Collective behaviour is typically thought to arise from individuals following fixed interaction rules. The possibility that interaction rules may change under different circumstances has thus only rarely been investigated. Here we show that local interactions in flocks of wild jackdaws (Corvus monedula) vary drastically in different contexts, leading to distinct group-level properties. Jackdaws interact with a fixed number of neighbours (topological interactions) when traveling to roosts, but coordinate with neighbours based on spatial distance (metric interactions) during collective anti-predator mobbing events. Consequently, mobbing flocks exhibit a dramatic transition from disordered aggregations to ordered motion as group density increases, unlike transit flocks where order is independent of density. The relationship between group density and group order during this transition agrees well with a generic self-propelled particle model. Our results demonstrate plasticity in local interaction rules and have implications for both natural and artificial collective systems.

Supplementary Fig. 1 | Flock morphology and evidence of pairing for flocks #2 to #6. a1-a5, Spatial distributions and velocities of birds in three-dimensional space. Paired birds are colored in red. b1-b5, Radial distribution functions G(r). c1-c5, Joint PDFs of D i, n=1 and D i, n=2 . Supplementary Fig. 2 | Pairing causes variations in local interaction for flocks #2 to #6. a1-a5, Change of distance between a bird and its nearest neighbour at time 0. Flock #6 shows a somewhat different trend from the other flocks for unpaired birds, which may be due to the relatively large value of in this flock. b1-b5, Acceleration in the direction away from the nearest neighbour; positive values are repulsive and negative values are attractive. Flock #5 shows a somewhat different trend from the other flocks for paired birds, which may be due to a tendency in this flock for paired birds to be located in front of or in back of each other. c1-c5, Alignment angle between a focal bird and its n th neighbour. Error bars show the standard error and are smaller than the symbols in most figures. Supplementary Fig. 3 | Wingbeat frequency as a function of flight speed during cruising flight for flocks #2 to #6. Paired birds typically have lower wingbeat frequency than unpaired birds at same flight speed. Error bars show the standard error. The magnitudes of |u| represent ground speeds.

Digital holographic microscopy is used for characterizing the profiles of mean velocity, viscous and Reynolds shear stresses, as well as turbulence level in the inner part of turbulent boundary layers over several super-hydrophobic surfaces (SHSs) with varying roughness/texture characteristics. The friction Reynolds numbers vary from 693 to 4496, and the normalized root mean square values of roughness $(k_{rms}^{+})$ vary from 0.43 to 3.28. The wall shear stress is estimated from the sum of the viscous and Reynolds shear stress at the top of roughness elements and the slip velocity is obtained from the mean profile at the same elevation. For flow over SHSs with $k_{rms}^{+}<1$, drag reduction and an upward shift of the mean velocity profile occur, along with a mild increase in turbulence in the inner part of the boundary layer. As the roughness increases above $k_{rms}^{+}\sim 1$, the flow over the SHSs transitions from drag reduction, where the viscous stress dominates, to drag increase where the Reynolds shear stress becomes the primary contributor. For the present maximum value of $k_{rms}^{+}=3.28$, the inner region exhibits the characteristics of a rough wall boundary layer, including elevated wall friction and turbulence as well as a downward shift in the mean velocity profile. Increasing the pressure in the test facility to a level that compresses the air layer on the SHSs and exposes the protruding roughness elements reduces the extent of drag reduction. Aligning the roughness elements in the streamwise direction increases the drag reduction. For SHSs where the roughness effect is not dominant ($k_{rms}^{+}<1$), the present measurements confirm previous theoretical predictions of the relationships between drag reduction and slip velocity, allowing for both spanwise and streamwise slip contributions.

As one of nature's most striking examples of collective behaviour, bird flocks have attracted extensive research. However, we still lack an understanding of the attractive and repulsive forces that govern interactions between individuals within flocks and how these forces influence neighbours' relative positions and ultimately determine the shape of flocks. We address these issues by analysing the three-dimensional movements of wild jackdaws ( Corvus monedula ) in flocks containing 2–338 individuals. We quantify the social interaction forces in large, airborne flocks and find that these forces are highly anisotropic. The long-range attraction in the direction perpendicular to the movement direction is stronger than that along it, and the short-range repulsion is generated mainly by turning rather than changing speed. We explain this phenomenon by considering wingbeat frequency and the change in kinetic and gravitational potential energy during flight, and find that changing the direction of movement is less energetically costly than adjusting speed for birds. Furthermore, our data show that collision avoidance by turning can alter local neighbour distributions and ultimately change the group shape. Our results illustrate the macroscopic consequences of anisotropic interaction forces in bird flocks, and help to draw links between group structure, local interactions and the biophysics of animal locomotion.