Impoverished soil supports more plateau pika through lowered diversity of plant functional traits in Tibetan alpine meadows

Abstract

Frequent outbreaks of plateau pika (Ochotona curzoniae) have become a major challenge for the sustainability of Tibetan rangeland, although top-down control has been widely employed since the 1960s over the plateau. Based on studies on plant trait-mediated soil-plant-herbivore relations, we hypothesized that impoverished soil was a consequence of rangeland degradation as well as a cause of increasing pika numbers through lowered functional trait diversity (FD) of the plant community that provides advantages for pika fitness. At three sites of alpine meadows on the eastern Tibetan Plateau, we counted active burrows of pika in 27 plots (i.e., 9 plots per site) distributed along a gradient of soil fertility. For every plant species within each plot, we measured five functional traits (e.g., mature height, SLA: specific leaf area, LDMC: leaf dry matter content, and leaf N and P: leaf nitrogen and phosphorus concentrations), indicating plant performance in response to changing soil fertility and impacting the fitness of pika. We employed Bayesian regression, structural equation modeling and network analyses to assess causal relations from soil fertility to FD and burrow density. The results show i) there was no significant difference in burrow density between three sites; spatial variation among plots in terms of soil fertility, plant species diversity and FD accounted for 57%–72%, 26%–41% and 59%–73%, respectively, of the variation in burrow density; ii) the burrow density was associated positively with soil moisture and organic matter, plant coverage, forb abundance and FD of leaf P, but negatively with soil nutrients, plant species diversity and FD for SLA, height and leaf N; and iii) deficiency of soil P potentially resulted in an increase in pika density, likely through lowered FD of SLA and height, with other soil factors and FD of leaf nutrients indirectly associated with burrow density. Our study suggested that under the joint effect of intensive grazing by livestock and output of livestock products from region, deficiency of soil P potentially increased pika density, likely by supporting more and diverse food resources while decreasing the risk of being preyed upon. Control of pika outbreak should pay more attention to bottom-up restoration of degraded soil and vegetation via traditional rotational grazing and promotion of recycling and compensation of lost soil P and plant trait diversity.