Plant antagonistic facilitation across environmental gradients: a soil-resource ecosystem engineering model.

Theory questions the persistence of nonreciprocal interactions in which one plant has a positive net effect on a neighbor that, in return, has a negative net impact on its benefactor - a phenomenon known as antagonistic facilitation. We develop a spatially explicit consumer-resource model for belowground plant competition between ecosystem engineers, plants able to mine resources and make them available for any other plant in the community, and exploiters. We use the model to determine in what environmental conditions antagonistic facilitation via soil-resource engineering emerges as an optimal strategy. Antagonistic facilitation emerges in stressful environments where ecosystem engineers' self-benefits from mining resources outweigh the competition with opportunistic neighbors. Among all potential causes of stress considered in the model, the key environmental parameter driving changes in the interaction between plants is the proportion of the resource that becomes readily available for plant consumption in the absence of any mining activity. Our results align with theories of primary succession and the stress gradient hypothesis. However, we find that the total root biomass and its spatial allocation through the root system, often used to measure the sign of the interaction between plants, do not predict facilitation reliably.