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The European hare Lepus europaeus is an iconic but rapidly declining farmland specialist with recently confirmed populations in urban areas. However, their behavioral responses and adaptability to urbanization and life in human-dominated areas are fully unexplored. Here, using infrared cameras, we explored escape behavior (measured as the flight initiation distance) using 965 hare observations in urban and farmland areas in the Czech Republic and Austria (Central Europe) and its association with habitat type, distance to the city center, patch size, season, hare age and initial behavior. We found that European hares adjusted their escape behavior to habitat type and escaped significantly earlier in farmland (rural) habitats than in urban habitats. However, escape distances of hares did not differ between farmland types with different degree of habitat heterogeneity. We also revealed that urban hares escaped earlier when located further from the city center or in a larger study patch. Moreover, adult hares escaped earlier than subadults and foraging individuals escaped earlier than resting hares but both only in rural areas. Our results support high behavioral adaptability of European hares to human-dominated urban areas.

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Anthropogenic noise can affect a number of behavioral, physiological, and ecological aspects of animals from major taxonomic groups, raising serious conservation concerns. For example, noise pollution impacts communicative behavior and perception of signals, movements and distribution, as well as predator-prey interactions, such as hunting success or predator detection and predation risk assessment. We have carried out an experimental playback study, in which we investigated whether exposure to anthropogenic noise (sound of a tractor) distracts free-ranging barn swallows Hirundo rustica from paying attention to an approaching human "predator" (the "cognitive distraction" hypothesis), or whether noise leads to increased responsiveness to this "predator" (the "increased threat" hypothesis). The subjects were male barn swallows attending their breeding territories during the time when the females were incubating. We found that barn swallow males initiated flight at significantly greater distances to the approaching human "predator" in the noise treatment than during the quiet control trials. These results suggest that anthropogenic noise causes increased vigilance and reactivity rather than a distraction, enabling birds to avoid the "predator" more quickly. We further discuss the mechanism behind the increased alertness in response to noise and contrast the "increased threat" mechanism, usually tested in previous studies, with an alternative "cognitive sensitization" mechanism.

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Temporal changes in environmental conditions may play a major role in the year-to-year variation in fitness consequences of behaviours. Identifying environmental drivers of such variation is crucial to understand the evolutionary trajectories of behaviours in natural contexts. However, our understanding of how environmental variation influences behaviours in the wild remains limited. Using data collected over 14 breeding seasons from a collared flycatcher (Ficedula albicollis) population, we examined the effect of environmental variation on the relationship between survival and risk-taking behaviour, a highly variable behavioural trait with great evolutionary and ecological significance. Specifically, using annual recapture probability as a proxy of survival, we evaluated the specific effect of predation pressure, food availability, and mean temperature on the relationship between annual recapture probability and risk-taking behaviour (measured as flight initiation distance [FID]). We found a negative trend, as the relationship between annual recapture probability and FID decreased over the study years and changed from positive to negative. Specifically, in the early years of the study, risk-avoiding individuals exhibited a higher annual recapture probability, whereas in the later years, risk-avoiders had a lower annual recapture probability. However, we did not find evidence that any of the considered environmental factors mediated the variation in the relationship between survival and risk-taking behaviour.

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Visual cues are important in both interspecific and intraspecific communication. The species confidence hypothesis proposes that animals are more attracted to conspecific colors and repelled by colors, not on their bodies. Studies on terrestrial lizards and birds have tested the species confidence hypothesis and shown that conspecific colors elicit reduced antipredator behavior. To date, the species confidence hypothesis has not been tested in the marine environment, specifically on coral reefs where color communication is of vital importance. We addressed this knowledge gap by measuring flight initiation distance (the distance an individual moves away from an approaching threat) in dusky damselfish (Stegastes nigricans) in response to an approaching disc of 1 of 4 different color treatments: conspecific, blue, yellow, and black. If the species confidence hypothesis explained variation in damselfish flight initiation distance, then we expected individuals to tolerate closer approaches when approached by a conspecific color. In addition, we calculated the color difference between each stimulus and its corresponding background as a potential alternative explanation for flight responses. Damselfish tolerated the closest approach from the conspecific color stimulus; there were no significant differences between other colors and there was no support for the alternative color difference hypothesis. As with similar terrestrial studies, these results are relevant to ecotourists' choice of swimsuit and wetsuit colors because color choice may modify natural antipredator behavior.

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The 2019 zoonotic pandemic (COVID-19), has led to a massive global lockdown that provides a good opportunity to study how wildlife responds to changes in human activity. Wearing a mask after the COVID-19 outbreak was widely used to prevent the spread of the causative pathogen. It has been shown that tree sparrows (Passer montanus) at two sites in south China exhibit reduced fear responses to people with face masks after a period of heavy exposure to them, whereas European studies showed the opposite, with no changes in the behaviour of the birds towards mask wearers in either rural or urban areas. To further study this, from October 2021 to January 2022, we conducted a flight initiation distance (FID) survey in Pakistan, Bangladesh, and Xi'an, China for a variety of field bird species by comparing the FID for researchers wearing masks to that for researchers not wearing masks to assess whether wearing masks in public places caused birds to adjust their flight response. Results from the three Asian countries showed that after a period of sustained contact with people wearing masks, in both rural and urban areas, birds were significantly more adapted to them and had a shorter FID to people wearing masks. We suggest that the rapid habituation of birds to people wearing masks with a reduced fear response could have some fitness advantage, allowing them to adapt rapidly to the new environmental conditions induced by COVID-19.

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Recognizing, assessing, and responding to threats is critical for survival in the wild. Birds, especially in their role as parents, must decide whether to flee or delay flight when threatened. This study examines how age, reproductive stage, and the presence of a mate influence flight initiation distance (FID) and nest recess duration in white storks. Analyzing the data with a generalized additive mixed model (GAMM), we found significant correlations between FID and age, reproductive stage, and presence of a mate. These results suggest that the trade-off between current and future reproduction shifts during critical breeding periods, such as incubation and nestling care. To increase breeding success, White Storks appear willing to take risks and extend their stay in the nest when offspring are most valuable and vulnerable. In the presence of a mate, individuals leave the nest earlier, suggesting possible sexual conflict over parental care. The duration of nest abandonment is consistent with FID, except for age. These results illustrate how parental age, brood value, vulnerability, and sexual dynamics influence white stork flight decisions in complex ways. Understanding these dynamics enriches our knowledge of bird behavior and adaptations to environmental challenges and highlights the complexity of parental decision making.

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Following the COVID-19 pandemic, many people around the world stayed home, drastically altering human activity in cities. This exceptional moment provided researchers the opportunity to test how urban animals respond to human disturbance, in some cases testing fundamental questions on the mechanistic impact of urban behaviours on animal behaviour. However, at the end of this 'anthropause', human activity returned to cities. How might each of these strong shifts affect wildlife in the short and long term? We focused on fear response, a trait essential to tolerating urban life. We measured flight initiation distance-at both individual and population levels-for an urban bird before, during and after the anthropause to examine if birds experienced longer-term changes after a year and a half of lowered human presence. Dark-eyed juncos did not change fear levels during the anthropause, but they became drastically less fearful afterwards. These surprising and counterintuitive findings, made possible by following the behaviour of individuals over time, has led to a novel understanding that fear response can be driven by plasticity, yet not habituation-like processes. The pandemic-caused changes in human activity have shown that there is great complexity in how humans modify a behavioural trait fundamental to urban tolerance in animals.

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Human disturbance, particularly road traffic, is one of the greatest threats to wildlife. Considering the association between alerting behavior and the survival of animals, it is important to study the effects of road traffic on alerting behavior of wildlife. Previous studies assessing the short-term impact of road traffic on alerting behavior of wildlife have focused on vigilance distances. However, studies on the use of alarm calls are scarce, and it is unclear whether such behavioral responses change after repeated exposure to road traffic. We assessed the alerting behavior of plateau pikas (Ochotona curzoniae) who were near or far from roads when facing a potential predator. We found that pikas near roads exhibited shorter vigilance and tolerance distances, and produced fewer alarm calls than those relatively far away from roads. Furthermore, both vigilance and tolerance distances of plateau pikas were significantly positively correlated with the distance from the burrow to the road. Road traffic reduced antipredator responses and shaped alerting behavior; that is, pikas near roads were bolder and more silent compared to those far away from roads. Our findings suggest that increasing urbanization will have corresponding effects on animal behavior, which may have significant fitness effects in the future.

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Escape behavior is a common antipredator strategy among wild animals. Here, we investigated the effect of four factors on the vigilance behavior of the endemic Cyprus rock agama (Laudakia cypriaca). Flight initiation distance (FID, the minimum distance to which an observer can approach a lizard before it flees) was measured in relation to the type of location (tourist vs. nontourist area), the observer's starting distance, air temperature, and substrate temperature. We collected data for 39 agamas in tourist areas and 34 of these lizards in nontourist areas. As a whole, the mean starting distance was 10.5 m and the FID was 3.6 m. The average substrate temperature was 34.0°C and the average air temperature 29.6°C. Only the type of area affected the agamas' escape decisions with FID being 1.8 m shorter in tourist areas than in nontourist areas (2.7 m vs. 4.5 m). This is probably due to the habituation of lizards to the presence of humans in the former areas. This study shows that tourism strongly affects the behavior of lizards, which may have consequences for the functioning of the population. Tourists can increase the safety of lizards by creating a human shield to deter predators. Once the tourist season is over, lizards may become more vulnerable to predators.

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In predator-prey interactions, various factors affect the prey's perception of risk and decision to flee. Gaze sensitivity, the ability to react to the presence, direction, or movement of the head and eyes, has been reported in many birds. However, few studies have focussed on variation in sensitivity to human gaze in relation to other risks and potential breeding costs. Here, we studied the influence of human gaze on the escape behaviour of Azure-winged magpies (Cyanopica cyanus) and investigated the effects of breeding state (breeding season and nonbreeding season) and approach direction on gaze sensitivity. In Experiment 1, we tested whether magpies showed different sensitivities to human gaze according to age class and breeding state when approached directly. The results showed that the breeding state could affect the flight initiation distance (FID), with adults in the breeding season having a shorter FID compared to those in the nonbreeding season. Meanwhile, only adults were found to be averse to direct human gaze and juveniles showed no sensitivity. In Experiment 2, we conducted three different gaze treatments on adult magpies in the breeding season under three bypass distances (0 m, 2.5 m, 5 m). The results showed that approach direction had no effect on FID, while the sensitivity to human gaze differed under three bypass distances. Adults could clearly recognise human head and eye direction at a certain bypass distance (2.5 m). Our study reveals the cognitive ability of Azure-winged magpies to human head and eye direction and the effects of age, breeding state and approach direction, which may provide further insights into human-wildlife interactions, especially for birds in urban habitats.

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With the increasing urbanization of the last decades, more and more bird species occur in urban habitats. Birds which thrive in urban habitats often have a higher tolerance toward human disturbance and show behaviors which differ from their rural counterparts. There is increasing evidence that many behaviors have a genetic basis. One candidate gene is the dopamine receptor D4 (DRD4), which has been associated with fear and thus, flight initiation distance (FID). In this study, we analyzed a segment of DRD4 in greylag geese Anser anser, describing the variability of this gene across several geographically distant populations, and comparing its variability between an urban and a rural site in south-west Germany. We additionally measured FIDs of urban and rural geese to test for a possible correlation with DRD4 genotypes. We found a high variation within DRD4, with 10 variable sites leading to 11 alleles and 35 genotypes. Two genotypes occurred in 60% of all geese and were thus defined as common genotypes versus 33 rare genotypes. Population differentiation was very low between the urban and rural sites in Germany but common genotypes occurred more often in the urban area and rare genotypes more often in the rural area. FID was significantly higher at the rural site, but no significant correlation between FID and DRD4 genotypes could be detected. Nevertheless, our results suggest that local site selection may be related to DRD4 genotypes.

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The Covid-19 lockdown reduced drastically human presence outdoors, providing an uncontrolled experiment for disentangling direct and indirect effects of human presence on animal fearfulness. We measured 18,494 flight initiation distances (FIDs, the distance at which individual animals fly away when approached by a human) from 1333 populations of 202 bird species taken in four European cities both before, during and after the lockdown. FIDs decreased during lockdown in rural habitats but increased in urban habitats, especially for singing birds. Height above ground increases during lockdown in non-singing birds only, and birds adjusted horizontal tolerance to approach according to height outside lockdown, in rural habitats and while not singing. Responses showed lagged effects after lockdown in urban but not in rural habitats. Differential responses to lockdown among habitats and between signing and non-singing birds were consistent with relaxation of direct disturbance effects on birds in rural habitats during lockdown, as well as with increased indirect fear effects mediated by predator release in cities. FIDs seemed to measure the balance of direct and indirect effects of humans on predations risk and food needs rather than direct effects of humans on fear.

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Human-induced disturbances affect animal behaviours such as anti-predatory responses. Animals in urban environments tend to exhibit a reduced escape response, measured as a shorter flight initiation distance (FID), compared to their rural counterparts. While FID has been evaluated in animals dwelling in contrasting habitats (e.g. urban versus rural), little is known about how this response varies within urban environments, especially in tropical cities. Here, we studied the FID of 15 resident bird species in Bogota, Colombia, at 22 sites grouped into four categories (natural sites, metropolitan parks, zonal parks and residential areas) that differed in landscape features and evaluated which factors affected the escape responses of birds. We showed that birds foraging in larger flocks are more tolerant when being approached but they do not seem to be influenced by other factors such as heterospecific flock size, noise levels, pedestrian density, predator density, natural cover or body length. Also, birds inhabiting residential areas and parks showed a shorter FID compared to birds in natural areas suggesting that they are more tolerant of human-related disturbances compared to their conspecifics that live in natural areas within the city. Our study shows important differences in bird anti-predatory responses within the city and suggests that social strategies (i.e. flocking patterns) may be a mechanism for adapting to human-induced disturbances in urban tropical environments.

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Plastic and selective mechanisms govern parental investment adjustments to predation threat. We investigated the relative importance of plasticity and selection in risk-taking propensity of incubating female common eiders Somateria mollissima facing unprecedented predation in SW Finland, Baltic Sea. Using a 12-year individual-based dataset, we examined within- and among-individual variation in flight initiation distance (FID), in relation to predation risk, nest detectability, individual traits and reproductive investment (NFID = 1009; Nindividual = 559). We expected females nesting in riskier environments (higher predation risk, lower nest concealment) to mitigate environmentally imposed risk by exhibiting longer FIDs, and females investing more in current reproduction (older, in better condition or laying larger clutches) to display shorter FIDs. The target of predation-adult or offspring-affected the mechanisms adapting risk-taking propensity; females plastically increased their FID under higher adult predation risk, while risk-avoiding breeders were predominant on islands with higher nest predation risk. Risk-taking females selected thicker nest cover, consistent with personality-matching habitat choice. Females plastically attenuated their anti-predator response (shorter FIDs) with advancing age, and females in better body condition were more risk-taking, a result explained by selection processes. Future research should consider predator type when investigating the fitness consequences of risk-taking strategies.

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Flight initiation distance (FID) procedures are used to assess the risk perception animals have for threats (e.g., natural predators, hunters), but it is unclear whether these assessments remain meaningful if animals have habituated to certain human stimuli (e.g., researchers, tourists). Our previous work showed that habituated baboons displayed individually distinct and consistent responses to human approaches, a tolerance trait, but it is unknown if the trait is resilient to life-threatening scenarios. If it were consistent, it would imply FIDs might measure specific human threat perception only and not generalize to other threats such as predators when animals have experienced habituation processes. We used FID procedures to compare baseline responses to the visual orientation distance, FID, and individual tolerance estimates assessed after a leopard predation on an adult male baboon (group member). All variables were consistent despite the predation event, suggesting tolerance to observers was largely unaffected by the predation and FID procedures are unlikely to be generalizable to other threats when habituation has occurred. FID approaches could be an important tool for assessing how humans influence animal behavior across a range of contexts, but careful planning is required to understand the type of stimuli presented.

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AbstractWar influences wildlife in a variety of ways but may influence their escape responses to approaching threats, including humans, because of its effect on human populations and behavior and landscape change. We collected 1,400 flight initiation distances (FIDs) from 157 bird species in the dry zone of Sri Lanka, where civil war raged for 26 years, ending in 2009. Accounting for factors known to influence FIDs (phylogeny, starting distance of approaches, body mass, prevailing human density, group size, and location), we found that birds have longer FIDs in the part of the dry zone that experienced civil war. Larger birds-often preferred by human hunters-showed greater increases in FID in the war zone, consistent with the idea that war was associated with greater hunting pressure and that larger birds experienced longer-lasting trauma or had more plastic escape behavior than smaller species. While the mechanisms linking the war and avian escape responses remain ambiguous, wars evidently leave legacies that extend to behavioral responses in birds.

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Flash behaviour is widespread in the animal kingdom and describes the exposure of a hidden conspicuous signal as an animal flees from predators. Recent studies have demonstrated that the signal can enhance survivorship by leading pursuing predators into assuming the flasher is also conspicuous at rest. Naturally, this illusion will work best if potential predators are ignorant of the flasher's resting appearance, which could be achieved if the prey flees while the predator is relatively far away. To test this hypothesis, we compared the survival of flashing and non-flashing computer-generated prey with different flight initiation distances (FIDs) using humans as model predators. This experiment found that flash displays confer a survivorship advantage only to those prey with a long FID. A complementary phylogenetic analysis of Australian bird species supports these results: after controlling for body size, species with putative flashing signals had longer FIDs than those without. Species with putative flashing signals also tended to be larger, as demonstrated in other taxa. The anti-predation benefit of flash displays is therefore related to the nature of escape behaviour. Since birds with hidden signals tend to flee at a distance, the flash display here is unlikely to function by startling would-be predators.

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Individuals which have invaded urbanized environments are reported to engage in riskier behaviors, possibly influenced by the scarcity of predators in urbanized areas. Here, we studied the risk-taking behavior of birds which had invaded a new natural environment, rather than an artificial urban environment, using recently established populations of the bull-headed shrike Lanius bucephalus, which naturally colonized three subtropical islands in Japan. We compared flight initiation distance (FID), the distance at which an individual approached by a human initiates flight, between the islands and the temperate mainland. FID was longer for the insular shrikes compared with the mainland shrikes after controlling for other factors, indicating that the individuals which had invaded a new natural environment had a lower propensity for risk-taking. A possible explanation for these results is that low risk-taking behavior might be adaptive on the islands due to predation by the black rat Rattus rattus, an unfamiliar predator not found in shrike habitats on the temperate mainland. Further studies are needed to examine the nest predation rate, predator species, and nest site selection of these insular shrike populations.

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Understanding how vulnerable species are to new stressors, such as anthropogenic changes, is crucial for mitigating their potential negative consequences. Many studies have investigated species sensitivity to human disturbance by focusing on single behavioral or physiological parameters, such as flight initiation distance and glucocorticoid levels. However, little is known about the differential effect that modulating factors might have on behavioral versus physiological stress responses across species. This lack of knowledge make difficult to understand the relationship between both types of reactions, and thus to assess to what extent a behavioral reaction is representative of an internal physiological stress response or vice versa. We collected published data on bird flight initiation distances (FID) and corticosterone (CORT) responses, the two most frequently used indicators of stress reaction. We then investigated how spatio-temporal factors or species-specific characteristics relate to these behavioral and physiological stress responses, and potentially modify the relationship between them. Additionally, we evaluated the strength of the correlation between the two stress responses (behavioral and physiological). Our findings showed that FID and CORT responses were poorly correlated across species, and the lack of correlation was attributable to modulating factors (e.g. latitude and body mass) which influence behavior and physiology differently. These modulating factors, therefore, should be taken into consideration to better interpret FID and CORT responses in the context of species vulnerability to stress.

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Humans have shaped carnivore behavior since at least the Middle Paleolithic period, about 42,000 years ago. In more recent times, spotted hyenas (Crocuta crocuta) in Ethiopia have adapted to living in urban areas, while humans have adapted to living with hyenas. Yet, relationships between coexisting humans and carnivores are rarely addressed beyond mitigating conflicts. We provided a case study for how to broadly think about coexistence and how to study it when measuring if humans and carnivores affect one another. We collected data in four Ethiopian cities: Mekelle, Harar, Addis Ababa, and Arba Minch. We held focus groups and key informant interviews that incorporated feedback from 163 people, representing a wide array of religious, economic, and educational backgrounds. We also determined how many hyenas resided in these cities, hyena behavioral responses to humans using a flight initiation test, and problem-solving abilities via puzzle box trials. We found that in three of the cities, hyenas and humans coexist at high densities and frequently encounter each other. While all participants recognized the importance of hyenas as scavengers to maintain a clean environment, there was pronounced variation in cultural perspectives across cities. For example, while the people of Harar revere hyenas in spiritual terms, in Arba Minch hyenas were regarded as nuisance animals. Hyenas were universally respected as a formidable predator across cities but reports of attacks on livestock and humans were few. Flight initiation tests revealed hyenas fled at significantly closer distances in Harar and Addis Ababa than in Mekelle. Hyenas succeeded at solving a puzzle box in Harar but not in Mekelle. These variable behavior in hyenas correlated to different human perceptions. Our case study results suggest that the hyena-human dynamic is highly variable across these locations. We conclude by exploring the implications of these findings for how humans and hyenas can shape one another's behavior. Developing studies to link human perceptions and animal behavior could advance wildlife conservation, especially in urban areas.