Experimental evidence of the long-term effects of reindeer on Arctic vegetation greenness and species richness at a larger landscape scale

Large herbivores influence plant community structure and ecosystem processes in many ecosystems. In large parts of the Arctic, reindeer (or caribou) are the only large herbivores present. Recent studies show that reindeer have the potential to mitigate recent warming-induced shrub encroachment in the Arctic and the associated greening of high-latitude ecosystems. This will potentially have large scale consequences for ecosystem productivity and carbon cycling. To date, information on variation in the interactions between reindeer and plants across Arctic landscapes has been scarce. We utilized a network of experimental sites across a latitudinal gradient in the Scandinavian mountains where reindeer have been excluded from 59 study plots for at least 15 years. We used this study system to test the effect of long-term exclusion of reindeer on the abundance of major plant functional groups, the greenness indexes Leaf Area Index (LAI) and Normalized Difference Vegetation Index (NDVI), soil mineral nitrogen (N) and phosphorous (P), and species richness, and to determine whether the effect of reindeer exclusion is dependent on reindeer density, productivity, soil fertility or climate. We found that NDVI and LAI, lichen and deciduous shrub abundances were largely reduced while soil mineral N was enhanced by reindeer. The direction and amplitude of other plant functional group responses to reindeer exclusion differed between forest and tundra as well as shrub- and herbaceous-dominated vegetation. Higher reindeer densities were related to decreased plant species richness in low-productive sites and to increased species richness in productive sites. The relative reduction in LAI and associated absolute reductions of deciduous shrubs in response to reindeer were positively related to reindeer density, while the relative reduction in NDVI was not. Further, relative reductions in LAI and NDVI in response to reindeer were unrelated to climate and soil fertility. Synthesis. Our results provide long-term experimental evidence highlighting the role of reindeer density in regulating plant species richness, global climate change induced greenness patterns and shrub encroachment at regional scales in the Arctic. These findings emphasize the need to consider reindeer in models predicting vegetation patterns and changes in high-latitude ecosystems.