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The dynamical critical exponent $z$ of natural swarms of insects is calculated using the renormalization group to order $\epsilon = 4-d$. A novel fixed point emerges, where both activity and inertia are relevant. In three dimensions the critical exponent at the new fixed point is $z = 1.35$, in agreement with both experiments ($1.37 \pm 0.11$) and numerical simulations ($1.35 \pm 0.04$)., Comment: Numerical simulations to validate RG predictions added. Experimental assesment of the dynamic critical exponent revised. 72 pages, 24 figures, 1 table

Speed fluctuations of individual birds in natural flocks are moderate, due to the aerodynamic and biomechanical constraints of flight. Yet the spatial correlations of such fluctuations are scale-free, namely they have a range as wide as the entire group, a property linked to the capacity of the system to collectively respond to external perturbations. Scale-free correlations and moderate fluctuations set conflicting constraints on the mechanism controlling the speed of each agent, as the factors boosting correlation amplify fluctuations, and vice versa. Here, using a statistical field theory approach, we suggest that a marginal speed confinement that ignores small deviations from the natural reference value while ferociously suppressing larger speed fluctuations, is able to reconcile scale-free correlations with biologically acceptable group's speed. We validate our theoretical predictions by comparing them with field experimental data on starling flocks with group sizes spanning an unprecedented interval of over two orders of magnitude., Comment: 18 pages, 5 figures. Latest version before publication uploaded, journal reference added

Many living and complex systems exhibit second order emergent dynamics. Limited experimental access to the configurational degrees of freedom results in data that appears to be generated by a non-Markovian process. This poses a challenge in the quantitative reconstruction of the model from experimental data, even in the simple case of equilibrium Langevin dynamics of Hamiltonian systems. We develop a novel Bayesian inference approach to learn the parameters of such stochastic effective models from discrete finite length trajectories. We first discuss the failure of naive inference approaches based on the estimation of derivatives through finite differences, regardless of the time resolution and the length of the sampled trajectories. We then derive, adopting higher order discretization schemes, maximum likelihood estimators for the model parameters that provide excellent results even with moderately long trajectories. We apply our method to second order models of collective motion and show that our results also hold in the presence of interactions., Comment: we correct previous inaccuracy about a reference; 29 pages, 9 figures

We study the critical behaviour of a model with non-dissipative couplings aimed at describing the collective behaviour of natural swarms, using the dynamical renormalization group. At one loop, we find a crossover between a conservative yet unstable fixed point, characterized by a dynamical critical exponent $z=d/2$, and a dissipative stable fixed point with $z=2$, a result we confirm through numerical simulations. The crossover is regulated by a conservation length scale that is larger the smaller the effective friction, so that in finite-size biological systems with low dissipation, dynamics is ruled by the conservative fixed point. In three dimensions this mechanism gives $z=3/2$, a value significantly closer to the experimental result $z\approx 1$ than the value $z\approx 2$ found in fully dissipative models, either at or off equilibrium. This result indicates that non-dissipative dynamical couplings are necessary to develop a theory of natural swarms fully consistent with experiments, Comment: 5 pages, 2 figure

Motivated by the collective behaviour of biological swarms, we study the critical dynamics of field theories with coupling between order parameter and conjugate momentum in the presence of dissipation. By performing a dynamical renormalization group calculation at one loop, we show that the violation of momentum conservation generates a crossover between a conservative yet IR-unstable fixed point, characterized by a dynamic critical exponent $z=d/2$, and a dissipative IR-stable fixed point with $z=2$. Interestingly, the two fixed points have different upper critical dimensions. The interplay between these two fixed points gives rise to a crossover in the critical dynamics of the system, characterized by a crossover exponent $\kappa=4/d$. Such crossover is regulated by a conservation length scale, $\mathcal R_0$, which is larger the smaller the dissipation: beyond $\mathcal R_0$ the dissipative fixed point dominates, while at shorter distances dynamics is ruled by the conservative fixed point and critical exponent, a behaviour which is all the more relevant in finite-size systems with weak dissipation. We run numerical simulations in three dimensions and find a crossover between the exponents $z=3/2$ and $z=2$ in the critical slowing down of the system, confirming the renormalization group results. From the biophysical point of view, our calculation indicates that in finite-size biological groups mode-coupling terms in the equation of motion can significantly change the dynamical critical exponents even in the presence of dissipation, a step towards reconciling theory with experiments in natural swarms. Moreover, our result provides the scale within which fully conservative Bose-Einstein condensation is a good approximation in systems with weak symmetry-breaking terms violating number conservation, as quantum magnets or photon gases., Comment: 33 pages, 7 figures

The correlated motion of flocks is an instance of global order emerging from local interactions. An essential difference with analogous ferromagnetic systems is that flocks are active: animals move relative to each other, dynamically rearranging their interaction network. The effect of this off-equilibrium element is well studied theoretically, but its impact on actual biological groups deserves more experimental attention. Here, we introduce a novel dynamical inference technique, based on the principle of maximum entropy, which accodomates network rearrangements and overcomes the problem of slow experimental sampling rates. We use this method to infer the strength and range of alignment forces from data of starling flocks. We find that local bird alignment happens on a much faster timescale than neighbour rearrangement. Accordingly, equilibrium inference, which assumes a fixed interaction network, gives results consistent with dynamical inference. We conclude that bird orientations are in a state of local quasi-equilibrium over the interaction length scale, providing firm ground for the applicability of statistical physics in certain active systems.

Three-dimensional tracking of animal systems is the key to the comprehension of collective behavior. Experimental data collected via a stereo camera system allow the reconstruction of the 3d trajectories of each individual in the group. Trajectories can then be used to compute some quantities of interest to better understand collective motion, such as velocities, distances between individuals and correlation functions. The reliability of the retrieved trajectories is strictly related to the accuracy of the 3d reconstruction. In this paper, we perform a careful analysis of the most significant errors affecting 3d reconstruction, showing how the accuracy depends on the camera system set-up and on the precision of the calibration parameters., Comment: 14 pages, 9 figures

We derive a new method to infer from data the out-of-equilibrium alignment dynamics of collectively moving animal groups, by considering the maximum entropy distribution consistent with temporal and spatial correlations of flight direction. When bird neighborhoods evolve rapidly, this dynamical inference correctly learns the parameters of the model, while a static one relying only on the spatial correlations fails. When neighbors change slowly and detailed balance is satisfied, we recover the static procedure. We demonstrate the validity of the method on simulated data. The approach is applicable to other systems of active matter.

Tracking multiple moving targets allows quantitative measure of the dynamic behavior in systems as diverse as animal groups in biology, turbulence in fluid dynamics and crowd and traffic control. In three dimensions, tracking several targets becomes increasingly hard since optical occlusions are very likely, i.e. two featureless targets frequently overlap for several frames. Occlusions are particularly frequent in biological groups such as bird flocks, fish schools, and insect swarms, a fact that has severely limited collective animal behavior field studies in the past. This paper presents a 3D tracking method that is robust in the case of severe occlusions. To ensure robustness, we adopt a global optimization approach that works on all objects and frames at once. To achieve practicality and scalability, we employ a divide and conquer formulation, thanks to which the computational complexity of the problem is reduced by orders of magnitude. We tested our algorithm with synthetic data, with experimental data of bird flocks and insect swarms and with public benchmark datasets, and show that our system yields high quality trajectories for hundreds of moving targets with severe overlap. The results obtained on very heterogeneous data show the potential applicability of our method to the most diverse experimental situations., Comment: 13 pages, 6 figures, 3 tables. Version 3 was slightly shortened, and new comprative results on the public datasets (thermal infrared videos of flying bats) by Z. Wu and coworkers (2014) were included. in A. Attanasi et al., "GReTA - A Novel Global and Recursive Tracking Algorithm in Three Dimensions", IEEE Trans. Pattern Anal. Mach. Intell., vol.37 (2015)

Flocks of starlings exhibit a remarkable ability to maintain cohesion as a group in highly uncertain environments and with limited, noisy information. Recent work demonstrated that individual starlings within large flocks respond to a fixed number of nearest neighbors, but until now it was not understood why this number is seven. We analyze robustness to uncertainty of consensus in empirical data from multiple starling flocks and show that the flock interaction networks with six or seven neighbors optimize the trade-off between group cohesion and individual effort. We can distinguish these numbers of neighbors from fewer or greater numbers using our systems-theoretic approach to measuring robustness of interaction networks as a function of the network structure, i.e., who is sensing whom. The metric quantifies the disagreement within the network due to disturbances and noise during consensus behavior and can be evaluated over a parameterized family of hypothesized sensing strategies (here the parameter is number of neighbors). We use this approach to further show that for the range of flocks studied the optimal number of neighbors does not depend on the number of birds within a flock; rather, it depends on the shape, notably the thickness, of the flock. The results suggest that robustness to uncertainty may have been a factor in the evolution of flocking for starlings. More generally, our results elucidate the role of the interaction network on uncertainty management in collective behavior, and motivate the application of our approach to other biological networks., Comment: 19 pages, 3 figures, 9 supporting figures

Bird flocking is a striking example of collective animal behaviour. A vivid illustration of this phenomenon is provided by the aerial display of vast flocks of starlings gathering at dusk over the roost and swirling with extraordinary spatial coherence. Both the evolutionary justification and the mechanistic laws of flocking are poorly understood, arguably because of a lack of data on large flocks. Here, we report a quantitative study of aerial display. We measured the individual three-dimensional positions in compact flocks of up to 2700 birds. We investigated the main features of the flock as a whole - shape, movement, density and structure - and discuss these as emergent attributes of the grouping phenomenon. We find that flocks are relatively thin, with variable sizes, but constant proportions. They tend to slide parallel to the ground and, during turns, their orientation changes with respect to the direction of motion. Individual birds keep a minimum distance from each other that is comparable to their wingspan. The density within the aggregations is non-homogeneous, as birds are packed more tightly at the border compared to the centre of the flock. These results constitute the first set of large-scale data on three-dimensional animal aggregations. Current models and theories of collective animal behaviour can now be tested against these results., Comment: To be published in Animal Behaviour

The study of collective animal behaviour must progress through a comparison between the theoretical predictions of numerical models and data coming from empirical observations. To this aim it is important to develop methods of three-dimensional (3D) analysis that are at the same time informative about the structure of the group and suitable to empirical data. In fact, empirical data are considerably noisier than numerical data, and they are subject to several constraints. We review here the tools of analysis used by the STARFLAG project to characterise the 3D structure of large flocks of starlings in the field. We show how to avoid the most common pitfalls i the quantitative analysis of 3D animal groups, with particular attention to the problem of the bias introduced by the border of the group. By means of practical examples, we demonstrate that neglecting border effects gives rise to artefacts when studying the 3D structure of a group. Moreover, we show that mathematical rigour is essential to distinguish important biological properties from trivial geometric features of animal groups., Comment: To be published in Animal Behaviour

The most startling examples of collective animal behaviour are provided by very large and cohesive groups moving in three dimensions. Paradigmatic examples are bird flocks, fish schools and insect swarms. However, because of the sheer technical difficulty of obtaining 3D data, empirical studies conducted to date have only considered loose groups of a few tens of animals. Moreover, these studies were very seldom conducted in the field. Recently the STARFLAG project achieved the 3D reconstruction of thousands of birds under field conditions, thus opening the way to a new generation of quantitative studies of collective animal behaviour. Here, we review the main technical problems in 3D data collection of large animal groups and we outline some of the methodological solutions adopted by the STARFLAG project. In particular, we explain how to solve the stereoscopic correspondence - or matching - problem, which was the major bottleneck of all 3D studies in the past., Comment: To be published in Animal Behaviour