Semi-arid shrub steppe occupies a vast geographic range that is characterized in part by distinct seasonal patterns in precipitation. Few studies have evaluated how variability in both the amount and timing of precipitation affect the structure and physiology of shrubs in these systems. We quantified changes in foliar crown parameters, xylem anatomy, gas exchange, and hydraulic transport capacity in deep-rooted Artemisia tridentata shrubs following 20 years of experimental manipulations in amount and seasonal timing of precipitation. We hypothesized that shrub growth (total leaf area per shrub and cover of shrub community), hydraulic transport efficiency, and gas exchange would increase in shrubs in irrigated plots compared to non-irrigated control plots, especially for irrigation applied in winter rather than summer. We also predicted similar changes in xylem anatomy (ring width, vessel size and frequency). Most treatment responses entailed changes in plant structure, and were generally consistent with our hypotheses: total-shrub leaf area, shrub basal area, canopy cover, and maximum sapwood-specific branch hydraulic conductivity were more than 2× greater in shrubs in winter-irrigated compared to control plots, while summer irrigation had few effects on these variables. Irrigation in either season did not affect xylem vessel size, but did increase xylem ring width by ~ 2 × and decreased xylem vessel frequency by about half. Anatomical, morphological, and stand-level abundance of A. tridentataappeared much more responsive to irrigation than state changes in gas exchange, particularly when the extra water is received during winter. Thus, it appears for sagebrush that seasonal timing is at least as important as the amount of precipitation, and that responses to changes in precipitation timing occur through changes in carbon allocation more so than changes in leaf-level carbon gain.
