Seeing the locust in the swarm: accounting for spatiotemporal hierarchy improves ecological models of insect populations

Ecological phenomena operate at different spatial scales and are not uniform across landscapes or through time. One ecological theory that attempts to account for scaling and spatiotemporal variances is hierarchical patch dynamics. It introduces a hierarchical patch network with smaller spatiotemporal scales being nested within larger scales. However, few studies have modeled its presence within animal population dynamics. Locusts are an excellent model for investigating the spatiotemporal hierarchy of animal population dynamics, due to their high migratory capacity, large geographic ranges that extend across widely differing environments, and available long-term data on distributions. Here, we investigated the influence of preceding vegetation growth on desert locust Schistocerca gregaria and Australian plague locust Chortoicetes terminifera outbreaks on three spatial levels (species range > geographic region > land unit) and between seasons. Both species are dryland herbivores with population dynamics linked to habitat productivity pulses after rain. We used NDVI data (MODIS imagery) as a measure of vegetation growth in hierarchical generalized additive models at different scales. Locust outbreaks were either preceded by vegetation growth between 78 and 32 days (Australian plague locusts) or 32 and 20 days before (desert locust) the observation. Although prior vegetation growth characterized outbreaks of both species, the temporal pattern of NDVI differed between spatiotemporal levels. All model selection criteria selected for a similar spatial hierarchy for both species: geographic region > land unit which supports the hierarchical patch dynamics paradigm. Further, it illuminates important timing differences between geographic regions and land units for preceding vegetation growth and locust outbreaks which can help locust managers identify when and where outbreaks occur. By acknowledging the spatiotemporal patterning of locust abundance, we account for heterogeneity of population dynamics throughout species ranges. Our findings demonstrate the importance of incorporating spatiotemporal variation in population models of insects and other animals.