Confronting the cycle synchronisation paradigm of defoliator outbreaks in space and time—Evidence from two systems in a mixed-species forest landscape

Abstract

Defoliators cause extensive damage in boreal and temperate forests of the world. Considerable effort has been invested to understand their individual population dynamics, and despite ample theorising, there is little empirical evidence on factors causing spatial synchrony of pest eruptions at landscape scales. We report on the landscape-level effect of forest configuration and composition on the intensity of outbreaks of spruce budworm and forest tent caterpillar in a mixedwood boreal forest in northern Minnesota (USA) and adjacent Ontario (Canada), and how this is related to the degree of spatial synchrony in each species' outbreak cycling. Using a large spatiotemporal tree-ring reconstruction of outbreak impacts across these two systems, we evaluate two contrasting theories governing defoliator outbreaks: harmonic oscillation (a.k.a. ‘clockwork’) and relaxation oscillation (a.k.a. ‘catastrophe’), each with consequences linked to top-down versus bottom-up influences on outbreak behaviour in time and space. We find synchrony varies temporally, among outbreak cycles and in direct proportion to cycle peak intensity; however, cycle peak intensities are distributed bimodally in time, and so, therefore, are synchrony coefficients. Spatially, the area where each pest species currently cycles with the greatest peak intensity and synchrony is where their preferred host trees are currently found in greatest proportion. Despite overall synchrony in cycling, we found, in both systems, a persistent negative spatial correlation among successive eruptive pulses of defoliation. Many of these eruptions failed to spread spatially and to coalesce with other spot eruptions to form extensive area-wide outbreaks. Eruptions often fail to spread at the hardwood-conifer interface, resulting in outbreak pulses that systematically bounce back and forth between landscape types, particularly when systems were cycling at low amplitude. These over-dispersed spatial patterns of pulse impact are consistent with a contagious theory of eruption and outbreak spread. They could be considered consistent with harmonic oscillation theory only for populations cycling at different frequencies, with cycling frequency determined by host forest landscape structure. Synthesis. We find that defoliator outbreak dynamics across systems include spatiotemporal signatures of each theoretical paradigm—suggesting a hybrid approach will better characterise outbreak behaviour. Host concentration influences which paradigm dominates the spatial dynamics in any given forest landscape context. Because of the synchronising effect of host concentration on forest insect spatial dynamics, mixedwoods appear to be less prone to intense, synchronised defoliator attack than forests of pure hardwood or pure conifer.