Causes of variation of darkness in flocks of starlings, a computational model

The coordinated motion of large flocks of starlings is fascinating for both laymen and scientists. During their aerial displays, the darkness of flocks often changes, for instance dark bands propagate through the flock (so-called agitation waves) and small or large parts of the flock darken. The causes of dark bands in agitation waves have recently been shown to depend on changes in orientation of birds relative to the observer rather than changes in density of the flock, but what causes other changes in darkness need to be studied still and this is the aim of the present investigation. Because we cannot empirically relate changes in darkness in flocks to quantities, such as position and orientation of the flock and of its members relative to the observer, we study this in a computational model. We use StarDisplay, a model of collective motion of starlings, because its flocks resemble empirical data in many properties, such as their three-dimensional shape, their manner of turning, the correlation of heading of its group-members, and its internal structure regarding density and stability of neighbors. We show that the change in darkness in the flocks perceived by an observer on the ground mostly depends on the observer’s distance to the flock and on the degree of exposure of the wing surface of flock members to the observer, and that darkness appears to decrease when birds roll during sharp turns. Remarkably, the darkness of the flock perceived by the observer was neither affected by the orientation of the flock relative to the observer nor by the density of the flock. Further studies are needed to investigate changes in darkness for flocks under predation

Damping of waves of agitation in starling flocks

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

Calibration of multiple cameras for large-scale experiments using a freely moving calibration target

Abstract: Obtaining accurate experimental data from Lagrangian tracking and tomographic velocimetry requires an accurate camera calibration consistent over multiple views. Established calibration procedures are often challenging to implement when the length scale of the measurement volume exceeds that of a typical laboratory experiment. Here, we combine tools developed in computer vision and non-linear camera mappings used in experimental fluid mechanics, to successfully calibrate a four-camera setup that is imaging inside a large tank of dimensions ∼10×25×6m3. The calibration procedure uses a planar checkerboard that is arbitrarily positioned at unknown locations and orientations. The method can be applied to any number of cameras. The parameters of the calibration yields direct estimates of the positions and orientations of the four cameras as well as the focal lengths of the lenses. These parameters are used to assess the quality of the calibration. The calibration allows us to perform accurate and consistent linear ray-tracing, which we use to triangulate and track fish inside the large tank. An open-source implementation of the calibration in Matlab is available. Graphic abstract: [Figure not available: see fulltext.]

Complex patterns of collective escape in starling flocks under predation

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 statementPatterns 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

The Effect of Sensory Blind Zones on Milling Behavior in a Dynamic Self-Propelled Particle Model

Emergent pattern formation in self-propelled particle (SPP) systems is extensively studied because it addresses a range of swarming phenomena which occur without leadership. Here we present a dynamic SPP model in which a sensory blind zone is introduced into each particle's zone of interaction. Using numerical simulations we discovered that the degradation of milling patterns with increasing blind zone ranges undergoes two distinct transitions, including a new, spatially nonhomogeneous transition that involves cessation of particles' motion caused by broken symmetries in their interaction fields. Our results also show the necessity of nearly complete panoramic sensory ability for milling behavior to emerge in dynamic SPP models, suggesting a possible relationship between collective behavior and sensory systems of biological organisms.Comment: 12 pages, 4 figure

An Individual-Oriented Model on the Emergence of Support in Fights, Its Reciprocation and Exchange

Complex social behaviour of primates has usually been attributed to the operation of complex cognition. Recently, models have shown that constraints imposed by the socio-spatial structuring of individuals in a group may result in an unexpectedly high number of patterns of complex social behaviour, resembling the dominance styles of egalitarian and despotic species of macaques and the differences between them. This includes affiliative patterns, such as reciprocation of grooming, grooming up the hierarchy, and reconciliation. In the present study, we show that the distribution of support in fights, which is the social behaviour that is potentially most sophisticated in terms of cognitive processes, may emerge in the same way. The model represents the spatial grouping of individuals and their social behaviour, such as their avoidance of risks during attacks, the self-reinforcing effects of winning and losing their fights, their tendency to join in fights of others that are close by (social facilitation), their tendency to groom when they are anxious, the reduction of their anxiety by grooming, and the increase of anxiety when involved in aggression. Further, we represent the difference in intensity of aggression apparent in egalitarian and despotic macaques. The model reproduces many aspects of support in fights, such as its different types, namely, conservative, bridging and revolutionary, patterns of choice of coalition partners attributed to triadic awareness, those of reciprocation of support and ‘spiteful acts’ and of exchange between support and grooming. This work is important because it suggests that behaviour that seems to result from sophisticated cognition may be a side-effect of spatial structure and dominance interactions and it shows that partial correlations fail to completely omit these effects of spatial structure. Further, the model is falsifiable, since it results in many patterns that can easily be tested in real primates by means of existing data

Some Causes of the Variable Shape of Flocks of Birds

Flocks of birds are highly variable in shape in all contexts (while travelling, avoiding predation, wheeling above the roost). Particularly amazing in this respect are the aerial displays of huge flocks of starlings (Sturnus vulgaris) above the sleeping site at dawn. The causes of this variability are hardly known, however. Here we hypothesise that variability of shape increases when there are larger local differences in movement behaviour in the flock. We investigate this hypothesis with the help of a model of the self-organisation of travelling groups, called StarDisplay, since such a model has also increased our understanding of what causes the oblong shape of schools of fish. The flocking patterns in the model prove to resemble those of real birds, in particular of starlings and rock doves. As to shape, we measure the relative proportions of the flock in several ways, which either depend on the direction of movement or do not. We confirm that flock shape is usually more variable when local differences in movement in the flock are larger. This happens when a) flock size is larger, b) interacting partners are fewer, c) the flock turnings are stronger, and d) individuals roll into the turn. In contrast to our expectations, when variability of speed in the flock is higher, flock shape and the positions of members in the flock are more static. We explain this and indicate the adaptive value of low variability of speed and spatial restriction of interaction and develop testable hypotheses