Stress-gradient hypothesis explains susceptibility to Bromus tectorum invasion and community stability in North America's semi-arid Artemisia tridentata wyomingensis ecosystems
Questions: (1) What combinations of overlapping water and heat stress and herbivory disturbance gradients are associated with shifts in interactions between Artemisia tridentata subsp. wyomingensis (Artemisia) and herbaceous beneficiary species? (2) Do interactions between Artemisia and beneficiaries shift from competition to facilitation with increasing stress-disturbance where facilitation and competition are most frequent and strongest at the highest and lowest levels, respectively? (3) Do such relationships differ for native and non-native beneficiaries? (4) What are the implications of any observed shifts in interactions between community compositional stability in space and susceptibility to invasion? Location: North American Artemisia communities. Methods: We tested the stress-gradient hypothesis (SGH) in an observational study consisting of 75 sites located along overlapping water and heat stress and disturbance gradients. We used spatial patterns of association among Artemisia and six native and two non-native beneficiary species; including the invasive annual grass Bromus tectorum, representing a diverse array of life history strategies, to infer whether the net outcome of interactions was facilitation or competition. We assessed implications for community stability by examining shifts in community composition in space and resistance to invasion. Results/Conclusions: Cattle herbivory, a novel disturbance and selective force, was a significant component of two overlapping stress gradients most strongly associated with observed shifts in interactions. Facilitation and competition were strongest and most frequent at the highest and lowest stress levels along both gradients, respectively. Contrasting ecological optima among native and non-native beneficiaries led to strikingly different patterns of interactions. The four native bunchgrasses with the strongest competitive response abilities exhibited the strongest facilitation at their upper limits of stress tolerance, while the two non-natives exhibited the strongest competition at the highest stress levels, which coincided with their maximum abundance. Artemisia facilitation enhanced stability at intermediate stress levels by providing a refuge for native bunchgrasses, which in turn reduced the magnitude of B. tectorum invasion. However, facilitation was a destabilizing force at the highest stress levels when native bunchgrasses became obligate beneficiaries dependent on facilitation for their persistence. B. tectorum dominated these communities, and the next fire may convert them to annual grasslands.