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.
There has been growing recent use of elevational gradients as tools for assessing effects of temperature changes on vegetation properties, because these gradients enable temperature effects to be considered over larger spatial and temporal scales than is possible through conventional experiments. While many studies have explored the direct effects of temperature, the indirect effects of temperature through its long-term influence on soil abiotic or biotic properties remain essentially unexplored. We performed two climate chamber experiments using soils from a subarctic elevational gradient in Abisko, Sweden to investigate the direct effects of temperature, and indirect effects of temperature via soil legacies, on growth of two grass species. The soils were collected from each of two vegetation types (heath, dominated by dwarf shrubs, and meadow, dominated by graminoids and herbs) at each of three elevations. We found that plants responded to both the direct effect of temperature and its indirect effect via soil legacies, and that direct and indirect effects were largely decoupled. Vegetation type was a major determinant of plant responses to both the direct and indirect effects of temperature; responses to soils from increasing elevation were stronger and showed a more linear decline for meadow than for heath soils. The influence of soil biota on plant growth was independent of elevation, with a positive influence across all elevations regardless of soil origin for meadow soils but not for heath soils. Taken together, this means that responses of plant growth to soil legacy effects of temperature across the elevational gradient were driven primarily by soil abiotic, and not biotic, factors. These findings emphasize that vegetation type is a strong determinant of how temperature variation across elevational gradients impacts on plant growth, and highlight the need for considering both direct and indirect effects of temperature on plant responses to future climate change.
Nitrogen (N), acquired by roots and mycorrhizal fungi and supplied to plant foliage, is a growth-limiting nutrient at the subarctic treeline. Due to this limitation, interspecific competition and acquisition of N is an important control on plant community composition and distribution. The ability of trees and shrubs to access N shapes community dynamics at this ecotone undergoing species range shifts and changes in primary productivity driven by climate change. Using 15N soil labelling we investigate the fate of soil inorganic N, and spatial distances over which trees and understorey shrubs access soil N, in a treeline forest. 15N was injected into soil rooting zones in discrete 1 m2 patches and foliar samples were collected from trees between 1 and 50 m away, and understorey shrubs between 0.5 and 11 m away from labelled soil. The 15N label was found in mountain birch trees up to 5 m, and in understorey shrubs up to 2 m, away from labelled soil. We estimate that 1.27% of pulse-derived N was found in foliage of birch trees, compared to 1.16% in the understorey. However, mountain birch trees contributed only 31% of ecosystem leaf area index (LAI), thus there was a disproportionate allocation of added label to the birch canopy compared with its contribution to ecosystem LAI. The difference in root and mycorrhizal exploration distances and community N partitioning between mountain birch trees and understorey shrubs may confer competitive advantage to trees with respect to nitrogen and nutrient patches, which may alter plant community structures within these forests. This is particularly important considering predicted climate-driven tree and tall shrub expansion in subarctic regions, with likely consequences for ecosystem N and carbon (C) cycling, as well as for community composition and biodiversity.
Global warming has created a need for studies along climatic gradients to assess the effects of temperature on ecological processes. Altitudinal and latitudinal gradients are often used as such, usually in combination with air temperature data from the closest weather station recorded at 1.5–2 m above the ground. However, many ecological processes occur in, at, or right above the soil surface. To evaluate how representative the commonly used weather station data are for the microclimate relevant for soil surface biota, we compared weather station temperatures for an altitudinal (500–900 m a.s.l.) and a latitudinal gradient (49–68°N) with data obtained by temperature sensors placed right below the soil surface at five sites along these gradients. The mean annual temperatures obtained from weather stations and adjusted using a lapse rate of −5.5°C km−1 were between 3.8°C lower and 1.6°C higher than those recorded by the temperature sensors at the soil surface, depending on the position along the gradients. The monthly mean temperatures were up to 10°C warmer or 5°C colder at the soil surface. The within‐site variation in accumulated temperature was as high as would be expected from a 300 m change in altitude or from a 4° change in latitude or a climate change scenario corresponding to warming of 1.6–3.8°C. Thus, these differences introduced by the decoupling are significant from a climate change perspective, and the results demonstrate the need for incorporating microclimatic variation when conducting studies along altitudinal or latitudinal gradients. We emphasize the need for using relevant temperature data in climate impact studies and further call for more studies describing the soil surface microclimate, which is crucial for much of the biota.
Treelines have drawn persistent research interest as they can respond markedly to climate. However, the mechanisms that determine tree seedling recruitment and the response of the forest-tundra ecotone to environmental changes remain poorly understood. We hypothesise that treeline tree seedling performance depends on the interplay between climatic and soil nutritional changes and facilitative and competitive interactions between trees and shrubs. We conducted a seedling transplantation experiment with Betula pubescens at a subarctic treeline, in northern Sweden, which followed a full factorial design with four treatment factors relating to environmental regimes of stress and resource availability: site (forest vs treeline); temperature (+/- passive warming); shrub presence (+/-Vaccinium myrtillus removal); and nutrient availability (+/- NPK addition). During three growing seasons we assessed the establishment and performance of Betula. The experimental manipulations caused highly significant effects on seedling performance. Although Vaccinium enhanced seedling survival and reduced the effects of excessive solar radiation and insect herbivory, the seedlings growing with the shrub had a poorer performance by the end of the experimental period. Also, seedlings in the forest had a poorer performance than those at the treeline. Betula seedlings showed a very pronounced and positive response to passive warming and to nutrient addition, but such effects were more evident at the treeline site and often interacted with the presence of Vaccinium. This experiment shows that shrubâtree interactions are important drivers of subarctic treeline dynamics and that they vary with time and space. Facilitation, competition, herbivory and environmental changes at the tree seedling stage act as important filters in structuring the forestâtundra ecotone. We demonstrate that changes in this ecotone cannot be simply predicted from changing temperature patterns alone, and that complex interactions need to be considered, not only between shrubs and trees, but also with herbivores and between warming and soil nutrient availability.
The relative importance of nutrient supply and food chain composition for the structure and function of ''two-level'' South Georgian (Sub-Antarctica) freshwater ecosystems was assessed. Along a productivity gradient of 19 lakes, we determined the abundance of bacteria, bacterivorous flagellates, and herbivorous macrozooplankters, as well as the concentrations of nutrients (phosphorus and nitrogen) and chlorophyll. Our aims were to test predictions from ''classical'' food chain theory on these simple ecosystems. and to include microbial components in the analysis. The abundance of all organisms increased with increasing productivity, although chlorophyll concentration showed a damped increase compared with lower latitude lakes with similar productivity. Moreover, the amount of chlorophyll per zooplankton grazer showed no increase with increasing productivity, indicating that grazing is a major structuring factor for algal biomass development. Similarly, there was no increase in the ratio between the abundances of flagellates (an alternative prey for zooplankters) and zooplankton with increasing productivity. Conversely, the number of bacteria per flagellate increased with increasing productivity, suggesting that bacteria are not regulated by flagellate grazing. We conclude that food chain theory may explain a considerable part of the variation in abundance of organisms of all size classes, including microorganisms.
Several different factors may determine where species range limits are located within regions of otherwise continuously available habitat and suitable climate. Within the Arctic tundra biome many bird species are migratory and their breeding distributions are affected by migration routes that are in turn limited by factors such as suitable winter habitat, migratory stopover sites, geographical barriers and historical routes of colonization. We identified longitudinal zones in the circumpolar Arctic of pronounced changes in the avian species composition (high species spatial turnover; ‘species divides’). We tested for the association between migratory status and the geographical location and numbers of such species divides for species with non-breeding habitats mainly within terrestrial, pelagic and coastal ecosystems. Our results demonstrate that migration is of profound importance for both the number and locations of species divides in the Arctic. Long-distance migration is associated with a large number of divides among terrestrial and coastal arctic birds but with a reduced number of divides among pelagic birds. We suggest that long-distance migration permits pelagic but not terrestrial and coastal birds to colonize large winter ranges, which in turn causes expansion of breeding ranges, with more homogenous communities and reduction of species divides as consequences, among the long-distance migrants of pelagic but not of terrestrial and coastal birds. Furthermore, the divides among long-distance migrants are situated in two main regions, the Beringia and Greenland zones, while divides among short-distance migrants are more evenly spaced throughout the circumpolar Arctic. The Beringia and Greenland divides result largely from inter-continental colonization of new breeding ranges but retainment of original winter quarters in a process of evolution through extension of migration routes, leading to aggregated divides in the meeting zones of major global flyways.
Recent studies have shown that biotic interactions can shape species’ distributions, but empirical data on multiple biotic interactions are scarce. Therefore, we examined effects of plant–plant and plant–herbivore interactions on plant survival, growth and reproduction at different altitudes. For these purposes we conducted a factorial neighbor removal and large herbivore exclusion experiment with six transplant species (three tall forbs with their main distribution at low altitudes and three small forbs with their main distribution at high altitudes) on Låktačohkka Mountain, northern Sweden, replicated at two altitudes (ca 600 and 900 m a.s.l.) and consequently a 2.1°C difference in summer air temperatures. Overall transplant survival was 93%. Two out of three tall forbs grew better at low than at high altitudes, while no significant differences in growth between altitudes were found for any of the three small forbs. Since the main difference in abiotic conditions between the altitudes was most likely in temperature (as the sites were topographically and edaphically matched as closely as possible), this result indicates that climatic warming could induce upward migration of tall low-altitude forbs. Negative plant–plant interactions prevailed at both altitudes, and we found indications that competition may set the lower altitudinal limits of some small tundra forbs. Thus, increased competition in response to climate warming may potentially shift the lower margins of high-altitude forbs’ distributions upward. Large mammalian grazers reduced the growth of tall forbs and enhanced the flowering of small forbs, and grazers could thus at least partly counteract the anticipated warming-induced distribution shifts.
The decomposition rates of plant litter mixtures often deviate from the averaged rates of monocultures of their component litter species. The mechanisms behind these non-additive effects in decomposition of litter mixtures are lively debated. One plausible explanation for non-additive effects is given by the improved microenvironmental condition (IMC) theory. According to this theory, plant litter species, whose physical characteristics improve the microclimatic conditions for decomposers, will promote the decomposition of their co-occurring litter species. We tested the IMC theory in relation to leaf litter and soil moisture in two contrasting moisture conditions in a dry subarctic mountain birch forest with vascular plant leaf litters of poor and high quality. The non-additive effects in mass loss of litter mixtures increased when moisture conditions in litter and soil became more favourable for plant litter decomposition. The sign of this increase (antagonistic or synergistic) in non-additive effects was more predictable for litter mixtures of poor litter quality. Although the specific mechanisms underlying the IMC theory depended on the litter quality of the litter mixtures, a standardized water holding capacity (WHC) was the litter trait most closely related to the non-additive effects in mixtures of both poor and high quality litter types. Furthermore, we found that higher dissimilarity in WHC traits between the component litter species in a mixture increased synergistic effects in litter mixtures under limiting moisture conditions. However, under improved moisture conditions, increased antagonistic effects were observed. Thus, we found clear support for the IMC theory and showed that climatic conditions and leaf litter physical traits determine whether the non-additive effects in litter mixtures are antagonistic or synergistic. Our study emphasizes the need to include litter physical traits into predictive models of mixing effects on plant litter decomposition and in general suggests climate specificity into these models.
Several important fitness components in herbaceous perennial plants are commonly related to plant size: flowering probability, reproductive allocation and fecundity. However, evidence for such size-dependence of fitness components is mostly anecdotal and unconnected to other life history traits. Here we report size-dependence for flowering probability and reproductive allocation in 11 populations of Pinguicula vulgaris and relate it to environmental factors. Flowering probability was size-dependent in all populations of P. vulgaris, and indicated the existence of a threshold size for reproduction. Populations at low altitudes and in wet soils showed a significantly higher threshold size for reproduction. Reproductive mass was also size-dependent in all populations. We found considerable between-population differences in the slope and the intercept of the regression between plant size and reproductive mass. This variation was weakly related to the environmental factors measured. In general, relationships between different size-dependent fitness components were low. Instead of showing a covariation of traits, in line with interpretations in terms of life history “tactics”, P. vulgaris seemed to independently vary each size-dependent fitness component in each locality. In particular, no significant relationship was found between threshold size for reproduction and the slope of size-dependent reproductive allocation, as predicted by previous authors. Neither we found a significant influence of somatic cost of reproduction on size-dependent fitness components.
Herbivores play a key role in the carbon (C) cycle of arctic ecosystems, but these effects are currently poorly represented within models predicting land–atmosphere interactions under future climate change. Although some studies have examined the influence of various individual species of herbivores on tundra C sequestration, few studies have directly compared the effects of different herbivore assemblages. We measured peak growing season instantaneous ecosystem carbon dioxide (CO2) exchange (photosynthesis, respiration and net ecosystem exchange) on replicated plots in arctic tundra which, for 14 years, have excluded different portions of the herbivore population (grazed controls, large mammals excluded, both small and large mammals excluded). Herbivory suppressed photosynthetic CO2 uptake, but caused little change in ecosystem respiration. Despite evidence that small mammals consume a greater portion of plant biomass in these ecosystems, the effect of excluding only large herbivores was indistinguishable from that of excluding both large and small mammals. The herbivory-induced decline in photosynthesis was not entirely attributable to a decline in leaf area but also likely reflects shifts in plant community composition and/or species physiology. One shrub species – Betula nana – accounted for only around 13% of total aboveground vascular plant biomass but played a central role in controlling ecosystem CO2 uptake and release, and was suppressed by herbivory. We conclude that herbivores can have large effects on ecosystem C cycling due to shifts in plant aboveground biomass and community composition. An improved understanding of the mechanisms underlying the distinct ecosystem impacts of different herbivore groups will help to more accurately predict the net impacts of diverse herbivore communities on arctic C fluxes.
The use of celestial or geomagnetic orientation cues can lead migratory birds along different migration routes during the migratory journeys, e.g. great circle routes (approximate), geographic or magnetic loxodromes. Orientation cage experiments have indicated that migrating birds are capable of detecting magnetic compass information at high northern latitudes even at very steep angles of inclination. However, starting a migratory journey at high latitudes and following a constant magnetic course often leads towards the North Magnetic Pole, which means that the usefulness of magnetic compass orientation at high latitudes may be questioned. Here, we compare possible long-distance migration routes of three species of passerine migrants breeding at high northern latitudes. The initial directions were based on orientation cage experiments performed under clear skies and simulated overcast and from release experiments under natural overcast skies. For each species we simulated possible migration routes (geographic loxodrome, magnetic loxodrome and sun compass route) by extrapolating from the initial directions and assessing a fixed orientation according to different compass mechanisms in order to investigate what orientation cues the birds most likely use when migrating southward in autumn. Our calculations show that none of the compass mechanisms (assuming fixed orientation) can explain the migration routes followed by night-migrating birds from their high Nearctic breeding areas to the wintering sites further south. This demonstrates that orientation along the migratory routes of arctic birds (and possibly other birds as well) must be a complex process, involving different orientation mechanisms as well as changing compass courses. We propose that birds use a combination of several compass mechanisms during a migratory journey with each of them being of a greater or smaller importance in different parts of the journey, depending on environmental conditions. We discuss reasons why birds developed the capability to use magnetic compass information at high northern latitudes even though following these magnetic courses for any longer distance will lead them along totally wrong routes. Frequent changes and recalibrations of the magnetic compass direction during the migratory journey are suggested as a possible solution.
Both theoretical arguments and empirical evidence suggests that herbivory in general and mammalian winter herbivory in particular is important in arctic-alpine ecosystems. Although knowledge of the effect of herbivores on specific plants and communities is quite extensive, little is known about the relative impact of large and small vertebrate herbivores and how it might vary among different habitats. To address this key issue, we established exclosures with two different mesh sizes in forest and nearby tundra at three different sites in four contrasting locations in the forest-tundra ecotone in northernmost Sweden and Norway. Plant community composition was recorded annually in three permanent plots within each exclosure and an unfenced control. Local densities of vertebrate herbivores were estimated in spring and autumn from 1998 to 2002. Reindeer (Rangifer tarandus) were the most abundant large vertebrate while Norwegian lemmings (Lemmus lemmus) and grey-sided voles (Clethrionomys rufocanus) were the most common small vertebrates. The study reveals that voles and lemmings have larger effects on the vegetation than reindeer in both habitats in all four locations, even though densities of reindeer differ between locations and only two locations experienced lemming peaks during the period of the experiment. The relative abundance of five of the fifteen most common species was significantly influenced by voles and lemmings whereas only a single species was significantly influenced by reindeer. Different analyses give contrasting results on the importance of herbivory in forest versus open heathlands. A principal component analyses revealed that herbivory influenced the vegetation more in open heathlands than in forests. However, an importance index of herbivores did not differ between forest and open heathlands. Moreover, none of the plant species responded differently in the two habitats, when herbivores were removed. Our results suggest that intense and localised selective foraging by small mammals may have a more marked effect on vegetation than transient feeding by reindeer.
In long-lived and monogamous species, with partners separated between breeding seasons, the previous mate usually returns and waiting is therefore rewarded, thus contributing to mate retention as an adaptive strategy. However, in the long-lived monogamous king penguin, Aptenodytes patagonicus, studied at South Georgia for six years, the overall average divorce rate was as high as 81% (range 76-85%). Outcome of previous breeding attempt had no significant effect on the probability of divorce when all years were pooled, but there was an effect when two years with total breeding failure were excluded. It was obvious that asynchrony in arrival of former mates had strong effect on the probability of divorce. I suggest that the explanation for the high divorce rate in king penguins consists of two parts. (1) Asynchrony in combination with "the expensive fat storing hypothesis". At arrival, birds have only about half their maximum body-reserves. and are close to exhaustion of reserves at the end of the fast ashore. Thus, even birds that switched mates, rather than waited, barely endured the period of fasting. This suggests a trade-off between the costs of divorce and the cost of storing thc extra fuel required to endure waiting and thus ensure mate retention. (2) "The better option hypothesis" in combination with "ideal free mate choice". Even among pairs arriving synchronously, greater than or equal to 50% divorced. To explain this. I suggest that many birds, due to the time constraint created by the near-inadequacy of the fat store, had previously bred with a partner of lower quality than it could have obtained and therefore may benefit from divorcing if a higher-quality partner was available. Readiness to switch mate would be enhanced by low search cost and easy access to potential new mates in this non-territorial species. Hence, mate choice would be potentially ideal free in all aspects but arrival time asynchrony and sometimes competition from the previous mate.
Elevational gradients are useful for predicting how plant communities respond to global warming, because communities at lower elevations experience warmer temperatures. Fine root traits and root trait variation could play an important role in determining plant community responses to warming in cold-climate ecosystems where a large proportion of plant biomass is allocated belowground. Here, we investigated the effects of elevation-associated temperature change on twelve chemical and morphological fine root traits of plant species and plant communities in a Swedish subarctic tundra. We also assessed the relative contributions of plant species turnover and intraspecific variation to the total amount of community-level root trait variation explained by elevation. Several root traits, both at the species and whole community levels, had significant linear or quadratic relationships with elevation, but the direction and strength of these relationships varied among traits and plant species. Further, we found no support for a unidirectional change from more acquisitive root trait values at the lower elevations towards trait values associated with greater nutrient conservation at the higher elevations, either at the species or community level. On the other hand, root trait coefficients of variation at the community level increased with elevation for several root traits. Further, for a large proportion of the community-level traits we found that intraspecific variation was relatively more important than species turnover, meaning that trait plasticity is important for driving community-level trait responses to environmental factors in this tundra system. Our findings indicate that with progressing global warming, intraspecific trait variation may drive plant community composition but this may not necessarily lead to shifts in root resource?acquisition strategy for all species.
An improved knowledge of how contrasting types of plant communities and their associated soil biota differ in their responses to climatic variables is important for better understanding the future impacts of climate change on terrestrial ecosystems. Elevational gradients serve as powerful study systems for answering questions on how ecological processes can be affected by changes in temperature and associated climatic variables. In this study, we evaluated how plant and soil microbial communities, and abiotic soil properties, change with increasing elevation in subarctic tundra in northern Sweden, for each of two dominant but highly contrasting vegetation types, namely heath (dominated by woody dwarf shrubs) and meadow (dominated by herbaceous species). To achieve this, we measured plant community characteristics, microbial community properties and several soil abiotic properties for both vegetation types across an elevation gradient of 500 to 1000 m. We found that the two vegetation types differed not only in several above- and belowground properties, but also in how these properties responded to elevation, pointing to important interactive effects between vegetation type and elevation. Specifically, for the heath, available soil nitrogen and phosphorus decreased with elevation whereas fungal dominance increased, while for the meadow, idiosyncratic responses to elevation for these variables were found. These differences in belowground responses to elevation among vegetation types were linked to shifts in the species and functional group composition of the vegetation. Our results highlight that these two dominant vegetation types in subarctic tundra differ greatly not only in fundamental aboveground and belowground properties, but also in how these properties respond to elevation and are therefore likely to be influenced by temperature. As such they highlight that vegetation type, and the soil abiotic properties that determine this, may serve as powerful determinants of how both aboveground and belowground properties respond to strong environmental gradients.
Sphagnum mosses are key to northern peatland carbon sequestration. They have a range of morphological and anatomical characteristics that allow them to cope with environmental stress. Sphagnum also produces a plethora of biochemicals that may prevent stress-induced cell-damage. However, the linkages between Sphagnum anatomical, morphological and biochemical traits (i.e. metabolites, pigments and antioxidant enzyme activities) are poorly known, neither are their joint responses to environmental change. Here, we quantify and link an array of Sphagnum anatomical, morphological and biochemical traits in five Sphagnum-dominated peatlands distributed along a latitudinal gradient in Europe, covering a range of regional and local environmental conditions. Sphagnum morphological and anatomical traits were intrinsically linked to Sphagnum metabolites and enzyme activities, and these relationships were driven by shared responses to local and regional environmental factors. More particularly, we found that Sphagnum traits can be grouped into four clusters related to growth, biomass, defense and water stress tolerance. We used regional and local environmental conditions data to further show that biochemicals and their specific linkages with some morphological traits describe dimensions of physiology not captured by anatomical and morphological traits alone. These results suggest that Sphagnum morphology and function is rooted in the metabolome, and that incorporating biochemicals into the functional trait space concept can enhance our mechanistic understanding and predictive power in Sphagnum ecology.
Global warming has begun to have a major impact on the species composition and functioning of plant and soil communities. However, long-term community and ecosystem responses to increased temperature are still poorly understood. In this study, we used a well-established elevational gradient in northern Sweden to elucidate how plant, microbial and nematode communities shift with elevation and associated changes in temperature in three highly contrasting vegetation types (i.e. heath, meadow and Salix vegetation). We found that responses of both the abundance and composition of microbial and nematode communities to elevation differed greatly among the vegetation types. Within vegetation types, changes with elevation of plant, microbial and nematode communities were mostly linked at fine levels of taxonomic resolution, but this pattern disappeared when coarser functional group levels were considered. Further, nematode communities shifted towards more conservative nutrient cycling strategies with increasing elevation in heath and meadow vegetation. Conversely, in Salix vegetation microbial communities with conservative strategies were most pronounced at the mid-elevation. These results provide limited support for increasing conservative nutrient cycling strategies at higher elevation (i.e. with a harsher climate). Our findings indicate that climate-induced changes in plant community composition may greatly modify or counteract the impact of climate change on soil communities. Therefore, to better understand and predict ecosystem responses to climate change, it will be crucial to consider vegetation type and its specific interactions with soil communities.