Leaf carbon isotope tracks the facilitation pattern of legume shrubs shaped by water availability and species replacement along a large elevation gradient in Trans-Himalayas.

BACKGROUND AND AIMS: Understanding patterns and mechanisms of nurse plant facilitation is important to predict the resilience of arid/semi-arid ecosystems to climate change. We investigate whether water availability and nurse species turnover interact to shape the facilitation pattern of widespread legume shrubs along a large elevation gradient. We also investigate whether leaf δ13C of nurse plants can track the facilitation pattern. METHODS: We measured the relative interaction index (RII) of the number of species within and outside the canopy of two widespread legume shrub species (Caragana gerardiana and Caragana versicolor) alternatively distributed along a large elevation gradient in the Trans-Himalayas. We also assessed the proportional increase of species richness (ISR) at the community level using the paired plot data. To determine site-specific water availability, we measured the leaf δ13C of nurse shrubs and calculated the Thornthwaite moisture index (MI) for each elevation site. KEY RESULTS: Elevational variations in RII, ISR and δ13C were mainly explained by the MI when the effects of soil nitrogen and plant traits (leaf nitrogen and shrub size) were controlled. Variations in RII and ISR across the two nurse species were explained better by δ13C than by smoothly changing climatic factors along elevation. At the transition zone between the upper limit of C. gerardiana (4100 m) and the lower limit of C. versicolor (4200 m), RII and ISR were much higher in C. versicolor than in C. gerardiana under a similar MI. Such an abrupt increase in facilitation induced by nurse species replacement was well tracked by the variation of δ13C. CONCLUSIONS: Water availability and nurse species replacement are crucial to shaping facilitation patterns by legume shrubs along a large elevation gradient in dry mountainous regions, such as the Trans-Himalayas. Turnover in nurse species under global change might significantly alter the pattern of nurse plant facilitation associated with water availability, which can be well tracked by leaf δ13C.

Shrub-mediated effects on soil nitrogen determines shrub-herbaceous interactions in drylands of the Tibetan Plateau.

Introduction: Shrub promotes the survival, growth and reproduction of understory species by buffering the environmental extremes and improving limited resources (i.e., facilitation effect) in arid and semiarid regions. However, the importance of soil water and nutrient availability on shrub facilitation, and its trend along a drought gradient have been relatively less addressed in water-limited systems. Methods: We investigated species richness, plant size, soil total nitrogen and dominant grass leaf δ13C within and outside the dominant leguminous cushion-like shrub Caragana versicolor along a water deficit gradient in drylands of Tibetan Plateau. Results: We found that C. versicolor increased grass species richness but had a negative effect on annual and perennial forbs. Along the water deficit gradient, plant interaction assessed by species richness (RIIspecies) showed a unimodal pattern with shift from increase to decrease, while plant interaction assessed by plant size (RIIsize) did not vary significantly. The effect of C. versicolor on soil nitrogen, rather than water availability, determined its overall effect on understory species richness. Neither the effect of C. versicolor on soil nitrogen nor water availability affected plant size. Discussion: Our study suggests that the drying tendency in association with the recent warming trends observed in drylands of Tibetan Plateau, will likely hinder the facilitation effect of nurse leguminous shrub on understories if moisture availability crosses a critical minimum threshold.