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Collective mechanical adaptation of honeybee swarms


Honeybee Apis mellifera swarms form large congested tree-hanging clusters made solely of bees attached to each other1. How these structures are maintained under the influence of dynamic mechanical forcing is unknown. To address this, we created pendant clusters and subject them to dynamic loads of varying orientation, amplitude, frequency and duration. We find that horizontally shaken clusters adapt by spreading out to form wider, flatter cones that recover their original shape when unloaded. Measuring the response of a cluster to an impulsive pendular excitation shows that flattened cones deform less and relax faster than the elongated ones (that is, they are more stable). Particle-based simulations of a passive assemblage suggest a behavioural hypothesis: individual bees respond to local variations in strain by moving up the strain gradient, which is qualitatively consistent with our observations of individual bee movement during dynamic loading. The simulations also suggest that vertical shaking will not lead to significant differential strains and thus no shape adaptation, which we confirmed experimentally. Together, our findings highlight how a super-organismal structure responds to dynamic loading by actively changing its morphology to improve the collective stability of the cluster at the expense of increasing the average mechanical burden of an individual.

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Fig. 1: A mechanically adaptive honeybee cluster.
Fig. 2: Quantifying adaptive response of the cluster to horizontal shaking.
Fig. 3: Computational model of mechanical adaptation.
Fig. 4: Response to vertical shaking.


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This work was supported by funding from the US NSF PoLS grant 1606895. We thank the Mahadevan laboratory for discussions and comments.

Author information




O.P., J.M.P. and L.M. conceived of the research study; O.P., J.M.P., M.K.S. and L.M. designed the experiments, O.P., J.M.P. and M.K.S. performed the experiments; O.P. analysed the data with the help of J.M.P; O.P. and L.M. conceived of the behavioural rule and designed the simulations; O.P. carried out the simulations; O.P., J.M.P. and L.M. wrote the paper; L.M. supervised the project.

Corresponding author

Correspondence to L. Mahadevan.

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The authors declare no competing interests.

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Supplementary information

Supplementary Information

Supplementary Information, Supplementary Figures 1–6, Supplementary Tables 1–3

Reporting Summary

Supplementary Video 1

Honeybee cluster in the wind

Supplementary Video 2

Time-lapse of horizontal shaking experiment

Supplementary Video 3

Before/after horizontal shaking experiment—response to continuous shaking

Supplementary Video 4

Before/after horizontal shaking experiment—response to a single sharp shake

Supplementary Video 5

Tracking individual bees during horizontal shaking experiment

Supplementary Video 6

Passive simulations to extract local strains

Supplementary Video 7

Active simulations

Supplementary Video 8

Active simulations

Supplementary Video 9

Honeybee cluster breakage

Supplementary Video 10

Before/after vertical shaking experiment—response to a single sharp shake

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Peleg, O., Peters, J.M., Salcedo, M.K. et al. Collective mechanical adaptation of honeybee swarms. Nature Phys 14, 1193–1198 (2018).

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