Effects of Trampling on Morphological and Mechanical Traits of Dryland Shrub Species Do Not Depend on Water Availability

In semiarid drylands water shortage and trampling by large herbivores are two factors limiting plant growth and distribution. Trampling can strongly affect plant performance, but little is known about responses of morphological and mechanical traits of woody plants to trampling and their possible interaction with water availability. Seedlings of four shrubs (Caragana intermedia, Cynanchum komarovi, Hedysarum laeve and Hippophae rhamnoides) common in the semiarid Mu Us Sandland were grown at 4% and 10% soil water content and exposed to either simulated trampling or not. Growth, morphological and mechanical traits were measured. Trampling decreased vertical height and increased basal diameter and stem resistance to bending and rupture (as indicated by the increased minimum bend and break force) in all species. Increasing water availability increased biomass, stem length, basal diameter, leaf thickness and rigidity of stems in all species except C. komarovii. However, there were no interactive effects of trampling and water content on any of these traits among species except for minimum bend force and the ratio between stem resistance to rupture and bending. Overall shrub species have a high degree of trampling resistance by morphological and mechanical modifications, and the effects of trampling do not depend on water availability. However, the increasing water availability can also affect trade-off between stem strength and flexibility caused by trampling, which differs among species. Water plays an important role not only in growth but also in trampling adaptation in drylands.