Collisions between birds and airplanes, bird strikes, can damage aircrafts, resulting in delays a... more Collisions between birds and airplanes, bird strikes, can damage aircrafts, resulting in delays and cancellation of flights, costing the international civil aviation industry more than 1.4 billion U.S. dollars annually. Bird deterrence is therefore crucial, but the effectiveness of all available deterrence methods is limited. For example, live avian predators can be a highly effective deterrent, because potential prey will not habituate to them, but live predators cannot be controlled with sufficient precision. Thus, there is an urgent need for new deterrence methods. To this end we developed the RobotFalcon, a device that we modelled after the peregrine falcon, a cosmopolitan predator that preys on a large range of bird species. Mimicking natural hunting behaviour, we tested the effectiveness of the RobotFalcon to deter flocks of corvids, gulls, starlings and lapwings. We compared its effectiveness with that of a drone, and of conventional methods routinely applied at a military ai...
This is a PDF file of an article that has undergone enhancements after acceptance, such as the ad... more This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
When a predator attacks a flock of starlings (Sturnus vulgaris), involving thousands of individua... more When a predator attacks a flock of starlings (Sturnus vulgaris), involving thousands of individuals, a typical collective escape response is the so-called agitation wave, consisting of one or more dark bands (pulses) propagating through the flock and moving away from the predator (usually a Peregrine falcon, Falco peregrinus). The mechanism underlying this collective behavior remains debated. A theoretical study has suggested that the individual motion underlying a pulse could be a skitter (in the form of a zigzag), that is copied by nearby neighbors, and causes us to temporarily observe a larger surface of the wing because the bird is banking during turning while zigzagging. It is not known, however, whether pulses during a wave event weaken over time. This is of interest, because whereas during the usual turning by an undisturbed flock the motion is copied completely without weakening, we may expect that pulses dampen during a wave event because individuals that are further away from a predator react less because of reduced fear. In the present paper, we show in empirical data that pulses during a wave event weaken over time. Our computational model, StarDisplay, reveals that this is most likely a consequence of a reduction of the maximum banking angle during the zigzag escape maneuver rather than by a reduced tendency to copy this maneuver with time. The response seems adaptive because of lowered danger at a larger distance to the location of attack. Significance statement Huge flocks of starlings display amazing patterns of collective escape when attacked by an avian predator, such as a Peregrine falcon. One of them is the "agitation wave" in which dark bands move away from the predator. Dark bands arise probably from the temporarily larger wing area, which is observed when birds perform a skitter escape motion (zigzag) while temporarily banking sideward. Whereas during regular flock turns birds copy each other's motion completely, it is unknown whether this happens during agitation waves, because individuals further away from the attack may be less frightened. Studying this both empirically at the group level only and in a computational model at both the level of the individual and the group, we show that pulses of waves fade out with time and that this is probably due to a reduced maximum banking angle during the zigzag maneuver rather than a lower tendency of copying. This seems an adaptive response.
Collective behaviour of animals has been a main focus of recent research, yet few empirical studi... more Collective behaviour of animals has been a main focus of recent research, yet few empirical studies deal with this issue in the context of predation, a major driver of social complexity in many animal species. When starling (Sturnus vulgaris) flocks are under attack by a raptor, such as a peregrine falcon (Falco peregrinus), they show a great diversity of patterns of collective escape. The corresponding structural complexity concerns rapid variation in density and shape of the flock over time. Here, we present a first step towards unravelling this complexity. We apply a time series analysis to video footage of 182 sequences of hunting by falcons on flocks of thousands of starlings close to two urban roosts during winter. We distinguish several types of collective escape by determining the position and movement of individuals relative to each other (which determines darkness and shape of the flock over time) as well as relative to the predator, namely 'flash expansion', 'blackening', 'wave event', 'vacuole', 'cordon' and 'split'. We show that the specific type of collective escape depends on the collective pattern that precedes it and on the level of threat posed by the raptor. A wave event was most likely to occur when the predator attacked at medium speed. Flash expansion occurred more frequently when the predator approached the flock at faster rather than slower speed and attacked from above rather than from the side or below. Flash expansion was often followed by split, but in many cases, the flock showed resilience by remaining intact. During a hunting sequence, the frequencies of different patterns of collective escape increased when the frequency of attack by the raptor was higher. Despite their complexity, we show that patterns of collective escape depend on the predatory threat, which resembles findings in fish. Significance statement Patterns of collective escape in flocks of starlings have always intrigued laymen and scientists. A detailed analysis of their complex dynamics has been lacking so far, and is the focus of our present study: we analysed video footage of hunting by falcons on flocks of thousands of starlings and show how patterns of collective escape (namely flash expansion, blackening, wave event, vacuole, cordon and split) depend on the preceding pattern and on details of attack. A higher frequency of attack during a hunting sequence resulted in a higher frequency of collective escape events. Flash expansion happened most often when the predator attacks at greater speed. A wave event was most likely when the raptor attacks at medium (rather than high or low) speed. These results provide a first quantitative approach to social complexity in collective avoidance of a predator.
Journal of the Marine Biological Association of the United Kingdom, 2015
The identification of foraging hotspots able to support the co-existence of multiple top predator... more The identification of foraging hotspots able to support the co-existence of multiple top predators provides a potential approach to addressing protection measures for marine ecosystems. In this study, we conducted visual surveys in the central Tyrrhenian Sea to determine areas with simultaneous presence of bottlenose dolphins, four species of seabirds (Audouin's gull, Yellow-legged gull, Yelkouan shearwater and Cory's shearwater), and baitballs occurring at the surface, indicating the presence of potential prey items. We also analysed their occurrence in relation to topography (depth, slope and distance from the shore) and seabed types. Kriging analysis identified areas with simultaneous presence of several marine top predators. Dolphin distribution appeared to be linked to gentle slope (6–10 m) and muddy seabed, possibly associated with prey distribution, whereas the four seabird species were more frequent in areas with a water depth of 100–150 m, gentle slope and muddy sea...
Avian embryos are exposed not only to endogenous sex steroids, which are produced by their gonads... more Avian embryos are exposed not only to endogenous sex steroids, which are produced by their gonads and have a key role in sexual differentiation, but also to maternal steroids transferred into the egg yolk, which can modulate the development of individual differences in behavior. Studies of maternal hormones have primarily focused on ultimate questions (evolutionary trade-offs, functional significance), whereas proximate mechanistic questions have been largely ignored. A central problem that must be addressed is how exposure to maternal hormones affects the individual phenotype without interfering with sexual differentiation. Separate effects could result from the action of different hormones, at different doses or at different times, on different targets.
associated with synaptic plasticity and synaptogenesis [4,8,9]. These socially related learning m... more associated with synaptic plasticity and synaptogenesis [4,8,9]. These socially related learning mechanisms mediate more than just mate choice and sexual responses, affecting social-affiliative processes, including the formation of pair bonds. This is in agreement with the ideas of social selection, whereby social selection embraces all forms of selection, including sexual selection, driven by social interactions [12]. Because social recognition, mate choice, and mate preferences are strongly intertwined it is highly likely that these co-evolved and are making use of similar neurobiological mechanisms. Social learning, by contrast, may differ with the emphasis being on the need to pay attention to salient information from social sources for a variety of behavioral responses, including that of mate choice. Overall, this large overlap in the conservation and utilization of neural mechanisms supports common neurobiological links between social behavior, social learning, social recognition, and group membership and individual mate choice and preferences.
Collisions between birds and airplanes, bird strikes, can damage aircrafts, resulting in delays a... more Collisions between birds and airplanes, bird strikes, can damage aircrafts, resulting in delays and cancellation of flights, costing the international civil aviation industry more than 1.4 billion U.S. dollars annually. Bird deterrence is therefore crucial, but the effectiveness of all available deterrence methods is limited. For example, live avian predators can be a highly effective deterrent, because potential prey will not habituate to them, but live predators cannot be controlled with sufficient precision. Thus, there is an urgent need for new deterrence methods. To this end we developed the RobotFalcon, a device that we modelled after the peregrine falcon, a cosmopolitan predator that preys on a large range of bird species. Mimicking natural hunting behaviour, we tested the effectiveness of the RobotFalcon to deter flocks of corvids, gulls, starlings and lapwings. We compared its effectiveness with that of a drone, and of conventional methods routinely applied at a military ai...
This is a PDF file of an article that has undergone enhancements after acceptance, such as the ad... more This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
When a predator attacks a flock of starlings (Sturnus vulgaris), involving thousands of individua... more When a predator attacks a flock of starlings (Sturnus vulgaris), involving thousands of individuals, a typical collective escape response is the so-called agitation wave, consisting of one or more dark bands (pulses) propagating through the flock and moving away from the predator (usually a Peregrine falcon, Falco peregrinus). The mechanism underlying this collective behavior remains debated. A theoretical study has suggested that the individual motion underlying a pulse could be a skitter (in the form of a zigzag), that is copied by nearby neighbors, and causes us to temporarily observe a larger surface of the wing because the bird is banking during turning while zigzagging. It is not known, however, whether pulses during a wave event weaken over time. This is of interest, because whereas during the usual turning by an undisturbed flock the motion is copied completely without weakening, we may expect that pulses dampen during a wave event because individuals that are further away from a predator react less because of reduced fear. In the present paper, we show in empirical data that pulses during a wave event weaken over time. Our computational model, StarDisplay, reveals that this is most likely a consequence of a reduction of the maximum banking angle during the zigzag escape maneuver rather than by a reduced tendency to copy this maneuver with time. The response seems adaptive because of lowered danger at a larger distance to the location of attack. Significance statement Huge flocks of starlings display amazing patterns of collective escape when attacked by an avian predator, such as a Peregrine falcon. One of them is the "agitation wave" in which dark bands move away from the predator. Dark bands arise probably from the temporarily larger wing area, which is observed when birds perform a skitter escape motion (zigzag) while temporarily banking sideward. Whereas during regular flock turns birds copy each other's motion completely, it is unknown whether this happens during agitation waves, because individuals further away from the attack may be less frightened. Studying this both empirically at the group level only and in a computational model at both the level of the individual and the group, we show that pulses of waves fade out with time and that this is probably due to a reduced maximum banking angle during the zigzag maneuver rather than a lower tendency of copying. This seems an adaptive response.
Collective behaviour of animals has been a main focus of recent research, yet few empirical studi... more Collective behaviour of animals has been a main focus of recent research, yet few empirical studies deal with this issue in the context of predation, a major driver of social complexity in many animal species. When starling (Sturnus vulgaris) flocks are under attack by a raptor, such as a peregrine falcon (Falco peregrinus), they show a great diversity of patterns of collective escape. The corresponding structural complexity concerns rapid variation in density and shape of the flock over time. Here, we present a first step towards unravelling this complexity. We apply a time series analysis to video footage of 182 sequences of hunting by falcons on flocks of thousands of starlings close to two urban roosts during winter. We distinguish several types of collective escape by determining the position and movement of individuals relative to each other (which determines darkness and shape of the flock over time) as well as relative to the predator, namely 'flash expansion', 'blackening', 'wave event', 'vacuole', 'cordon' and 'split'. We show that the specific type of collective escape depends on the collective pattern that precedes it and on the level of threat posed by the raptor. A wave event was most likely to occur when the predator attacked at medium speed. Flash expansion occurred more frequently when the predator approached the flock at faster rather than slower speed and attacked from above rather than from the side or below. Flash expansion was often followed by split, but in many cases, the flock showed resilience by remaining intact. During a hunting sequence, the frequencies of different patterns of collective escape increased when the frequency of attack by the raptor was higher. Despite their complexity, we show that patterns of collective escape depend on the predatory threat, which resembles findings in fish. Significance statement Patterns of collective escape in flocks of starlings have always intrigued laymen and scientists. A detailed analysis of their complex dynamics has been lacking so far, and is the focus of our present study: we analysed video footage of hunting by falcons on flocks of thousands of starlings and show how patterns of collective escape (namely flash expansion, blackening, wave event, vacuole, cordon and split) depend on the preceding pattern and on details of attack. A higher frequency of attack during a hunting sequence resulted in a higher frequency of collective escape events. Flash expansion happened most often when the predator attacks at greater speed. A wave event was most likely when the raptor attacks at medium (rather than high or low) speed. These results provide a first quantitative approach to social complexity in collective avoidance of a predator.
Journal of the Marine Biological Association of the United Kingdom, 2015
The identification of foraging hotspots able to support the co-existence of multiple top predator... more The identification of foraging hotspots able to support the co-existence of multiple top predators provides a potential approach to addressing protection measures for marine ecosystems. In this study, we conducted visual surveys in the central Tyrrhenian Sea to determine areas with simultaneous presence of bottlenose dolphins, four species of seabirds (Audouin's gull, Yellow-legged gull, Yelkouan shearwater and Cory's shearwater), and baitballs occurring at the surface, indicating the presence of potential prey items. We also analysed their occurrence in relation to topography (depth, slope and distance from the shore) and seabed types. Kriging analysis identified areas with simultaneous presence of several marine top predators. Dolphin distribution appeared to be linked to gentle slope (6–10 m) and muddy seabed, possibly associated with prey distribution, whereas the four seabird species were more frequent in areas with a water depth of 100–150 m, gentle slope and muddy sea...
Avian embryos are exposed not only to endogenous sex steroids, which are produced by their gonads... more Avian embryos are exposed not only to endogenous sex steroids, which are produced by their gonads and have a key role in sexual differentiation, but also to maternal steroids transferred into the egg yolk, which can modulate the development of individual differences in behavior. Studies of maternal hormones have primarily focused on ultimate questions (evolutionary trade-offs, functional significance), whereas proximate mechanistic questions have been largely ignored. A central problem that must be addressed is how exposure to maternal hormones affects the individual phenotype without interfering with sexual differentiation. Separate effects could result from the action of different hormones, at different doses or at different times, on different targets.
associated with synaptic plasticity and synaptogenesis [4,8,9]. These socially related learning m... more associated with synaptic plasticity and synaptogenesis [4,8,9]. These socially related learning mechanisms mediate more than just mate choice and sexual responses, affecting social-affiliative processes, including the formation of pair bonds. This is in agreement with the ideas of social selection, whereby social selection embraces all forms of selection, including sexual selection, driven by social interactions [12]. Because social recognition, mate choice, and mate preferences are strongly intertwined it is highly likely that these co-evolved and are making use of similar neurobiological mechanisms. Social learning, by contrast, may differ with the emphasis being on the need to pay attention to salient information from social sources for a variety of behavioral responses, including that of mate choice. Overall, this large overlap in the conservation and utilization of neural mechanisms supports common neurobiological links between social behavior, social learning, social recognition, and group membership and individual mate choice and preferences.
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