Competition and facilitation: Contrasting effects of Artemisia Tridentata on desert vs. montane pines

Circumstantial evidence suggests that Artemisia tridentata may out-compete Pinus ponderosa and P. jefferyi for water at ecotones between shrub steppe and montane forest vegetation in the Great Basin. Other studies indicate that within the shrub steppe Artemisia may act as a nurse plant for a third species of pine, P. monophylla. We used field experiments to study these contrasting effects of Artemisia on P. ponderosa and P. monophylla within the context of the distributional patterns in western Nevada of all three species on andesite, and on sites where hydrothermal activity has altered the andesite. At intermediate elevations in the Great Basin Artemisia and P. monophylla are restricted to unaltered desert soils, whereas P. ponderosa is restricted to acidic, nutrient-poor altered andesite. Although mature P. monophylla were virtually absent in our study plots on altered andesite, first- and second-year seedlings were common. On adjacent unaltered andesite, all size classes of P. monophylla occurred, and P. monophylla seedlings were associated with Artemisia shrubs. Pinus ponderosa and P. jefferyi adults and seedlings were rare on unaltered andesite, but a wide range of size classes was found on altered andesite. In experiments, all P. ponderosa seedlings on unaltered andesite were consumed by predators regardless of positive or negative spatial association with shrubs. Of the P. monophylla seedlings that germinated on unaltered andesite, all that were under shrubs survived, but only 6% of those that germinated in the intershrub spaces survived. On the open altered andesite the mortality of P. monophylla seedlings due to abiotic stress was high, with a final survival of only 3%, whereas 28% of P. ponderosa seedlings survived the first growing season on altered andesite. On unaltered andesite, survival and conductance of P. ponderosa saplings was enhanced by shrub removal, but P. monophylla survival was significantly higher under shrubs than in shrub-removal plots or in intershrub spaces. In Artemisia-removal experiments, we found that Artemisia competed with P. ponderosa seedlings and saplings for water. Removal of Artemisia decreased water use efficiency (WUE) of P. monophylla seedlings. The absence of Artemisia may restrict Pinus monophylla from outcrops of altered andesite in the Great Basin, but provide refuges for P. ponderosa.