Intermittent precipitation and spatial Allee effects drive irregular vegetation patterns in semiarid ecosystems

Abstract

Vegetation in semi-arid ecosystems frequently organizes into spatially heterogeneous mosaics that regulate ecosystem functioning, productivity, and resilience. These patterns arise from local biological interactions, including facilitation among neighboring plants and competition for limiting resources. Classical theoretical approaches have attributed such organization to scale-dependent feedbacks, predicting regular spatial patterns and abrupt transitions to collapse. However, growing empirical and theoretical evidence reveal that environmental variability and demographic stochasticity can fundamentally reshape spatial organization, driving irregular clusters, dynamic mosaics, and gradual rather than catastrophic vegetation declines. In drylands, rainfall variability is a dominant source of environmental forcing: precipitation typically occurs in short, irregular pulses that transiently enhance survival and recruitment before competitive interactions again dominate. Near persistence thresholds, ecosystem dynamics are therefore governed not only by average climatic conditions but also by the timing and spatial coincidence of favorable events. Under these conditions, positive density dependence and local facilitation can critically determine whether vegetation patches persist, expand, or collapse. Here, we develop an individual-based model that integrates intermittent precipitation with local Allee effects to examine how stochastic rainfall shapes spatial organization and persistence. We show that the interaction between pulsed resource availability and density-dependent survival generates irregular cluster structures and strongly modulates extinction risk, with resilience emerging from local spatial covariance and neighborhood density rather than from total biomass alone. These results highlight the importance of individual-level, stochastic processes in determining ecosystem resilience.