No abstract

Large herbivores can control plant community composition and, under certain conditions, even induce vegetation shifts to alternative ecosystem states. As different plant assemblages maintain contrasting carbon (C) cycling patterns, herbivores have the potential to alter C sequestration at regional scales. Their influence is of particular interest in the Arctic tundra, where a large share of the world's soil C reservoir is stored. We assessed the influence of grazing mammals on tundra vegetation and C stocks by resampling two sites located along pasture rotation fences in northern Norway. These fences have separated lightly grazed areas from heavily grazed areas (in close proximity to the fences) and moderately grazed areas (further away from the fences) for the past 50 years. Fourteen years earlier, the lightly and moderately grazed areas were dominated by dwarf shrubs, whereas heavy grazing had promoted the establishment of graminoid‐dominated vegetation. Since then, both reindeer densities and temperatures have increased, and more time has passed for transient dynamics to be expressed. We expected that the vegetation and C stocks would have changed under all grazing intensities, but not necessarily in the same way. At the site where relative reindeer numbers and trampling intensity had increased the most, graminoid‐dominated vegetation was now also found in the moderately grazed area. At the other site, the dominant vegetation types under all grazing intensities were the same as 14 years earlier. We show that the heavily grazed, graminoid‐dominated areas stored less C above‐ground than the lightly grazed, shrub‐dominated areas. Yet, the below‐ground consequences of grazing‐induced grassification varied between the sites: Grazing did not alter organic soil C stocks at the site where both evergreen and deciduous shrubs were abundant in the lightly grazed area, whereas heavy grazing increased organic soil C stocks at the site where the deciduous shrub Betula nana was dominant. Our results indicate that, despite the negative impacts of grazers on above‐ground C storage, their impact on below‐ground C may even be positive. We suggest that the site‐specific responses of organic soil C stocks to grazing could be explained by the differences in vegetation under light grazing. This would imply that the replacement of deciduous shrubs by graminoids, as a consequence of grazing could be beneficial for C sequestration in tundra soils. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13029/suppinfo is available for this article.

Selective herbivory of palatable plant species provides a competitive advantage for unpalatable plant species, which often have slow growth rates and produce slowly decomposable litter. We hypothesized that through a shift in the vegetation community from palatable, deciduous dwarf shrubs to unpalatable, evergreen dwarf shrubs, selective herbivory may counteract the increased shrub abundance that is otherwise found in tundra ecosystems, in turn interacting with the responses of ecosystem carbon (C) stocks and CO2 balance to climatic warming. We tested this hypothesis in a 19-year field experiment with factorial treatments of warming and simulated herbivory on the dominant deciduous dwarf shrub Vaccinium myrtillus. Warming was associated with a significantly increased vegetation abundance, with the strongest effect on deciduous dwarf shrubs, resulting in greater rates of both gross ecosystem production (GEP) and ecosystem respiration (ER) as well as increased C stocks. Simulated herbivory increased the abundance of evergreen dwarf shrubs, most importantly Empetrum nigrum ssp. hermaphroditum, which led to a recent shift in the dominant vegetation from deciduous to evergreen dwarf shrubs. Simulated herbivory caused no effect on GEP and ER or the total ecosystem C stocks, indicating that the vegetation shift counteracted the herbivore-induced C loss from the system. A larger proportion of the total ecosystem C stock was found aboveground, rather than belowground, in plots treated with simulated herbivory. We conclude that by providing a competitive advantage to unpalatable plant species with slow growth rates and long life spans, selective herbivory may promote aboveground C stocks in a warming tundra ecosystem and, through this mechanism, counteract C losses that result from plant biomass consumption.

Ungulate trampling modifies soils and interlinked ecosystem functions across biomes. Until today, most research has focused on temperate ecosystems and mineral soils while trampling effects on cold and organic matter‐rich tundra soils remain largely unknown. We aimed to develop a general model of trampling effects on soil structure, biota, microclimate and biogeochemical processes, with a particular focus on polar tundra soils. To reach this goal, we reviewed literature about the effects of trampling and physical disturbances on soils across biomes and used this to discuss the knowns and unknowns of trampling effects on tundra soils. We identified the following four pathways through which trampling affects soils: (a) soil compaction; (b) reductions in soil fauna and fungi; (c) rapid losses in vegetation biomass and cover; and (d) longer term shifts in vegetation community composition. We found that, in polar tundra, soil responses to trampling pathways 1 and 3 could be characterized by nonlinear dynamics and tundra‐specific context dependencies that we formulated into testable hypotheses. In conclusion, trampling may affect tundra soil significantly but many direct, interacting and cascading responses remain unknown. We call for research to advance the understanding of trampling effects on soils to support informed efforts to manage and predict the functioning of tundra systems under global changes. A free Plain Language Summary can be found within the Supporting Information of this article.

The circumpolar Arctic is currently facing multiple global changes that have the potential to alter the capacity of tundra soils to store carbon. Yet, predicting changes in soil carbon is hindered by the fact that multiple factors simultaneously control processes sustaining carbon storage and we do not understand how they act in concert. Here, we investigated the effects of warmer temperatures, enhanced soil nitrogen availability, and the combination of these on tundra carbon stocks at three different grazing regimes: on areas with over 50‐yr history of either light or heavy reindeer grazing and in 5‐yr‐old exlosures in the heavily grazed area. In line with earlier reports, warming generally decreased soil carbon stocks. However, our results suggest that the mechanisms by which warming decreases carbon storage depend on grazing intensity: under long‐term light grazing soil carbon losses were linked to higher shrub abundance and higher enzymatic activities, whereas under long‐term heavy grazing, carbon losses were linked to drier soils and higher enzymatic activities. Importantly, under enhanced soil nitrogen availability, warming did not induce soil carbon losses under either of the long‐term grazing regimes, whereas inside exclosures in the heavily grazed area, also the combination of warming and enhanced nutrient availability induced soil carbon loss. Grazing on its own did not influence the soil carbon stocks. These results reveal that accounting for the effect of warming or grazing alone is not sufficient to reliably predict future soil carbon storage in the tundra. Instead, the joint effects of multiple global changes need to be accounted for, with a special focus given to abrupt changes in grazing currently taking place in several parts of the Arctic.

The climate‐driven encroachment of shrubs into the Arctic is accompanied by shifts in soil fungal communities that could contribute to a net release of carbon from tundra soils. At the same time, arctic grazers are known to prevent the establishment of deciduous shrubs and, under certain conditions, promote the dominance of evergreen shrubs. As these different vegetation types associate with contrasting fungal communities, the belowground consequences of climate change could vary among grazing regimes. Yet, at present, the impact of grazing on soil fungal communities and their links to soil carbon have remained speculative. Here we tested how soil fungal community composition, diversity and function depend on tree vicinity and long‐term reindeer grazing regime and assessed how the fungal communities relate to organic soil carbon stocks in an alpine treeline ecotone in Northern Scandinavia. We determined soil carbon stocks and characterized soil fungal communities directly underneath and >3 m away from mountain birches (Betula pubescens ssp. czerepanovii) in two adjacent 55‐year‐old grazing regimes with or without summer grazing by reindeer (Rangifer tarandus). We show that the area exposed to year‐round grazing dominated by evergreen dwarf shrubs had higher soil C:N ratio, higher fungal abundance and lower fungal diversity compared with the area with only winter grazing and higher abundance of mountain birch. Although soil carbon stocks did not differ between the grazing regimes, stocks were positively associated with root‐associated ascomycetes, typical to the year‐round grazing regime, and negatively associated with free‐living saprotrophs, typical to the winter grazing regime. These findings suggest that when grazers promote dominance of evergreen dwarf shrubs, they induce shifts in soil fungal communities that increase soil carbon sequestration in the long term. Thus, to predict climate‐driven changes in soil carbon, grazer‐induced shifts in vegetation and soil fungal communities need to be accounted for.

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