March 18, 2020 7 be adaptive with respect to predation; group members may benefit from behaving more uniformly as predators 11 disproportionately target individuals whose behaviour differs from the majority [3] . Conversely, animals living 12 alone may benefit from behaving in a less predictable manner to confound predators; a mechanism known as 13 protean behaviour [4] . The contrasting selection pressures of group-and lone-living are therefore expected to 14 lead to adaptive differences in the extent of behavioural variability [5] . 15 Reduction in either within-or between-individual variability of behaviour contributes to greater homo- 16 geneity of animal collectives [6] . Previous studies have largely focussed on changes in behavioural variability 17 as an instantaneously reactive process, when animals behave in one way when alone, but immediately adjust 18 their behaviour to be less variable when in the presence of conspecifics [2] . These changes often arise through 19 simple local rules governing movement in response to neighbours [7] and can be interpreted as 'contextual 20 plasticity' [5] . Most research into the effect of predation on behavioural variability uses the framework of 21 contextual plasticity [8,9] , with individuals increasing their within-individual variability in the presence of 22 predator cues. However, a framework that is entirely based on context-driven plasticity of within-individual 23 variability makes it harder to clarify the evolution of changes in behavioural variability, because contextual 24 plasticity is thought to shield genotypes from selection [10] . 25 An alternative explanation for how changes in behavioural variability occur is that they are an intrin-26 sic process, where prior external conditions induce changes in behavioural variability which then become 27 intrinsic to the individual, persisting beyond the triggering conditions [11][12][13] . This implies that the extent 28 of behavioural variability is regulated by intrinsic mechanisms and forms part of an animal's phenotype [14] . 29 Indeed, very recent studies report evidence for a genetic component to between-individual differences in 30 within-individual variability [15] . While such differences in intrinsic behavioural variability are predicted by 31 theory to be subject to selection, empirical evidence for adaptive changes is largely lacking [5] .
32Here, we present robust evidence demonstrating that selective pressures have shaped intrinsic behavioural 33 variability in the desert locust Schistocerca gregaria. In this species, the same genotype can manifest in 34 two very distinct eco-phenotypes or phases, a lone-living 'solitarious' phase and a 'gregarious' phase that 35 forms large, dense groups [16] . Solitarious individuals show low levels of activity, have cryptic green or 36 beige colouration and avoid other locusts, whereas gregarious individuals are highly active, are attracted 37 to conspecifics, develop a striking aposematic yellow-and-black colouration, and can aggregate into swarms 38 containing...