Abstract

Grassland biodiversity is vulnerable to land use change. How to best manage semi-natural grasslands for maintaining biodiversity is still unclear in many cases because land-use processes may depend on environmental conditions and the indirect effects of land-use on biodiversity mediated by altered abiotic and biotic factors are rarely considered. Here we evaluate the relative importance of the direct and indirect effects of grazing intensity on plant communities along an elevational gradient on a large topographic scale in the Eastern Carpathians in Ukraine. We sampled for two years 31 semi-natural grasslands exposed to cattle grazing. Within each grassland site we measured plant community properties such as the number of species, functional groups, and the proportion of species undesirable for grazing. In addition, we recorded cattle density (as a proxy for grazing intensity), soil properties (bare soil exposure, soil organic carbon, and soil pH) and densities of soil decomposers (earthworms and soil microorganisms). We used structural equation modelling to explore the direct and indirect effects of grazing intensity on plant communities along the elevation gradient. We found that cattle density decreased plant species and functional diversity but increased the proportion of undesirable species. Some of these effects were directly linked to grazing intensity (i.e., species richness), while others (i.e., functional diversity and proportion of undesirable species) were mediated via bare soil exposure. Although grazing intensity decreased with elevation, the effects of grazing on the plant community did not change along the elevation gradient. Generally, elevation had a strong positive direct effect on plant species richness as well as a negative indirect effect, mediated via altered soil acidity and decreased decomposer density. Our results indicate that plant diversity and composition are controlled by the complex interplay among grazing intensity and changing environmental conditions along an elevation gradient. Furthermore, we found lower soil pH, organic carbon and decomposer density with elevation, indicating that the effects of grazing on soil and related ecosystem functions and services in semi-natural grasslands may be more pronounced with elevation. This demonstrates that we need to account for environmental gradients when attempting to generalize effects of land-use intensity on biodiversity.

Highlights

  • Grasslands cover more than 40% of the global terrestrial area [1] and are among the most species-rich habitats in Europe [2]

  • Our results indicate that plant diversity and composition are controlled by the complex interplay among grazing intensity and changing environmental conditions along an elevation gradient

  • We evaluate the simultaneous direct and indirect effects of grazing intensity and elevation on plant species richness, number of key plant functional groups (legumes, grasses (Poaceae), other monocots, and forbs), and on the proportion of species undesirable for grazing in semi-natural grasslands at large topographic scale ranging from the Carpathian Mountains in Ukraine, across the adjacent foothills to the plain areas (Fig 2), which is a largely under-represented region in scientific literature

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Summary

Introduction

Grasslands cover more than 40% of the global terrestrial area [1] and are among the most species-rich habitats in Europe [2]. Grassland biodiversity compared to the diversity in other ecosystem types is among the most vulnerable to human impact, to land-use change [21]. In order to survive and to function, semi-natural grassland communities require regular grass removal, for example through grazing [2]. The effects of grazing on grassland biodiversity have been found to depend on the land-use intensity [2, 22] and on particular environmental conditions [23], yielding contrasting results. The inconsistency in the observed patterns may depend upon the balance of different mechanisms underlying the relationships among grazing and plant community composition and diversity along environmental gradients (the hypothesized mechanisms derived from literature are summarized in S1 Table in S1 Appendix and Fig 1)

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