Environmental conditions can control the structure and composition of plant communities by changing the direction and intensity of plant–plant interactions. In extreme arid regions, accompanied by water and soil nutrient limitation, positive shrub–herb interactions may vary along an aridity gradient, leading to changes in the ecological consequences of shrub encroachment. We investigated the vegetation and soil within 60 shrub patches and their paired interspaces at 20 sites from the northeast to southwest desert steppe of Inner Mongolia, China, encroached by the Caragana microphylla shrub. The results show that aridity, soil organic matter (SOM), and soil total phosphorus (TP) were the main factors driving shrub–herb interactions. The positive shrub–herb interaction first increased and then decreased with increasing aridity (in the range of De Martonne Index (DMI) 0.54 to 1.85). The DMI indirectly affected shrub–herb interaction through TP, and the facilitation of shrubs on herbs coverage and biomass increased with the increase in TP. The SOM can directly affect the shrub–herbs interaction, and the facilitation of shrubs on herb diversity decreases with the increase in SOM. Our results indicate that the shrub–herb interaction changes along the environmental stress gradient; in general, shrubs have a positive effect on herbaceous communities along the aridity gradient. This study underscores the positive effects of shrubs on vegetation restoration in desert steppes, and changing environmental conditions by increasing precipitation, increasing TP content, and reducing SOM content can enhance the facilitation of shrub on herbs to accelerate the ecological restoration of degraded desert steppe.

Leymus is a perennial genus that belongs to the tribe Triticeae (Poaceae) which has an adaptive capacity to ecological conditions and strong resistance to cold, drought, and salinity. Most Leymus species are fine herbs that can be used for agriculture, conservation, and landscaping. Due to confusion taxonomy within genera, the complete chloroplast (cp) genome of 13 Leymus species was sequenced, assembled, and compared with those of three other previously published Leymus species (Leymus condensatus, Leymus angustus, and Leymus mollis) to clarify the issue. Overall, the whole cp genome size ranged between 135,057 (L. condensatus) and 136,906 bp (Leymus coreanus) and showed a typical quadripartite structure. All studied species had 129 genes, including 83 protein-coding genes, 38 transfer RNAs, and 8 ribosomal RNAs. In total, 800 tandem repeats and 707 SSR loci were detected, most of which were distributed in the large single-copy region, followed by the inverted repeat (IR) and small single-copy regions. The sequence identity of all sequences was highly similar, especially concerning the protein-coding and IR regions; in particular, the protein-coding regions were significantly similar to those in the IR regions, regardless of small sequence differences in the whole cp genome. Moreover, the coding regions were more conserved than the non-coding regions. Comparisons of the IR boundaries showed that IR contraction and expansion events were reflected in different locations of rpl22, rps19, ndhH, and psbA genes. The close phylogenetic relationship of Leymus and Psathyrostachys indicated that Psathyrostachys possibly is the donor of the Ns genome sequence identified in Leymus. Altogether, the complete cp genome sequence of Leymus will lay a solid foundation for future population genetics and phylogeography studies, as well as for the analysis of the evolution of economically valuable plants.