Interactive Effects Between Reindeer and Habitat Fertility Drive Soil Nutrient Availabilities in Arctic Tundra

Herbivores impact nutrient availability and cycling, and the net effect of herbivory on soil nutrients is generally assumed to be positive in nutrient-rich environments and negative in nutrient-poor ones. This is, however, far from a uniform pattern, and there is a recognized need to investigate any interactive effects of herbivory and habitat fertility (i.e., plant C/N ratios) on soil nutrient availabilities. We determined long-term effects of reindeer on soil extractable nitrogen (N) and phosphorus (P) and their net mineralization rates along a fertility gradient of plant carbon (C) to N and P ratios in arctic tundra. Our results showed that reindeer had a positive effect on soil N in the more nutrient-poor sites and a negative effect on soil P in the more nutrient-rich sites, which contrasts from the general consensus. The increase in N availability was linked to a decrease in plant and litter C/N ratios, suggesting that a shift in vegetation composition toward more graminoids favors higher N cycling. Soil P availability was not as closely linked to the vegetation and is likely regulated more by herbivore-induced changes in soil physical and chemical properties. The changes in soil extractable N and P resulted in higher soil N/P ratios, suggesting that reindeer could drive the vegetation toward P-limitation. This research highlights the importance of including both the elements N and P and conducting studies along environmental gradients in order to better understand the interactive effects of herbivory and habitat fertility on nutrient cycling and primary production.     

Effects of summer grazing by reindeer on composition of vegetation, productivity and nitrogen cycling

In this study, we investigated the effect of reindeer grazing on tundra heath vegetation in northern Norway. Fences, erected 30 yr ago, allowed us to compare winter grazed, lightly summer grazed and heavily summer grazed vegetation at four different sites. At two sites, graminoids dominated the heavily grazed zone completely, while ericoid dwarf shrubs had almost disappeared. In the other two areas, the increase of graminoids was almost significant. At one of the sites where graminoids dominated the heavily grazed area, we also measured plant biomass, primary production and nitrogen cycling. In this site, heavy grazing increased primary production and rate of nitrogen cycling, while moderate grazing decreased primary production. These results were inconsistent with the view that the highest productivity is found at intermediate grazing pressure. These results rather support the hypothesis that intensive grazing can promote a transition of moss-rich heath tundra into productive, graminoid-dominated steppe-like tundra vegetation. Moreover the results suggests that intermittent intensive reindeer grazing can enhance productivity of summer ranges.     

Herbivores inhibit climate-driven shrub expansion on the tundra

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana , has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (>0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (<0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.     

Non-parallel changes in soil microbial carbon and nitrogen dynamics due to reindeer grazing in northern boreal forests

Reindeer grazing has a considerable influence on mineralization processes in northern Fennoscandian boreal forests, but the mechanisms underlying the observed differences between grazed and ungrazed areas are not well understood. We studied the below-ground impacts of reindeer grazing by comparing the carbon and nitrogen mineralization rates inside and outside long-term fenced reindeer exclosure areas in five oligotrophic, lichen-dominated and five mesotrophic, dwarf-shrub dominated forests. The soil C mineralization rates and microbial metabolic activity (qCO₂) were significantly lower in the grazed than the ungrazed areas in both oligotrophic and mesotrophic forests. The reductions occurred irrespective of the impact on soil moisture. We conclude that reindeer grazing causes a reduction in the supply of labile C substrates to microbes, resulting in reduced organic matter decomposition rates through changes in the activity of the microbial biomass. Simultaneously, grazing had no consistent effect on the microbial N dynamics, but the impact ranged from no change to increased or decreased in N mineralization rates at the different study sites. The impact of grazing on the N mineralization potential thus seems to be site-specific and uncoupled from the impact of grazing on soil C mineralization. Reciprocal transplant incubations showed no interactions between N mineralization rates and the reindeer-mediated impact on the soil microclimate. We suggest that plant root damage due to trampling by reindeer may be an important mechanism for the deceleration of soil C cycling. In some cases, however, the impact of grazing on the soil active N pool may be strong enough to outweigh the reduction in soil organic matter decomposition, and by these means uncouple soil N dynamics from soil C quality.     

Direct and indirect control by snow cover over decomposition in alpine tundra along a snowmelt gradient

We assessed direct and indirect effects of snow cover on litter decomposition and litter nitrogen release in alpine tundra. Direct effects are driven by the direct influence of snow cover on edaphoclimatic conditions, whereas indirect effects result from the filtering effect of snow cover on species’ abundance and traits. We compared the in situ decomposition of leaf litter from four dominant plant species (two graminoids, two shrubs) at early and late snowmelt locations using a two-year litter-bag experiment. A seasonal experiment was also performed to estimate the relative importance of winter and summer decomposition. We found that growth form (graminoids vs. shrubs) are the main determinants of decomposition rate. Direct effect of snow cover exerted only a secondary influence. Whatever the species, early snowmelt locations showed consistently reduced decomposition rates and delayed final stages of N mineralization. This lower decomposition rate was associated with freezing soil temperatures during winter. The results suggest that a reduced snow cover may have a weak and immediate direct effect on litter decomposition rates and N availability in alpine tundra. A much larger impact on nutrient cycling is likely to be mediated by longer term changes in the relative abundance of lignin-rich dwarf shrubs.     

Insensitivity of Soil Microbial Activity to Temporal Variation in Soil N in Subarctic Tundra: Evidence from Responses to Large Migratory Grazers

Large migratory grazers commonly influence soil processes in tundra ecosystems. However, the extent to which grazing effects are limited to intensive grazing periods associated with migration has not previously been investigated. We analyzed seasonal patterns in soil nitrogen (N), microbial respiration and extracellular enzyme activities (EEAs) in a lightly grazed tundra and a heavily grazed tundra that has been subjected to intensive grazing during reindeer (Rangifer tarandus L.) migration for the past 50 years. We hypothesized that due to the fertilizing effect of the reindeer, microbial respiration and EEAs related to microbial C acquisition should be higher in heavily grazed areas compared to lightly grazed areas and that the effects of grazing should be strongest during reindeer migration. Reindeer migration caused a dramatic peak in soil N availability, but in contrast to our predictions, the effect of grazing was more or less constant over the growing season and the seasonal patterns of microbial activities and microbial N were strikingly uniform between the lightly and heavily grazed areas. Microbial respiration and the EEAs of β-glucosidase, acid-phosphatase, and leucine-aminopeptidase were higher, whereas that of N-acetylglucosamidase was lower in the heavily grazed area. Experimental fertilization had no effect on EEAs related to C acquisition at either level of grazing intensity. Our findings suggest that soil microbial activities were independent of grazing-induced temporal variation in soil N availability. Instead, the effect of grazing on soil microbial activities appeared to be mediated by substrate availability for soil microorganisms. Following a shift in the dominant vegetation in response to grazing from dwarf shrubs to graminoids, the effect of grazing on soil processes is no longer sensitive to temporal grazing patterns; rather, grazers exert a consistent positive effect on the soil microbial potential for soil C decomposition.     

Interactive Effects of Ungulate Herbivores, Soil Fertility, and Variable Rainfall on Ecosystem Processes in a Semi-arid Savanna

Large herbivores can both positively and negatively affect primary productivity and rates of nutrient cycling in different ecosystems. Positive effects of grazers in grasslands have been attributed to migratory behavior of the dominant ungulate species and soil fertility. We studied the effects of grazers on aboveground net primary productivity (ANPP) and N cycling on central Kenyan rangeland characterized by intense, chronic grazing by a mixed community of cattle and resident native ungulates. Exclosure studies conducted at high and low levels of soil fertility showed that both soil fertility and annual rainfall patterns mediate the effects of grazers on ANPP and N cycling. In a low-rainfall year with short (1 month) growing seasons, grazers reduced aboveground productivity regardless of soil nutrient availability. However, in a high-rainfall year with a 5-month growing season, grazers increased ANPP on nutrient-rich glades and suppressed ANPP on nutrient-poor bushland sites. Concomitant studies of grazer effects on N cycling revealed complex interactions with the seasonal pattern of N-mineralization and inorganic N availability. Grazers increased the size of the inorganic N pool available to plants at the onset of the growing season, particularly in nutrient-rich glades. However, grazers also decreased N mineralization rates at all sites early in the growing season. Measures of N availability via ion-exchange resin bags suggested that the combined effects of grazers on inorganic N pool fluctuations and N-mineralization rates resulted in a net increase in N availability at glade sites and a net decrease in N availability at bushland sites. The net effect of grazers on soil N availability mirrored grazer effects on ANPP in the high-rainfall year. Overall, our results suggest that grazer effects on N dynamics are closely linked to effects on productivity and resilience to drought. Furthermore, even under optimal conditions of high soil fertility and above-average rainfall, grazer promotion of ANPP in this chronically grazed system dominated by resident ungulates was small compared to systems dominated by migratory ungulates.     

Nutrient content and dynamics in north Swedish shrub tundra areas

Forbs and leaves of deciduous shrubs had high concentrations of Ca, Mg, K, P, and N. Deciduous dwarf shrubs had intermediate concentrations but higher than evergreen dwarf shrubs. Monocots, cryptogams, woody and belowground tissues had low concentrations.
Plant nutrient concentrations and nutrient content in soil organic matter increased from dry towards moist tundra areas.
The residence time of nutrients was considerably less than ten years in surface litter, but several decades or centuries in total organic matter. The longest residence time was found in the moist part of the tundra.
N, K, and P in Betula nana leaves were translocated to a great extent prior to leaf fall, whereas Mg and particularly Ca were only slightly translocated. As on other tundra areas shortage of nutrients probably limits plant growth. In that case short supply of N and F seems most probable due to retention in litter and soil organic matter.     

Plant responses to fertilization and exclusion of grazers on an arctic tundra heath

This study investigated the impacts of fertilization and grazing by Norwegian lemmings (Lemmus lemmus), grey‐sided voles (Clethrionomys rufocanus), and reindeer (Rangifer tarandus) on a diverse tundra plant community dominated by deciduous shrubs. Four out of eight study areas, having a size of 2500 m² each, were fertilized with a N‐P‐K fertilizer and four areas served as unfertilized controls. Two types of exclosures were used within each study area, one to exclude solely reindeer, and one to exclude both rodents and reindeer. Open, grazed plots served as controls. During 5 years following the fertilization event the changes in vegetation inside and outside the exclosures were monitored using a point frequency method. The densities of rodents on the fertilized and unfertilized areas were investigated by live trapping and by counting nests of overwintering individuals. Reindeer do not graze on the study area during the growing season but migrate through this area in autumn and spring. Fertilization increased the abundance of vascular plants while grazing by reindeer and rodents decreased the abundance of vascular plants significantly on both fertilized and unfertilized areas. Rodents preferred clearly the fertilized areas during winter, decreasing the abundance of Vaccinium myrtillus and Vaccinium vitis‐idaea, while very little grazing occurred during summer. Graminoids showed the strongest positive response to fertilization and dominated the plant community on ungrazed plots, while winter grazing by both reindeer and rodents significantly decreased the abundance of graminoids. Deciduous shrubs (Betula nana, Vaccinium myrtillus) increased slightly but significantly due to fertilization and evergreen dwarf shrubs showed no response to fertilization. However, the use of functional growth forms for predicting the responses of nutrient enrichment and grazing must be questioned, as responses to fertilization as well as preferences by herbivores were shown to be species‐specific rather than uniform within functional groups based on plant growth forms.     

Depth‐based differentiation in nitrogen uptake between graminoids and shrubs in an Arctic tundra plant community

Questions

The rapid climate warming in tundra ecosystems can increase nutrient availability in the soil, which may initiate shifts in vegetation composition. The direction in which the vegetation shifts will co‐determine whether Arctic warming is mitigated or accelerated, making the understanding of successional trajectories urgent. One of the key factors influencing the competitive relationships between plant species is their access to nutrients, depending on the depth where they take up most nutrients. However, nutrient uptake at different soil depths by tundra plant species that differ in rooting depth is unclear.

Location

Kytalyk Nature Reserve, northeast Siberia, Russia.

Methods

We injected ¹⁵N to 5 cm, 15 cm and the thaw front of the soil in a moist tussock tundra. The absorption of ¹⁵N by grasses, sedges, deciduous shrubs and evergreen shrubs from the three depths was compared.

Results

The results clearly show a vertical differentiation of N uptake by these plant functional types, corresponding to their rooting strategy. Shallow‐rooting dwarf shrubs were more capable of absorbing nutrients from the upper soil than from deeper soil. Deep‐rooting grasses and sedges were more capable of absorbing nutrients from deeper soil than the dwarf shrubs. The natural ¹⁵N abundances in control plants also indicate that graminoids can absorb more nutrients from the deeper soil than dwarf shrubs.

Conclusions

Our results show that graminoids and shrubs in the Arctic differ in their N uptake strategies, with graminoids profiting from nutrients released at the thaw front, while shrubs mainly forage in upper soil layers. Our results suggest that tundra vegetation will become graminoid‐dominated as permafrost thaw progresses and nutrient availability increases in the deep soil.     

The effect of reindeer grazing on decomposition, mineralization and soil biota in a dry oligotrophic Scots pine forest

Reindeer grazing in the Fennoscandian area has a considerable influence on the ground vegetation, and this is likely in turn to have important consequences for the soil biota and decomposition processes. The effects of reindeer grazing on soil biota, decomposition and mineralization processes, and ecosystem properties in a lichen‐dominated forest in Finnish Lapland were studied inside and outside a large long term fenced reindeer exclosure area. Decomposition rates of Vaccinium myrtillus leaves in litter bags were retarded in the grazed area relative to the ungrazed area, as well as in subplots from which lichens had been artificially removed to simulate grazing. The effect of reindeer grazing on soil respiration and microbial C was positive in the lichen and litter layers of the soil profile, but retarded in the humus layer. There was no effect of grazing on gross N mineralization and microbial biomass N in the humus and upper mineral soil layer, but net N mineralization was increased by grazing. In these layers soil respiration was reduced by grazing, indicating that reindeer effects reduce the ratio of C to N mineralized by soil microorganisms. Grazing stimulated populations of all trophic groupings of nematodes in the lichen layer and microbe feeding nematodes in the litter layer, indicating that grazing by reindeer has multitrophic effects on the decomposer food‐web. Grazing decreased lichen and dwarf shrub biomasses and increased the mass of litter present in the litter layer on an areal basis, but did not significantly alter total C storage per unit area in the humus and mineral soil layers. The N concentration of lichens was increased by grazing, but the N concentrations of both living and dead Pinus sylvestris needles and Empetrum hermaphroditum leaves were not affected.
There was some evidence for each of three mechanisms which could account for the grazing effects that we observed in our study. Firstly, reindeer may have changed the composition and quality of litter input by affecting plant species composition and through addition of N from urine and faeces, resulting in a lack of available C relative to N for decomposer organisms. Secondly, the organic matter in the soil may be older in the grazed area, because of reduction of recent production of lichen litter relative to the ungrazed area. The organic matter in the grazed area may have been in a different phase of decomposition from that in the exclosure. Thirdly, the soil microclimate is likely to be affected by reindeer grazing through physical removal of lichen cover on the ground, and this can have a significant influence on soil microbial processes. This is supported by the strong observed effects of experimental removal of lichens on decomposer processes. The impact of reindeer grazing on soil processes may be a result of complex interactions between different mechanisms, and this could help to explain why the below‐ground effects of reindeer grazing have different consequences to those which have been observed in recent investigations on other grazing systems.     

The effects of reindeer grazing on the composition and species richness of vegetation in forest–tundra ecotone

We conducted an 8-year exclosure experiment (1999–2006) in a forest–tundra ecotonal area in northwestern Finnish Lapland to study the effects of reindeer grazing on vegetation in habitats of variable productivity and microhabitat structure. The experimental sites included tundra heath, frost heath and riparian habitats, and the two latter habitats were characterized by hummock-hollow ground forms. The total cover of vegetation, cover of willow (Salix spp.), dwarf birch (Betula nana), dwarf shrubs, forbs and grasses (Poaceae spp.) increased in exclosures in all habitats. The increase in the total cover of vegetation and in the covers of willow and dwarf birch tended to be greatest in the least productive tundra heath. Opposing to the increase in the dominant vascular plant groups, the cover and species number of bryophytes decreased in exclosures. We conclude that the effects of reindeer grazing on vegetation composition depend on environmental heterogeneity and the responses vary among plant groups. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal grazing effects by semi-domesticated reindeer on subarctic mountain birch forests

In northern Fennoscandia, the spatial and temporal grazing practices of semi-domesticated reindeer (Rangifer tarandus tarandus) vary, which implies different grazing effects dependent on natural conditions as well as management regime (i.e., timing and intensity of grazing). We compared density and biomass of main plant groups in semi-dry mountain birch forests exposed to either long-term summer or winter grazing in three reindeer herding districts in the northernmost Finland. Percent plant cover, height, and biomass of reindeer lichens (Cladonia spp.) and dwarf shrubs (Vaccinium uliginosum, Calluna vulgaris, and Betula nana) were lower on summer ranges compared with winter ranges. The biomass of other dwarf shrubs (Vaccinium myrtillus and V. vitis-idaea), and graminoids and herbs, and the % cover of non-vegetated bare soil and litter were, however, higher on summer ranges than on winter ranges. Young mountain birch shoots (Betula pubescens ssp. czerepanovii) were less frequent on summer ranges than on winter ranges. The total leaf biomass under the browsing height of reindeer (<1.5 m) was also lower on summer ranges compared with winter ranges. Especially in drier and nutrient poor mountain birch forests, intensive summer grazing reduces the quantity of lichens and total plant biomass which affects the ecological state and productivity of these forests and also reduces especially their winter grazing value for reindeer. Therefore, in addition to regulating the maximum sustained numbers of reindeer, pasture rotation systems that effectively protect dry and nutrient poor vegetation from summer grazing and trampling should be encouraged.     

Regulation of Microbial Community Composition and Activity by Soil Nutrient Availability, Soil pH, and Herbivory in the Tundra

Soil nitrogen (N) availability and pH constitute major abiotic controls over microbial community composition and activity in tundra ecosystems. On the other hand, mammalian grazers form an important biotic factor influencing resource coupling between plants and soil microorganisms. To investigate individual effects and interactions among soil nutrients, pH, and grazing on tundra soils, we performed factorial treatments of fertilization, liming, and grazer exclusion in the field for 3 years at 2 contrasting tundra habitats, acidic (N-poor) and non-acidic (N-rich) tundra heaths. The effects of all treatments were small in the non-acidic tundra heaths. In the acidic tundra heaths, fertilization decreased the fungal:bacterial ratio as analyzed by soil PLFAs, but there were no effects of liming. Fertilization increased soil N concentrations more drastically in ungrazed than grazed plots, and in parallel, fertilization decreased the fungal:bacterial ratio to a greater extent in the ungrazed plots. Liming, on the other hand, partly negated the effects of fertilization on both soil N concentrations and PLFAs. Fertilization drastically increased the activity of phenol oxidase, a microbial enzyme synthesized for degradation of soil phenols, in grazed plots, but had no effect in ungrazed plots. Taken together, our results demonstrate that grazers have the potential to regulate the fungal:bacterial ratio in soils through influencing N availability for the soil microorganisms.     

Reindeer influence on ecosystem processes in the tundra

Reindeer have been recorded to increase nutrient cycling rate and primary production in studies from fences almost 40 years old that separate areas with different grazing regimes in northern Fennoscandia. To further understand the mechanism behind the effects of herbivores on primary production, we measured the size of the major C and N pools, soil temperature, litter decomposition rate and N mineralization rate in lightly, moderately and heavily grazed areas along two of these fences.
Plant N found in new biomass, indicative of plant N assimilation, was significantly higher in moderately and heavily grazed areas than in lightly grazed areas, which corresponded to a decreased amount of N in old plant parts. The amount of N found in plant litter or organic soil layer did not differ between the grazing treatments. Together with soil N concentrations and litter decomposition rates, soil temperatures were significantly higher in moderately and heavily grazed areas.
We conclude that the changes in soil temperature are important for the litter decomposition rate and thus on the nutrient availability for plant uptake. However, the changes in plant community composition appear to be more important for the altered N pools and thus the enhanced primary production. The results provide some support for the keystone herbivore hypothesis, which states that intensive grazing can promote a transition from moss-rich tundra heath to productive grasslands. Grazing altered N fluxes and pools, but the total N pools were similar in all grazing treatments. Our study thus indicates that grazing can increase the primary production through enhancing the soil nutrient cycling rate, even in a long term perspective on an ecological timescale.     

Role of litter decomposition for the increased primary production in areas heavily grazed by reindeer: a litterbag experiment

Heavy grazing and trampling by reindeer increase nutrient cycling and primary production in areas where grasslands have replaced shrub and moss tundra. One way in which herbivores can affect nutrient cycling is through changing the litter decomposition processes. We studied the effect of herbivory on litter decomposition rate by reciprocal transplantation of litter between lightly grazed and heavily grazed areas, using the litterbag technique. We used litter from two of the most common species on the lightly grazed side, Betula nana and Empetrum nigrum , and two of the most common species on the heavily grazed side, Carex bigelowii and Deschampsia flexuosa . We found that herbivory improved litter quality by favouring species with easily decomposed litter. However, herbivory also improved litter quality by increasing the nitrogen content and lowering the C/N ratio of each species. Decomposition rates even correlated with the abundance of the plant category in question. Shrub litter decomposed faster in the lightly grazed area where shrubs were common, and graminoid litter decomposed faster in the heavily grazed area where graminoids were common. These results indicate that the decomposer micro-organisms are adapted to the most common litter types. This study shows that detailed information about the effect of herbivory on litter quality is important to understand differences between the short-term and long-term effects of herbivory on nutrient cycling and primary production.     

Grazing history effects on above- and below-ground litter decomposition and nutrient cycling in two co-occurring grasses

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment.     

Effects of reindeer on boreal forest floor vegetation: Does grazing cause vegetation state transitions?

Intensive reindeer grazing has been hypothesized to drive vegetation shifts in the arctic tundra from a low-productive lichen dominated state to a more productive moss dominated state. Although the more productive state can potentially host more herbivores, it may still be less suitable as winter grazing grounds for reindeer, if lichens, the most preferred winter forage, are less abundant. Therefore, such a shift towards mosses may have severe consequences for reindeer husbandry if ground-growing lichens have difficulties to recover. We tested if reindeer cause this type of vegetation state shifts in boreal forest floor vegetation, by comparing plant species composition and major soil processes inside and outside of more than 40-year-old exclosures. Lichen biomass was more than twice as high inside exclosures than in grazed controls and almost 5 times higher than in heavily grazed patches. Contrary to our predictions, net N mineralization and plant production were higher in the exclosures than in the grazed controls. The lack of response of phytometer plants in a common garden bioassay indicated that changed soil moisture may drive effects of reindeer on plant productivity in these dry Pine forest ecosystems.     

Vertebrate herbivores and ecosystem control: cascading effects of faeces on tundra ecosystems

We tested the hypothesis that large herbivores manipulate their own food supply by modifying soil nutrient availability. This was investigated experimentally the impact of faeces on grasses, mosses and soil biological properties in tundra ecosystems. For this, we increased the density of reindeer Rangifer tarandus platyrhynchus faeces and studied the response of a tundra system on Spitsbergen to this single faecal addition treatment for four subsequent years. From the third year onwards faecal addition had unambiguously enhanced the standing crop of grasses, as evidenced by an increase in both shoot density and mass per shoot. Although reindeer grazing across experimental plots was positively related to the abundance of grasses in anyone year, the increase in grass abundance in fouled plots failed to result in greater grazing pressure in those plots. Faecal addition enhanced soil microbial biomass C and N, particularly under wet conditions where faecal decay rates were greatest, whilst grasses appeared to benefit from faeces under dry conditions. Whilst growth of grasses and soil microbial biomass were stimulated by faecal addition, the depth of the extensive moss layer that is typical of tundra ecosystems was significantly reduced in fouled plots four years after faecal addition. The greatest reduction in moss depth occurred where fouling increased soil microbial biomass most, suggesting that enhanced decomposition of moss by a more abundant microbial community may have caused the reduced moss layer depth in fouled plots. Our field experiment demonstrates that by the production of faeces alone, vertebrate herbivores greatly impact on both above‐ and belowground components of tundra ecosystems and in doing so manipulate their own food supply. Our findings verify the assertion that grazing is of fundamental importance to tundra ecosystem productivity, and support the hypothesis that herbivory is instrumental in promoting grasses whilst suppressing mosses. The widely observed inverse relationship between grass and moss abundance in the field may therefore reflect the long history of plant‐herbivore interactions in tundra ecosystems.     

Effects of plant species on nutrient cycling

Plant species create positive feedbacks to patterns of nutrient cycling in natural ecosystems. For example, in nutrient-poor ecosystems, plants grow slowly, use nutrients efficiently and produce poor-quality litter that decomposes slowly and deters herbivores. /n contrast, plant species from nutrient-rich ecosystems grow rapidly, produce readily degradable litter and sustain high rates of herbivory, further enhancing rates of nutrient cycling. Plants may also create positive feedbacks to nutrient cycling because of species' differences in carbon deposition and competition with microbes for nutrients in the rhizosphere. New research is showing that species' effects can be as or more important than abiotic factors, such as climate, in controlling ecosystem fertility.