Herbivory prevents positive responses of lowland plants to warmer and more fertile conditions at high altitudes

Warm-adapted low elevation plants are expected to exhibit considerable range shifts to higher altitudes and latitudes as a result of climate warming and increased nutrient loads. However, empirical studies show that the magnitude and direction of plant responses are highly species- and site-specific, suggesting that several additional drivers interact with warmer climate. We experimentally tested the interactive effects of climate warming, mammalian herbivory and soil fertility on low elevation plants. Seedlings of three warm-adapted lowland forbs (Epilobium angustifolium, Silene dioica and Solidago virgaurea) were transplanted to an open tundra site with native mountain tundra vegetation, and the effects of full factorial combinations of herbivore exclosures, warming and fertilization on transplant survival, growth and flowering were studied for two growing seasons. We also investigated the response of native vegetation biomass to the same treatments and compared it with the responses of transplanted lowland forbs. Effects of both warming and fertilization on the transplanted lowland forbs strongly hinged on herbivore exclusion, resulting in 2-13-fold increase in biomass in warmed and fertilized plots without herbivores compared with warmed and fertilized plots with herbivores present, the magnitude depending on the species. While warm-adapted transplants benefited from warming, the native tundra plant community biomass did not respond to warming treatment. Our results show that grazing limits the growth of transplants under warmer and more productive conditions, indicating that the expansion of lowland plant species to higher altitudes with warming may be hampered by mammalian herbivory. Furthermore, our results also suggest that migration of warm-adapted species into lightly grazed high-altitude tundra ecosystems might transform these communities to be more responsive to warmer climate and nutrient loads. Studies that do not consider species' upward shifts from lower altitudes might thus have underestimated vegetation responses to global warming, as well as the potential of herbivory to influence these responses.