ABSTRACT: After detecting an approaching predator, animals make a decision when to flee. Prey will initiate flight soon after detecting a predator so as to minimize attentional costs related to on-going monitoring of the whereabouts of the predator. Such costs may compete with foraging and other maintenance activities and hence be larger than the costs of immediate flight. The drivers of interspecific variation in escape strategy are poorly known. Here we investigated the morphological, life history and natural history traits that correlate with variation in avian escape strategy across a sample of 96 species of birds. Brain mass, body size, habitat structure and group size were the main predictors of escape strategy. The direction of the effect of these traits was consistent with selection for a reduction of monitoring costs. Therefore, attentional costs depend on relative brain size, which determines the ability to monitor the whereabouts of potential predators and the difficulty of this task as reflected by habitat and social complexity. Thus brain size, and the cognitive functions associated with it, constitute a general framework for explaining the effects of body size, habitat structure and sociality identified as determinants of avian escape strategy.
Project description:All animals flee from potential predators, and the distance at which this happens is optimized so the benefits from staying are balanced against the costs of flight. Because predator diversity and abundance decreases with increasing latitude, and differs between rural and urban areas, we should expect escape distance when a predator approached the individual to decrease with latitude and depend on urbanization. We measured the distance at which individual birds fled (flight initiation distance, FID, which represents a reliable and previously validated surrogate measure of response to predation risk) following a standardized protocol in nine pairs of rural and urban sites along a ca. 3000 km gradient from Southern Spain to Northern Finland during the breeding seasons 2009-2010. Raptor abundance was estimated by means of standard point counts at the same sites where FID information was recorded. Data on body mass and phylogenetic relationships among bird species sampled were extracted from the literature. An analysis of 12,495 flight distances of 714 populations of 159 species showed that mean FID decreased with increasing latitude after accounting for body size and phylogenetic effects. This decrease was paralleled by a similar cline in an index of the abundance of raptors. Urban populations had consistently shorter FIDs, supporting previous findings. The difference between rural and urban habitats decreased with increasing latitude, also paralleling raptor abundance trends. Overall, the latitudinal gradient in bird fear was explained by raptor abundance gradients, with additional small effects of latitude and intermediate effects of habitat. This study provides the first empirical documentation of a latitudinal trend in anti-predator behavior, which correlated positively with a similar trend in the abundance of predators.
Project description:Habitat reduction could drive biodiversity loss if the capacity of food webs to support predators is undermined by habitat-size constraints on predator body size. Assuming that (i) available space restricts predator body size, (ii) mass-specific energy needs of predators scale with their body size, and (iii) energy availability scales with prey biomass, we predicted that predator biomass per unit area would scale with habitat size (quarter-power exponent) and prey biomass (three-quarter-power exponent). We found that total predator biomass scaled with habitat size and prey resources as expected across 29 New Zealand rivers, such that a unit of habitat in a small ecosystem supported less predator biomass than an equivalent unit in a large ecosystem. The lower energetic costs of large body size likely mean that a unit of prey resource supports more biomass of large-bodied predators compared to small-bodied predators. Thus, contracting habitat size reduces the predator mass that can be supported because of constraints on predator body size, and this may be a powerful mechanism exacerbating reductions in biodiversity due to habitat loss.
Project description:Flight initiation distance (FID), the distance at which individuals take flight when approached by a potential (human) predator, is a tool for understanding predator-prey interactions. Among the factors affecting FID, tests of effects of group size (i.e., number of potential prey) on FID have yielded contrasting results. Group size or flock size could either affect FID negatively (i.e., the dilution effect caused by the presence of many individuals) or positively (i.e., increased vigilance due to more eyes scanning for predators). These effects may be associated with gregarious species, because such species should be better adapted to exploiting information from other individuals in the group than nongregarious species. Sociality may explain why earlier findings on group size versus FID have yielded different conclusions. Here, we analyzed how flock size affected bird FID in eight European countries. A phylogenetic generalized least square regression model was used to investigate changes in escape behavior of bird species in relation to number of individuals in the flock, starting distance, diet, latitude, and type of habitat. Flock size of different bird species influenced how species responded to perceived threats. We found that gregarious birds reacted to a potential predator earlier (longer FID) when aggregated in large flocks. These results support a higher vigilance arising from many eyes scanning in birds, suggesting that sociality may be a key factor in the evolution of antipredator behavior both in urban and rural areas. Finally, future studies comparing FID must pay explicit attention to the number of individuals in flocks of gregarious species.
Project description:The kinematic and behavioral components of the escape response can affect the outcomes of predator-prey interactions. For example, because sensory perception range can have spatial bias, and because turn duration before the initiation of escape locomotion can be smaller when prey is oriented away from predators, the prey's body orientation relative to a predator at the onset of the escape response (initial orientation) could affect whether prey successfully evade predators. We tested this hypothesis by recording the escape responses of juvenile red sea bream (Pagrus major) to the predatory scorpion fish (Sebastiscus marmoratus). Flight initiation distance tended to be small when prey were attacked from behind, suggesting that prey have spatial bias in detecting attacking predators. An increase in flight initiation distance increased escape probability. An increase in initial orientation decreased turn duration and increased escape probability when the effect of flight initiation distance was offset. These results suggest that initial orientation affects escape probability through two different pathways: changes in flight initiation distance and turn duration. These findings highlight the importance of incorporating initial orientation into other studies of the kinematics of predator-prey interactions.
Project description:Optimal escape theory states that animals should counterbalance the costs and benefits of flight when escaping from a potential predator. However, in apparent contradiction with this well-established optimality model, birds and mammals generally initiate escape soon after beginning to monitor an approaching threat, a phenomena codified as the "Flush Early and Avoid the Rush" (FEAR) hypothesis. Typically, the FEAR hypothesis is tested using correlational statistics and is supported when there is a strong relationship between the distance at which an individual first responds behaviorally to an approaching predator (alert distance, AD), and its flight initiation distance (the distance at which it flees the approaching predator, FID). However, such correlational statistics are both inadequate to analyze relationships constrained by an envelope (such as that in the AD-FID relationship) and are sensitive to outliers with high leverage, which can lead one to erroneous conclusions. To overcome these statistical concerns we develop the phi index (?), a distribution-free metric to evaluate the goodness of fit of a 1:1 relationship in a constraint envelope (the prediction of the FEAR hypothesis). Using both simulation and empirical data, we conclude that ? is superior to traditional correlational analyses because it explicitly tests the FEAR prediction, is robust to outliers, and it controls for the disproportionate influence of observations from large predictor values (caused by the constrained envelope in AD-FID relationship). Importantly, by analyzing the empirical data we corroborate the strong effect that alertness has on flight as stated by the FEAR hypothesis.
Project description:Escape enables prey to avoid an approaching predator. The escape decision-making process has traditionally been interpreted using theoretical models that consider ultimate explanations based on the cost/benefit paradigm. Ultimate approaches, however, suffer from inseparable extra-assumptions due to an inability to accurately parameterize the model's variables and their interactive relationships. In this study, we propose a mathematical model that uses intensity of predator-mediated visual stimuli as a basic cue for the escape response. We consider looming stimuli (i.e. expanding retinal image of the moving predator) as a cue to flight initiation distance (FID; distance at which escape begins) of incubating Mallards (Anas platyrhynchos). We then examine the relationship between FID, vegetation cover and directness of predator trajectory, and fit the resultant model to experimental data. As predicted by the model, vegetation concealment and directness of predator trajectory interact, with FID decreasing with increased concealment during a direct approach toward prey, but not during a tangential approach. Thus, we show that a simple proximate expectation, which involves only visual processing of a moving predator, may explain interactive effects of environmental and predator-induced variables on an escape response. We assume that our proximate approach, which offers a plausible and parsimonious explanation for variation in FID, may serve as an evolutionary background for traditional, ultimate explanations and should be incorporated into interpretation of escape behavior.
Project description:A perennial question in ornithology is whether flight has evolved mostly to facilitate access to food or as an anti-predator strategy. However, flight is an expensive mode of locomotion and species using flight regularly are associated with an expensive lifestyle. Using heart rate (HR) data loggers implanted in 13 female common eiders (Somateria mollissima), our objective was to test the hypothesis that a high level of flight activity increases their energy budget. We used the long-term recording (seven months) of HR as an index of energy expenditure and the HR flight signature to compile all flight events. Our results indicate that the eider is one of the thriftiest volant birds with only 10 minutes of flight time per day. Consequently, we were not able to detect any effect of flight activity on their energy budget despite very high flight costs (123-149 W), suggesting that flight was controlled by energy budget limitations. However, the low flight activity of that species may also be related to their prey landscape requiring few or no large-scale movements. Nevertheless, we suggest that the (fitness) benefits of keeping flight ability in this species exceed the costs by allowing a higher survival in relation to predation and environmental harshness.
Project description:A temporal separation of energetically costly life history events like reproduction and maintenance of the integumentary system is thought to be promoted by selection to avoid trade-offs and maximize fitness. It has therefore remained somewhat of a paradox that certain vertebrate species can undergo both events simultaneously. Identifying potential costs of overlapping two demanding life history stages will further our understanding of the selection pressures that shape the temporal regulation of life history events in vertebrates. We studied free-living tropical Slaty brush-finches (Atlapetes schistaceus), in which individuals spontaneously overlap reproduction and moult or undergo both events in separation. To assess possible costs of such an overlap we quantified feather quality and flight performance of individuals in different states. We determined individual's life history state by measuring gonad size and scoring moult stage, and collected a newly grown 7(th) primary wing feather for later analysis of feather quality. Finally, we quantified flight performance for each individual in the wild. Overlapping individuals produced lighter and shorter wing feathers than individuals just moulting, with females decreasing feather quality more strongly during the overlap than males. Moreover, overlapping individuals had a reduced flight speed during escape flights, while their foraging flight speed was unaffected. Despite overlappers being larger and having a smaller wing area, their lower body mass resulted in a similar wing load as in breeders or moulters. Individuals measured repeatedly in different states also showed significant decreases in feather quality and escape flight speed during the overlap. Reduced escape flight speed may represent a major consequence of the overlap by increasing predation risk. Our data document costs to undergoing two life history stages simultaneously, which likely arise from energetic trade-offs. Impairments in individual quality and performance may represent important factors that select for temporal separation of life history stages in other species.
Project description:Since 1986, studies about the escape decisions made by prey are grounded in optimal escape theory (OET) which states that prey will initiate escape when the risk of remaining and the costs of leaving are equal. However, a recent hypothesis, Flush Early and Avoid the Rush (FEAR), acknowledged that the cost of monitoring approaching predators might be a ubiquitous cost. The FEAR hypothesis predicts that prey will generally flee soon after they detect a predator so as to minimize the costs incurred by monitoring the predator. Knowing whether animals flee to reduce monitoring costs is of applied interest because wildlife managers use escape behavior to create set-back zones to reduce human-wildlife conflict. Here we provide the most comprehensive assessment of the FEAR hypothesis using data collected from 178 bird species representing 67 families from two continents. The FEAR hypothesis explains escape behavior in 79% of studied species. Because the FEAR hypothesis is a widespread phenomenon that drives escape behavior in birds, alert distance must be systematically incorporated into the design of set-back zones to protect vulnerable species.
Project description:BACKGROUND:Many animals rely on their escape performance during predator encounters. Because of its dependence on body size and temperature, escape velocity is fully characterized by three measures, absolute value, size-corrected value, and its response to temperature (thermal sensitivity). The primary target of the selection imposed by predators is poorly understood. We examined predator (dragonfly larva)-imposed selection on prey (newt larvae) body size and characteristics of escape velocity using replicated and controlled predation experiments under seminatural conditions. Specifically, because these species experience a wide range of temperatures throughout their larval phases, we predict that larvae achieving high swimming velocities across temperatures will have a selective advantage over more thermally sensitive individuals. RESULTS:Nonzero selection differentials indicated that predators selected for prey body size and both absolute and size-corrected maximum swimming velocity. Comparison of selection differentials with control confirmed selection only on body size, i.e., dragonfly larvae preferably preyed on small newt larvae. Maximum swimming velocity and its thermal sensitivity showed low group repeatability, which contributed to non-detectable selection on both characteristics of escape performance. CONCLUSIONS:In the newt-dragonfly larvae interaction, body size plays a more important role than maximum values and thermal sensitivity of swimming velocity during predator escape. This corroborates the general importance of body size in predator-prey interactions. The absence of an appropriate control in predation experiments may lead to potentially misleading conclusions about the primary target of predator-imposed selection. Insights from predation experiments contribute to our understanding of the link between performance and fitness, and further improve mechanistic models of predator-prey interactions and food web dynamics.