the behavior (feeding and dispersal behaviors) and performance (population abundance) of aphids (Myzus persicae). Aphids reduced phloem ingestion when feeding in close proximity to lady beetle cues, however, predator odors did not drive aphids to relocate on plants or to disperse across plants away from the odor source. The lack of dispersal we observed is in contrast to studies that show increased aphid movement in the presence of predators that can make physical contact with the aphids. We also found that lady beetle odor cues had a population-level effect on aphids, with 25% reduction in aphid population abundance in the presence of the odor cues. This study highlights that anti-predator strategies may differ depending on the mechanism of detection of predation risk (i.e. olfactory versus physical stimuli), or the context in which they experience them, and raises the question of whether anti-predator responses observed are maladaptive or beneficial to aphid survival. These conclusions are critical for both our basic understanding of the mechanisms driving predator-prey interactions as well as providing insight into pest-natural enemy interactions within agricultural landscapes.

Age differences in the responses of vervet monkeys, Chlorocebus pygerythrus, to terrestrial alarm calls

The high costs of predation and the opportunity costs associated with predator avoidance are likely to select for flexibility in the development of antipredator responses based on local socioecological conditions. As group size is hypothesized to vary across populations as a function of predation risk, the development of antipredator behaviours throughout ontogeny may be influenced by the size of an individual's social group. Here, we explore the development of alarm call responses in wild vervet monkeys living at the Samara Game Reserve, South Africa. Vervet monkeys at this site live in relatively large social groups, and adult responses to alarms have previously shown less uniformity than in other sites where group sizes are smaller. We presented monkeys playback recordings of terrestrial alarm calls produced by individuals of different age–sex classes. We then videorecorded and scored the responses of receivers along an ordinal maturity scale. We used a mixed-effects ordinal logistic regression model within a Bayesian framework to explore how response intensity is affected by the age–sex of the caller, and the age of receivers. Our analysis showed that younger monkeys (<2 years old) exhibit strong evasive responses to call stimuli, regardless of the age–sex class of the caller. The intensity of these responses decreases with age, with responses to nonalarm calls decreasing earlier in development than responses to alarm calls. Adult responses to alarm calls in this population are as likely to be characterized by a general increase in vigilance as they are to consist of an evasive response. We suggest that responses in younger individuals at Samara are mediated by generalized startle responses to loud, plosive noises. Development of more specific responses throughout ontogeny is likely to be dependent on learning from adult models, whose milder responses reflect local socioecological conditions specific to Samara.

Size-dependent response of the mussel collective behaviour to plastic leachates and predator cues

Both individual and collective anti-predator behaviours are essential for the survival of many species. This is particularly true for ecosystem engineers such as intertidal mussels, which through their collective behaviour create novel habitats for a range of organisms and biodiversity hotspots. However, contaminants may disrupt these behaviours and consequently indirectly affect exposure to predation risk at the population level. Among these, plastic litter is a major and ubiquitous contaminant of the marine environment. Here, we assessed the impact of microplastic (MP) leachates of the most produced plastic polymer, polypropylene (PlasticsEurope, 2022), at a high but locally relevant concentration (i.e. ca. 12 g L−1) on the collective behaviours and anti-predator responses of both small and large Mytilus edulis mussels. Indeed, in contrast to large mussels, small ones reacted to MP leachates, showing a taxis towards conspecifics and stronger aggregations. All mussels reacted to the chemical cues of the predatory crab, Hemigrapsus sanguineus, but with two different collective anti-predator behaviours. Small mussels only showed a taxis towards conspecifics when exposed to predator cues. This response was also found in large ones with a tendency to form more strongly bound aggregations and a considerable reduced activity, i.e. they significantly delayed their time to start to form aggregations and decreased their gross distance. These anti-predator behaviours were respectively inhibited and impaired in small and large mussels by MP leachates. The observed collective behavioural changes may reduce individual fitness by enhancing predation risk, particularly in small mussels that are the crab H. sanguineus's favourite preys. Given the key role of mussels as ecosystem engineers, our observations suggest that plastic pollution may have implication on M. edulis at the species level, but also enhancing a cascading effect towards a higher level of organisation such as population, community and ultimately structure and function of intertidal ecosystem.

Adaptive echolocation behavior of bats and toothed whales in dynamic soundscapes.

Journal of Experimental Biology has a long history of reporting research discoveries on animal echolocation, the subject of this Centenary Review. Echolocating animals emit intense sound pulses and process echoes to localize objects in dynamic soundscapes. More than 1100 species of bats and 70 species of toothed whales rely on echolocation to operate in aerial and aquatic environments, respectively. The need to mitigate acoustic clutter and ambient noise is common to both aerial and aquatic echolocating animals, resulting in convergence of many echolocation features, such as directional sound emission and hearing, and decreased pulse intervals and sound intensity during target approach. The physics of sound transmission in air and underwater constrains the production, detection and localization of sonar signals, resulting in differences in response times to initiate prey interception by aerial and aquatic echolocating animals. Anti-predator behavioral responses of prey pursued by echolocating animals affect behavioral foraging strategies in air and underwater. For example, many insect prey can detect and react to bat echolocation sounds, whereas most fish and squid are unresponsive to toothed whale signals, but can instead sense water movements generated by an approaching predator. These differences have implications for how bats and toothed whales hunt using echolocation. Here, we consider the behaviors used by echolocating mammals to (1) track and intercept moving prey equipped with predator detectors, (2) interrogate dynamic sonar scenes and (3) exploit visual and passive acoustic stimuli. Similarities and differences in animal sonar behaviors underwater and in air point to open research questions that are ripe for exploration.

Defensive tail displays in dice snakes (Natrix tessellata) are influenced by colour morph and sex

Abstract Defensive tail displays are common in squamates; however, the factors underlying the occurrence of such displays are rarely studied. We tested 268 adult dice snakes (Natrix tessellata) in a population with three colour morphs, exploring the effects of morph, sex, body size, tail length and temperature on the occurrence of tail waving in two stages of the antipredator response. When captured, the blotched morph tail waved the most, followed by melanistic then green morphs. Males displayed tail waving during post-capture immobility more often than females, possibly due to sexual dimorphism in tail length and/or fleeing speed. The occurrence of tail waving was low in response to a researcher as a “predator” (∼2.6% on capture; ∼6.8% during immobility), however the occurrence of this behaviour during encounters with real predators might be different. Large sample size is crucial for revealing fine scale effects on rare behaviours such as tail waving.

Behaviour of mesopredatory coral reef fishes in response to threats from sharks and humans

Both sharks and humans present a potentially lethal threat to mesopredatory fishes in coral reef systems, with implications for both population dynamics and the role of mesopredatory fishes in reef ecosystems. This study quantifies the antipredator behaviours mesopredatory fishes exhibit towards the presence of large coral reef carnivores and compares these behavioural responses to those elicited by the presence of snorkelers. Here, we used snorkelers and animated life-size models of the blacktip reef shark (Carcharhinus melanopterus) to simulate potential predatory threats to mesopredatory reef fishes (lethrinids, lutjanids, haemulids and serranids). The responses of these reef fishes to the models and the snorkelers were compared to those generated by three non-threatening controls (life-size models of a green turtle [Chelonia mydas], a PVC-pipe [an object control] and a Perspex shape [a second object control]). A Remote Underwater Stereo-Video System (Stereo-RUV) recorded the approach of the different treatments and controls and allowed accurate measurement of Flight Initiation Distance (FID) and categorization of the type of flight response by fishes. We found that mesopredatory reef fishes had greater FIDs in response to the approach of threatening models (1402 ± 402–1533 ± 171 mm; mean ± SE) compared to the controls (706 ± 151–896 ± 8963 mm). There was no significant difference in FID of mesopredatory fishes between the shark model and the snorkeler, suggesting that these treatments provoked similar levels of predator avoidance behaviour. This has implications for researchers monitoring behaviour in situ or using underwater census as a technique to estimate the abundance of reef fishes. Our study suggests that, irrespective of the degree to which sharks actually consume these mesopredatory reef fishes, they still elicit a predictable and consistent antipredator response that has the potential to create risk effects.

Anti-predator strategies of adult male Central Himalayan Langurs (Semnopithecus schistaceus) in response to domestic dogs in a human-dominated landscape.

The evolution of predator-prey relationships is an important topic in primatology. Many aspects of primate society have been explained as a response to predation pressure. While predation has been discussed in broad theoretical terms, few systematically collected data exist on the subject. Furthermore, little information exists regarding the inter-male variation in responses to predators. To address this data gap, predatory dog-primate interactions were studied in a 78-member group of habituated, individually recognized Central Himalayan Langurs (CHL) (Semnopithecus schistaceus) living in a high-altitude subsistence agricultural landscape of northern India. We recorded 312 langur-dog interactions over 2 years. These predation events resulted in 15 serious attacks on adult females, infants, juveniles and sub-adults, in eight of which the prey was killed and consumed on the spot. In response to dog predation, adult males performed three types of anti-predator response behaviors: direct fighting with a predator, emitting alarm calls, fleeing and/or freezing. Differences were noted in each male's response to village dogs. The results showed that the likelihood of CHL adult males engaging in more costly counterattacks or attention getting alarm calls were better predicted by the level of investment in the group (genetic relatedness, duration of residency, social relationships), but not rank and mating rate. Long-duration resident adult males performed high and/or intermediate cost behaviors to protect vulnerable members of the group; their potential offspring, maternal siblings or cousins, and adult female social partners. Short-term residents or recent immigrant males exhibited two less energetically costly, more self-preserving behaviors, depending on their rank: (1) high-ranking short-tenure duration males, with high mating frequencies, performed flee and freeze responses; (2) low-ranking, low-mating-frequency males performed more alarm calls. Counterattacks and alarm calls were performed by adult males with relatively more experience with village dogs and were directed towards dogs with predatory histories significantly more often than dogs with non-predatory histories. Natural selection and kin selection have both contributed to the evolution of CHL anti-predator tactics.

Anti-predatory responses of two native gastropods to an invasive predator

As native prey species lack a co-evolutionary history with invasive predators, they often have limited capacity to identify and respond to the threat of predation. This phenomenon has led to the prey naivety hypothesis. Invasive predators can have devastating impacts on native prey populations. Thus, there is a need to better understand the ability of native species to respond to invasive predators. Here, laboratory behavioural experiments tested the anti-predatory responses of two gastropods, Cominella lineolata and Dicathais orbita, against an invasive predator, Asterias amurensis, and a native predator, Coscinasterias muricata. There was no chemical exudate released by any D. orbita to either species of seastar or a wax seastar control. Only a small percentage of D. orbita individuals fled from the invasive (12.5%) and native (17%) seastars. The overall separation distance between D. orbita and the two seastars was recorded for a further three minutes. While the measured distance between the invasive seastar and D. orbita was greater (61 ± 7 mm) than the distance between the native seastar (39 ± 10 mm), this was not statistically significant. In contrast, C. lineolata exhibited strong responses to the native seastar, but less strongly to the invasive seastar. Only 50% of snails fled and 15% released an exudate in response to A. amurensis, compared to 80% of snails fleeing and releasing an exudate to the native seastar. In contrast to expectations, C. lineolata collected from naïve populations tended to respond less frequently (25% of snails) by fleeing from the invasive seastar than snails collected from exposed (50% of snails), however this was not significant. This same trend was observed for C. muricata with 70% and 90% of snails fleeing from exposed and naïve populations, respectively. After recording the behavioural responses of both seastars and snails for three minutes, the release of exudate resulted in an increased separation distance between C. lineolata and C. muricata of 122 mm (± 31 mm) compared with just 28 mm (± 16 mm) when the snails failed to release the exudate, whereas the separation distances between C. lineolata and A. amurensis was 143 mm (± 31 mm) and 87 mm (± 10 mm) with and without exudate release, respectively. This latter result indicates that while the exudate appears to be an effective anti-predatory mechanism against the native seastar, it appears less effective against the invasive seastar.

Landscape of fear or landscape of food? Moose hunting triggers an antipredator response in brown bears.

Hunters can affect the behavior of wildlife by inducing a landscape of fear, selecting individuals with specific traits, or altering resource availability across the landscape. Most research investigating the influence of hunting on wildlife resource selection has focused on target species and less attention has been devoted to nontarget species, such as scavengers that can be both attracted or repelled by hunting activities. We used resource selection functions to identify areas where hunters were most likely to kill moose (Alces alces) in south-central Sweden during the fall. Then, we used step-selection functions to determine whether female brown bears (Ursus arctos) selected or avoided these areas and specific resources during the moose hunting season. We found that, during both day and nighttime, female brown bears avoided areas where hunters were more likely to kill moose. We found evidence that resource selection by brown bears varied substantially during the fall and that some behavioral changes were consistent with disturbance associated with moose hunters. Brown bears were more likely to select concealed locations in young (i.e., regenerating) and coniferous forests and areas further away from roads during the moose hunting season. Our results suggest that brown bears react to both spatial and temporal variations in apparent risk during the fall: moose hunters create a landscape of fear and trigger an antipredator response in a large carnivore even if bears are not specifically targeted during the moose hunting season. Such antipredator responses might lead to indirect habitat loss and lower foraging efficiency and the resulting consequences should be considered when planning hunting seasons.

Perverse outcomes from fencing fauna: Loss of antipredator traits in a havened mammal population

Populations of threatened animals are increasingly preserved within predator-free havens, where populations tend to grow rapidly, resource competition increases, and traits relevant to avoiding predation may be selected against. We examine this phenomenon using a ten-year longitudinal dataset on a threatened Australian mammal; the woylie (Bettongia penicillata ogilbyi). Behavioural and morphological data were collected during routine monitoring of a havened woylie population and an adjacent wild population where predators occur, from which six traits relating to predator escape were extracted. Paired comparisons revealed that havened woylies were less likely to show injuries from excessive agitation in traps, were less likely to eject pouch young, and had shorter approach distances compared to woylies outside the haven, suggesting a dampened antipredator response. Further, body mass and relative leg length declined over time in the havened population, compared with no change outside of the haven, suggesting selection in the haven against body size. Population density affected body size and agitation in traps differently in havened and non-havened populations, indicating an interaction between resource competition and relaxed selection that likely hastens the loss of anti-predator traits. Our study offers a mechanistic understanding of the loss of anti-predator responses in havens, which is essential for guiding how populations could be managed to better realise their potential for recovering threatened fauna.

Multilevel social structure predicts individual helping responses in a songbird

Multilevel societies are formed when stable groups of individuals spatially overlap and associate preferentially with other groups, producing a hierarchical social structure.1 Once thought to be exclusive to humans and large mammals, these complex societies have recently been described in birds.2,3 However, it remains largely unclear what benefits individuals gain by forming multilevel societies.1 One hypothesis-based on food sharing in hunter-gatherers4-is that multilevel societies facilitate access to a range of cooperative relationships, with individual investment varying across the hierarchical levels of the society. We tested experimentally whether such graded cooperation occurs in the multilevel society of a songbird, the superb fairy-wren (Malurus cyaneus). Specifically, we measured whether responses to playbacks of distress calls-used to recruit help when in extreme danger-varied according to the social level at which the focal individual is connected with the caller. We predicted that anti-predator responses should be highest within breeding groups (the core social unit), intermediate between groups from the same community, and lowest across groups from different communities. Our results confirm that birds exhibit the predicted hierarchical pattern of helping and that, within breeding groups, this pattern is independent of kinship. This pattern of graded helping responses supports the hypothesis that multilevel social structures can sustain stratified cooperative relationships and reveals similarity in cooperation in qualitatively different behaviors-anti-predator behavior and food sharing-in the multilevel societies of songbirds and humans.

Prey species increase activity in refugia free of terrestrial predators

The decline of terrestrial predator populations across the globe is altering top-down pressures that drive predator-prey interactions. However, a knowledge gap remains in understanding how removing terrestrial predators affects prey behavior. Using a bifactorial playback experiment, we exposed fox squirrels to predator (red-tailed hawks, coyotes, dogs) and non-predator control (Carolina wren) calls inside terrestrial predator exclosures, accessible to avian predators, and in control areas subject to ambient predation risk. Fox squirrels increased their use of terrestrial predator exclosures, a pattern that corresponded with 3years of camera trapping. Our findings suggest fox squirrels recognized that exclosures had predictably lower predation risk. However, exclosures had no effect on their immediate behavioral response towards any call, and fox squirrels responded most severely to hawk predator calls. This study shows that anthropogenically driven predator loss creates predictably safer areas (refugia) that prey respond to proactively with increased use. However, the persistence of a lethal avian predator is sufficient to retain a reactive antipredator response towards an immediate predation threat. Some prey may benefit from shifting predator-prey interactions by gaining refugia without sacrificing a sufficient response towards potential predators.

Functionally redundant multimodal predator cues elicit changes in prey foraging behavior

Abstract Many prey species can assess the risk of predation from information acquired through different sensory systems. For many animals, this information is detected with sensory organs specialized for visual (sight) or chemical (smell or taste) stimuli. It is unclear; however, whether information acquired through multiple sensory systems is functionally redundant or interchangeable, especially if the message is the same. Here, we assess prey response to unimodal visual and chemical cues as well as multimodal (visual + chemical) cues. We specifically test if a foraging individual shows a stronger behavioral response to risk when they can perceive that risk through multimodal versus unimodal cues. To do this, we measured the functional response (prey abundance–foraging rate relationship) of Tibellus oblongus spiders foraging on midges while exposing them to visual stimuli, chemical stimuli, or a combination of both visual and chemical stimuli from potential predators. We then determined if the spider’s functional response for the multimodal treatment differed more strongly from a control treatment than from either unimodal treatment. We found that under any simulated predation risk (multimodal and both unimodal), T. oblongus spiders showed longer handling times than in control groups without risk. However, we saw no elevated anti-predator response in the multimodal treatment, suggesting that information from visual and chemical modalities is interchangeable and sufficient to indicate reliably predation risk.

Chemical communication in wood frog (Rana sylvatica) tadpoles is influenced by early-life exposure to naphthenic acid fraction compounds

Environmental pollutants can disrupt chemical communication between aquatic organisms by interfering with the production, transmission, and/or detection of, as well as responses to, chemical cues. Here, we test the hypothesis that early-life exposure to naphthenic acid fraction compounds (NAFCs) from oil sands tailings disrupts antipredator-associated chemical communication in larval amphibians. Wild adult wood frogs (Rana sylvatica) captured during their natural breeding period were combined (1 female:2 males) in six replicate mesocosms filled with either uncontaminated lakewater or with NAFCs isolated from an active tailings pond in Alberta, Canada, at nominal 5 mg/L concentrations. Egg clutches were incubated and tadpoles maintained in their respective mesocosms for ∼40 days post-hatch. Tadpoles (Gosner stage 25-31) were then transferred individually to trial arenas filled with uncontaminated water and exposed to one of six chemical alarm cue (AC) stimuli solutions following a 3 × 2 × 2 design (3 AC types × 2 stimulus carriers × 2 rearing exposure groups). Relative to control tadpoles, NAFC-exposed tadpoles demonstrated higher baseline activity levels (line crosses and direction changes) when introduced to uncontaminated water. Antipredator responses differed in graded fashion with AC type, with control ACs eliciting the greatest latency to resume activity, water the least, and NAFC-exposed ACs intermediate. Pre- to post-stimulus difference scores were non-significant in control tadpoles, while NAFC-exposed tadpoles demonstrated significantly greater variation. While this suggests that exposure to NAFCs from fertilization through hatching may have interfered with AC production, it is unclear whether the quality or quantity of cues was affected. There was also no clear evidence that NAFC carrier water interfered with ACs or the alarm response in unexposed control tadpoles. These results emphasize the importance of understanding how behavioral and physiological effects of early-life NAFC exposure on critical antipredator responses may persist across life history stages.

Threespine stickleback do not use social cues to learn about predation risk

Animals can use a variety of sources of information to learn about local predation threats, including the behavior of conspecifics. Socially-acquired information about predation risk has been demonstrated in a variety of fish species, so the phenomenon is likely taxonomically widespread. Threespine stickleback fish are a geographically widespread species that encounters a variety of native and introduced predators throughout its range; as such, learning to recognize predators may be an important component of survival. We assessed whether laboratory-bred, predator-naïve stickleback could learn to fear the odor of a live rainbow trout by first observing videos of conspecifics exhibiting antipredator responses in the presence of trout odor. We found that this is not the case: following one conditioning trial in which trout odor was paired with videos of frightened conspecifics, stickleback did not exhibit an increase in antipredator behavior (e.g., a decrease in activity or an increase in hiding) in the presence of trout odor. Although there is evidence that stickleback use social information to find foraging patches, it does not appear that they do the same to learn about predation threat, at least in the context of our experimental conditions.

Western Australian magpies respond to urgency information contained in conspecific alarm calls

Abstract Many animals provide information about predator proximity in their alarm calls. In response to predators further away, Western Australian magpies (Gymnorhina tibicen dorsalis) produce alarm calls containing fewer notes compared to those produced when predators are closer. Since the ability to make fine-scale adjustments to antipredator responses by being sensitive to the level of urgency in calls may be beneficial, receivers are expected to be able to appropriately decipher and respond to this information. We conducted playbacks to test whether magpies can respond to urgency information in conspecific alarm calls. Magpies were exposed to low-urgency calls (calls with one note), high-urgency calls (calls with four notes), and one- and four-note control calls. Receivers showed greater levels of responsiveness following playbacks of high-urgency calls compared to playbacks of low-urgency and control calls, providing evidence that magpies can respond to information about the urgency of a predator threat from conspecific alarm calls.

The habitat connectivity hypothesis of escape in urban woodland birds.

Habitat destruction and fragmentation increasingly bring humans into close proximity with wildlife, particularly in urban contexts. Animals respond to humans using nuanced anti-predator responses, especially escape, with responses influenced by behavioral and life history traits, the nature of the risk, and aspects of the surrounding environment. Although many studies examine associations between broad-scale habitat characteristics (i.e., habitat type) and escape response, few investigate the influence of fine-scale aspects of the local habitat within which escape occurs. We test the "habitat connectivity hypothesis," suggesting that given the higher cost of escape within less connected habitats (due to the lack of protective cover), woodland birds should delay escape (tolerate more risk) than when in more connected habitat. We analyze flight-initiation distances (FIDs) of five species of woodland birds in urban Melbourne, south-eastern Australia. A negative effect of habitat connectivity (the proportion of the escape route with shrubs/trees/perchable infrastructure) on distance fled was evident for all study species, suggesting a higher cost of escape associated with lower connectivity. FID did not vary with connectivity at the location at which escape was initiated (four species), apart from a positive effect of habitat connectivity on FID for Noisy Miner Manorina melanocephala. We provide some support for two predictions of the "habitat connectivity hypothesis" in at least some taxa, and conclude it warrants further investigation across a broader range of taxa inhabiting contrasting landscapes. Increasing habitat connectivity within urban landscapes may reduce escape stress experienced by urban birds.

Susceptibility to Predation Varies with Body Mass, Foraging Niche, and Anti-Predator Responses among Bird Species

Predation is a major source of mortality for many avian species. Species that face more predators, and those with less effective anti-predator responses, are presumably more likely to die from predation over time. Predation rate, as a measure of susceptibility to predation, is difficult to measure in the field. Radio-tracking studies, however, allow researchers to determine the time and cause of death of marked individuals, making it possible to estimate predation rate. I used estimates of predation rates from a large number of published radio-tracking studies in birds to assess in a phylogenetic framework the effect of several potential determinants. I obtained 393 estimates of predation rates from 129 species. Predation rates were lower in areas with fewer predators, such as islands and aquatic habitats, and for species with fewer potential predators, such as larger species. The predation rate was also lower for prey species with effective anti-predator responses, such as those that forage in flocks. Radio-tracking studies provide a unique opportunity to estimate overall predation rate in the field. Broadening the range of species and the range of habitats involved in such studies will help to further elucidate the factors that affect susceptibility to predation.