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Question: Herbivores exert strong influences on vegetation through activities such as trampling, defoliation, and fertilization. The combined effect of these activities on plant performance may cause dramatic vegetation shifts. Because herbivore pressures and the relative importance of their different activities are not equally distributed across the landscape, it is important to understand their isolated effect. One example of an herbivore‐induced vegetation shift is the reindeer‐driven transition from a subarctic tundra vegetation dominated by dwarf shrubs into a more productive, graminoid‐dominated state. Here, we asked how each of the grazing activities by reindeer separately and combined shape vegetation composition.

Location: Nordreisa, Norway.

Methods: We used a field experiment over six summers to study the separate and interacting effects of reindeer trampling, defoliation, addition of faeces and removal of moss on tundra heath vegetation, and to identify which of these factors were most important in driving the plant community towards a graminoid‐dominated state.

Results: The combination of all treatments resulted in the strongest changes in vegetation, but trampling was the single most important factor altering the vegetation composition by reducing the abundance of both evergreen and deciduous dwarf shrubs. In contrast to what was expected, none of our treatments, separate or combined, resulted in an increased abundance of graminoids in 5 years, although such rapid vegetation changes have been observed in the field in similar environmental conditions.

Conclusions: Trampling is the key process by which reindeer influence the abundance of functional groups, but only many processes combined result in strong changes in community composition. Moreover, additional factors not included in this experiment, such as urine, may be important in causing a state shift to a graminoid‐dominated community.

Herbivory is one of the key drivers shaping plant community dynamics. Herbivores can strongly influence plant productivity directly through defoliation and the return of nutrients in the form of dung and urine, but also indirectly by reducing the abundance of neighbouring plants and inducing changes in soil processes. However, the relative importance of these processes is poorly understood. We, therefore, established a common garden experiment to study plant responses to defoliation, dung addition, moss cover, and the soil legacy of reindeer grazing. We used an arctic tundra grazed by reindeer as our study system, and Festuca ovina, a common grazing-tolerant grass species as the model species. The soil legacy of reindeer grazing had the strongest effect on plants, and resulted in higher growth in soils originating from previously heavily-grazed sites. Defoliation also had a strong effect and reduced shoot and root growth and nutrient uptake. Plants did not fully compensate for the tissue lost due to defoliation, even when nutrient availability was high. In contrast, defoliation enhanced plant nitrogen concentrations. Dung addition increased plant production, nitrogen concentrations and nutrient uptake, although the effect was fairly small. Mosses also had a positive effect on aboveground plant production as long as the plants were not defoliated. The presence of a thick moss layer reduced plant growth following defoliation. This study demonstrates that grasses, even though they suffer from defoliation, can tolerate high densities of herbivores when all aspects of herbivores on ecosystems are taken into account. Our results further show that the positive effect of herbivores on plant growth via changes in soil properties is essential for plants to cope with a high grazing pressure. The strong effect of the soil legacy of reindeer grazing reveals that herbivores can have long-lasting effects on plant productivity and ecosystem functioning after grazing has ceased.

1. Mammalian herbivores can strongly influence nitrogen (N) cycling and herbivore urine could be a central component of the N cycle in grazed ecosystems. Despite its potential role for ecosystem productivity and functioning, the fate of N derived from urine has rarely been investigated in grazed ecosystems. 2. This study explored the fate of N-15-enriched urea in tundra sites that have been either lightly or intensively grazed by reindeer for more than 50years. We followed the fate of the N-15 applied to the plant canopy, at 2weeks and 1year after tracer addition, in the different ecosystem N pools. 3. N-15-urea was rapidly incorporated in cryptogams and in above-ground parts of vascular plants, while the soil microbial pool and plant roots sequestered only a marginal proportion. Furthermore, the litter layer constituted a large sink for the N-15-urea, at least in the short term, indicating a high biological activity in the litter layer and high immobilization in the first phases of organic matter decomposition. 4. Mosses and lichens still constituted the largest sink for the N-15-urea 1year after tracer addition at both levels of grazing intensity demonstrating their large ability to capture and retain N from urine. Despite large fundamental differences in their traits, deciduous and evergreen shrubs were just as efficient as graminoids in taking up the N-15-urea. The total recovery of N-15-urea was lower in the intensively grazed sites, suggesting that reindeer reduce ecosystem N retention. 5. Synthesis. The rapid incorporation of the applied N-15-urea indicates that arctic plants can take advantage of a pulse of incoming N from urine. In addition, N-15 values of all taxa in the heavily grazed sites converged towards the N-15 values for urine, bringing further evidence that urine is an important N source for plants in grazed tundra ecosystems.

1. Herbivores play a key role in shaping ecosystem structure and functions by influencing plant and microbial community composition and nutrient cycling.

2. This study investigated the long-term effects of herbivores on plant resource acquisition. We explored differences in the natural delta N-15 signatures in plant, microbial and soil N pools, and examined mycorrhizal colonization in two tundra sites that have been either lightly or heavily grazed by reindeer for more than 50 years. The study examined changes in nutrient acquisition in five common tundra plants with contrasting traits and mycorrhiza status; the mycorrhizal dwarf shrubs, Betula nana, Vaccinium myrtillus and Empetrum hermaphroditum; a mycorrhizal grass, Deschampsia flexuosa, and a non-mycorrhizal sedge, Carex bigelowii.

3. There were large variations in delta N-15 among coexisting plant species in the lightly grazed sites. This variation was dramatically reduced in the heavily grazed sites. At an individual species level, delta N-15 was higher in E. hermaphroditum and lower in C. bigelowii in the heavily grazed sites. Mycorrhizal colonization in B. nana and E. hermaphroditum roots were also lower in the heavily grazed sites. The delta N-15 signatures of the total soil N pool and of the microbial N pools were higher in the heavily grazed sites.

4. Since the strong delta N-15 differentiation among plant species has been interpreted as a result of plants with different mycorrhizal types using different sources of soil nitrogen, we suggest that the lower variation in delta N-15 in heavily grazed sites indicates a lower niche differentiation in nitrogen uptake among plants. Reduced mycorrhizamediated nitrogen uptake by some of the species, a shift towards a more mineral nutrition due to higher nutrient turnover, and uptake of labile nitrogen from dung and urine in the heavily grazed sites could all contribute to the changes in plant delta N-15.

5. We conclude that herbivores have the potential to influence plant nutrient uptake and provide the first data suggesting that herbivores decrease nutrient partitioning on the basis of chemical N forms among plant species. Reduced niche complementarity among species is potentially important for estimates of the effects of -herbivory on plant nutrient availability and species coexistence.

Reindeer appear to have strong positive effects on plant productivity and nutrient cycling in strongly nutrient-limited ecosystems. While the direct effects of grazing on vegetation composition have been intensively studied, much less is known about the indirect effect of grazing on plant-soil interactions. This thesis investigated the indirect effects of ungulate grazing on arctic plant communities via soil nutrient availability and plant nutrient uptake.

At high density, the deposition of dung alone increased plant productivity both in nutrient rich and nutrient poor tundra habitats without causing major changes in soil possesses. Plant community responses to dung addition was slow, with a delay of at least some years. By contrast, a ¹⁵N-urea tracer study revealed that nutrients from reindeer urine could be rapidly incorporated into arctic plant tissues. Soil and microbial N pools only sequestered small proportions of the tracer. This thesis therefore suggests a strong effect of dung and urine on plant productivity by directly providing nutrient-rich resources, rather than by stimulating soil microbial activities, N mineralization and ultimately increasing soil nutrient availability. Further, defoliation alone did not induce compensatory growth, but resulted in plants with higher nutrient contents. This grazing-induced increase in plant quality could drive the high N cycling in arctic secondary grasslands by providing litter of a better quality to the belowground system and thus increase organic matter decomposition and enhance soil nutrient availability. Finally, a ¹⁵N natural abundance study revealed that intense reindeer grazing influences how plants are taking up their nutrients and thus decreased plant N partitioning among coexisting plant species.

Taken together these results demonstrate the central role of dung and urine and grazing-induced changes in plant quality for plant productivity. Soil nutrient concentrations alone do not reveal nutrient availability for plants since reindeer have a strong influence on how plants are taking up their nutrients. This thesis highlights that both direct and indirect effects of reindeer grazing are strong determinants of tundra ecosystem functioning. Therefore, their complex influence on the aboveground and belowground linkages should be integrated in future work on tundra ecosystem N dynamic.

Deposition of feces is a key mechanism by which herbivores influence soil nutrient cycling and plant production, but the knowledge about its importance for plant production and community structure is still rudimental since experimental evidence is scarce. We thus performed a 7-year long reindeer feces manipulation experiment in two tundra vegetation types with contrasting nutrient availability and analyzed effects on plant community composition and soil nutrient availability. Despite feces being fairly nutrient poor, feces manipulation had strong effect on both the nutrient-poor heath and the nutrient-rich meadow. The strongest effect was detected when feces were added at high density, with a substantial increase in total vascular plant productivity and graminoids in the two communities. Doubling natural deposition of reindeer feces enhanced primary production and the growth of deciduous shrubs in the heath. By contrast, removal of feces decreased only the production of graminoids and deciduous shrubs in the heath. Although the response to feces addition was faster in the nutrient-rich meadow, after 7 years it was more pronounced in the nutrient-poor heath. The effect of feces manipulation on soil nutrient availability was low and temporarily variable. Our study provides experimental evidence for a central role of herbivore feces in regulating primary production when herbivores are abundant enough. Deposition of feces alone does, however, not cause dramatic vegetation shifts; to drive unproductive heath to a productive grass dominated state, herbivore trampling, and grazing are probably also needed.

Herbivores can strongly influence plant growth directly through defoliation and the return of nutrients in the form of dung and urine but also indirectly by reducing the abundance of neighbouring plants and inducing changes in soil processes. The relative importance of these driving mechanisms of plant response to herbivory are still poorly understood. In a common garden experiment, we studied the aboveground and belowground responses of Festuca ovina, a grazing tolerant grass common in arctic secondary grassland, to defoliation, reindeer dung addition, changes in soil microclimate induced by the presence or the absence of a moss cover, and soil grazing legacy. Defoliation strongly reduced shoot and root growth and plant nutrient uptake. Plants did thus not compensate for the tissue lost due to defoliation, even at a higher nutrient availability. By contrast, defoliation enhanced plant N concentration and decreased plant C to N ratio. Soil from heavily grazed sites and dung addition increased plant production, plant N concentrations and nutrient uptake, although the effects of dung addition were only small. Mosses had a strong negative effect of root biomass and reduced plant compensatory growth after defoliation. Interestingly mosses also had facilitative effects on aboveground plant growth in absence of defoliation and on plant nutrient uptake and N concentrations. Although plants suffered severely from defoliation, they were also strongly favoured by the increased nutrient availability associated with herbivory. After two years, plants produced as much biomass when all positive and negative effects of herbivores were considered (defoliation, soil communities and nutrient availability under heavily grazing, dung addition and no moss cover) as in the ungrazed conditions (no defoliation, soil communities and nutrient availability under lightly grazing, no dung addition, a thick moss cover). This study indicates that graminoids can tolerate high densities of herbivores, although it suffer from defoliation directly, and suggests that changes in plant quality following defoliation and grazing-induced changes in soil processes are two key mechanisms through which herbivores can control plant productivity in arctic secondary grasslands. Plant tolerance to herbivory will depends on how herbivores utilise a pasture area and on the balance between the positive and the negative effects of grazing on plant growth.

Mammalian herbivores can strongly influence nitrogen cycling and herbivore urine could be an important component of the nutrient cycle in grazed ecosystems. Despite its potential role for ecosystem productivity and soil processes, the distribution of N from urine in the different ecosystem compartments is poorly understood. This study investigates the fate of ¹⁵N enriched urea applied above the plant canopy in two tundra sites either heavily or lightly grazed by reindeer for the last 50 years. We explored the fate of the ¹⁵N in the different ecosystem N pools at 2 weeks and 1 years following tracer addition. We hypothesized that cryptogams would take up most N under light grazing, but graminoids most N under heavy grazing. The ¹⁵N-urea was rapidly incorporated in cryptogams and aboveground parts of vascular plants, while the soil microbial pool and plant roots sequestered only a marginal proportion of the labelled N applied. Hence, urine addition supports a higher primary production in tundra since most of the nutrients released from urine could be assimilated by the aboveground components with little N reaching the belowground compartments. Mosses and lichens still constituted the largest sink of the ¹⁵N-urea 1 year after tracer addition at both levels of grazing intensity demonstrating their large ability to capture and retain N  from urine. Deciduous and evergreen shrubs were just as efficient as graminoids in taking up the ¹⁵N-urea. The total recovery of the labelled urea was lower in the heavily grazed sites, suggesting that reindeer reduce the N retention in the system. Rapid incorporation of the applied ¹⁵N-urea indicates that arctic plants can take advantage of a pulse of incoming N in the form of urea, which supports a higher primary production. However, whether urine also maintains a high production of forage plants depend on plant community composition, since most urea was recovered in non-forage plants for reindeer.