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Fire and herbivory are important natural disturbances in grassy biomes. Both drivers are likely to influence belowground microbial communities but no studies have unravelled the long-term impact of both fire and herbivory on bacterial and fungal communities. We hypothesized that soil bacterial communities change through disturbance-induced shifts in soil properties (e.g. pH, nutrients) while soil fungal communities change through vegetation modification (biomass and species composition). To test these ideas, we characterised soil physico-chemical properties (pH, acidity, C, N, P and exchangeable cations content, texture, bulk density, moisture), plant species richness and biomass, microbial biomass and bacterial and fungal community composition and diversity (using 16S and ITS rRNA amplicon sequencing, respectively) in six long-term (18 to 70 years) ecological research sites in South African savanna and grassland ecosystems. We found that fire and herbivory regimes profoundly modified soil physico-chemical properties, plant species richness and standing biomass. In all sites, an increase in woody biomass (ranging from 12 to 50%) was observed when natural disturbances were excluded. The intensity and direction of changes in soil properties were highly dependent on the topo-pedo-climatic context. Overall, fire and herbivory shaped bacterial and fungal communities through distinct driving forces: edaphic properties (including Mg, pH, Ca) for bacteria, and vegetation (herbaceous biomass and woody cover) for fungi. Fire and herbivory explained on average 7.5 and 9.8% of the fungal community variability, respectively, compared to 6.0 and 5.6% for bacteria. The relatively small changes in microbial communities due to natural disturbance is in stark contrast to dramatic vegetation and edaphic changes and suggests that soil microbial communities, having evolved with disturbance, are resistant to change. This represents both a buffer to short-term anthropogenic-induced change

Fire and herbivory are important natural disturbances in grassy biomes. Both drivers are likely to influence belowground microbial communities but no studies have unravelled the long-term impact of both fire and herbivory on bacterial and fungal communities. We hypothesized that soil bacterial communities change through disturbance-induced shifts in soil properties (e.g. pH, nutrients) while soil fungal communities change through vegetation modification (biomass and species composition). To test these ideas, we characterised soil physico-chemical properties (pH, acidity, C, N, P and exchangeable cations content, texture, bulk density, moisture), plant species richness and biomass, microbial biomass and bacterial and fungal community composition and diversity (using 16S and ITS rRNA amplicon sequencing, respectively) in six long-term (18 to 70 years) ecological research sites in South African savanna and grassland ecosystems. We found that fire and herbivory regimes profoundly modified soil physico-chemical properties, plant species richness and standing biomass. In all sites, an increase in woody biomass (ranging from 12 to 50%) was observed when natural disturbances were excluded. The intensity and direction of changes in soil properties were highly dependent on the topo-pedo-climatic context. Overall, fire and herbivory shaped bacterial and fungal communities through distinct driving forces: edaphic properties (including Mg, pH, Ca) for bacteria, and vegetation (herbaceous biomass and woody cover) for fungi. Fire and herbivory explained on average 7.5 and 9.8% of the fungal community variability, respectively, compared to 6.0 and 5.6% for bacteria. The relatively small changes in microbial communities due to natural disturbance is in stark contrast to dramatic vegetation and edaphic changes and suggests that soil microbial communities, having evolved with disturbance, are resistant to change. This represents both a buffer to short-term anthropogenic-induced change

Fire and herbivory are important natural disturbances in grassy biomes. Both drivers are likely to influence belowground microbial communities but no studies have unravelled the long-term impact of both fire and herbivory on bacterial and fungal communities. We hypothesized that soil bacterial communities change through disturbance-induced shifts in soil properties (e.g. pH, nutrients) while soil fungal communities change through vegetation modification (biomass and species composition). To test these ideas, we characterised soil physico-chemical properties (pH, acidity, C, N, P and exchangeable cations content, texture, bulk density, moisture), plant species richness and biomass, microbial biomass and bacterial and fungal community composition and diversity (using 16S and ITS rRNA amplicon sequencing, respectively) in six long-term (18 to 70 years) ecological research sites in South African savanna and grassland ecosystems. We found that fire and herbivory regimes profoundly modified soil physico-chemical properties, plant species richness and standing biomass. In all sites, an increase in woody biomass (ranging from 12 to 50%) was observed when natural disturbances were excluded. The intensity and direction of changes in soil properties were highly dependent on the topo-pedo-climatic context. Overall, fire and herbivory shaped bacterial and fungal communities through distinct driving forces: edaphic properties (including Mg, pH, Ca) for bacteria, and vegetation (herbaceous biomass and woody cover) for fungi. Fire and herbivory explained on average 7.5 and 9.8% of the fungal community variability, respectively, compared to 6.0 and 5.6% for bacteria. The relatively small changes in microbial communities due to natural disturbance is in stark contrast to dramatic vegetation and edaphic changes and suggests that soil microbial communities, having evolved with disturbance, are resistant to change. This represents both a buffer to short-term anthropogenic-induced change

Established theory addresses the idea that herbivory can have positive feedbacks on nutrient flow to plants. Positive feedbacks likely emerge from a greater availability of organic carbon that primes the soil by supporting nutrient turnover through consumer and especially microbially-mediated metabolism in the detrital pool. We developed an entirely novel stoichiometric model that demonstrates the mechanism of a positive feedback. In particular, we show that sloppy or partial feeding by herbivores increases detrital carbon and nitrogen allowing for greater nitrogen mineralization and nutritive feedback to plants. The model consists of differential equations coupling flows among pools of: plants, herbivores, detrital carbon and nitrogen, and inorganic nitrogen. We test the effects of different levels of herbivore grazing completion and of the stoichiometric quality (carbon to nitrogen ratio, C:N) of the host plant. Our model analyses show that partial feeding and plant C:N interact because when herbivores are sloppy and plant biomass is diverted to the detrital pool, more mineral nitrogen is available to plants because of the stoichiometric difference between the organisms in the detrital pool and the herbivore. This model helps to identify how herbivory may feedback positively on primary production, and it mechanistically connects direct and indirect feedbacks from soil to plant production.

Invasive exotic plant species effects on soil biota and processes in their new range can promote or counteract invasions via changed plant–soil feedback interactions to themselves or to native plant species. Recent meta-analyses reveale that soil influenced by native and exotic plant species is affecting growth and performance of natives more strongly than exotics. However, the question is how uniform these responses are across contrasting life forms. Here, we test the hypothesis that life form matters for effects on soil and plant–soil feedback. In a meta-analysis we show that exotics enhanced C cycling, numbers of meso-invertebrates and nematodes, while having variable effects on other soil biota and processes. Plant effects on soil biota and processes were not dependent on life form, but patterns in feedback effects of natives and exotics were dependent on life form. Native grasses and forbs caused changes in soil that subsequently negatively affected their biomass, whereas native trees caused changes in soil that subsequently positively affected their biomass. Most exotics had neutral feedback effects, although exotic forbs had positive feedback effects. Effects of exotics on natives differed among plant life forms. Native trees were inhibited in soils conditioned by exotics, whereas native grasses were positively influenced in soil conditioned by exotics. We conclude that plant life form matters when comparing plant–soil feedback effects both within and between natives and exotics. We propose that impact analyses of exotic plant species on the performance of native plant species can be improved by comparing responses within plant life form.

Invasive exotic plant species effects on soil biota and processes in their new range can promote or counteract invasions via changed plant–soil feedback interactions to themselves or to native plant species. Recent meta-analyses reveale that soil influenced by native and exotic plant species is affecting growth and performance of natives more strongly than exotics. However, the question is how uniform these responses are across contrasting life forms. Here, we test the hypothesis that life form matters for effects on soil and plant–soil feedback. In a meta-analysis we show that exotics enhanced C cycling, numbers of meso-invertebrates and nematodes, while having variable effects on other soil biota and processes. Plant effects on soil biota and processes were not dependent on life form, but patterns in feedback effects of natives and exotics were dependent on life form. Native grasses and forbs caused changes in soil that subsequently negatively affected their biomass, whereas native trees caused changes in soil that subsequently positively affected their biomass. Most exotics had neutral feedback effects, although exotic forbs had positive feedback effects. Effects of exotics on natives differed among plant life forms. Native trees were inhibited in soils conditioned by exotics, whereas native grasses were positively influenced in soil conditioned by exotics. We conclude that plant life form matters when comparing plant–soil feedback effects both within and between natives and exotics. We propose that impact analyses of exotic plant species on the performance of native plant species can be improved by comparing responses within plant life form., Invasive exotic plant species effects on soil biota and processes in their new range can promote or counteract invasions via changed plant–soil feedback interactions to themselves or to native plant species. Recent meta-analyses reveale that soil influenced by native and exotic plant species is affecting growth and performance of natives more strongly than exotics. However, the question is how uniform these responses are across contrasting life forms. Here, we test the hypothesis that life form matters for effects on soil and plant–soil feedback. In a meta-analysis we show that exotics enhanced C cycling, numbers of meso-invertebrates and nematodes, while having variable effects on other soil biota and processes. Plant effects on soil biota and processes were not dependent on life form, but patterns in feedback effects of natives and exotics were dependent on life form. Native grasses and forbs caused changes in soil that subsequently negatively affected their biomass, whereas native trees caused changes in soil that subsequently positively affected their biomass. Most exotics had neutral feedback effects, although exotic forbs had positive feedback effects. Effects of exotics on natives differed among plant life forms. Native trees were inhibited in soils conditioned by exotics, whereas native grasses were positively influenced in soil conditioned by exotics. We conclude that plant life form matters when comparing plant–soil feedback effects both within and between natives and exotics. We propose that impact analyses of exotic plant species on the performance of native plant species can be improved by comparing responses within plant life form.

Invasive exotic plant species effects on soil biota and processes in their new range can promote or counteract invasions via changed plant–soil feedback interactions to themselves or to native plant species. Recent meta-analyses reveale that soil influenced by native and exotic plant species is affecting growth and performance of natives more strongly than exotics. However, the question is how uniform these responses are across contrasting life forms. Here, we test the hypothesis that life form matters for effects on soil and plant–soil feedback. In a meta-analysis we show that exotics enhanced C cycling, numbers of meso-invertebrates and nematodes, while having variable effects on other soil biota and processes. Plant effects on soil biota and processes were not dependent on life form, but patterns in feedback effects of natives and exotics were dependent on life form. Native grasses and forbs caused changes in soil that subsequently negatively affected their biomass, whereas native trees caused changes in soil that subsequently positively affected their biomass. Most exotics had neutral feedback effects, although exotic forbs had positive feedback effects. Effects of exotics on natives differed among plant life forms. Native trees were inhibited in soils conditioned by exotics, whereas native grasses were positively influenced in soil conditioned by exotics. We conclude that plant life form matters when comparing plant–soil feedback effects both within and between natives and exotics. We propose that impact analyses of exotic plant species on the performance of native plant species can be improved by comparing responses within plant life form., Invasive exotic plant species effects on soil biota and processes in their new range can promote or counteract invasions via changed plant–soil feedback interactions to themselves or to native plant species. Recent meta-analyses reveale that soil influenced by native and exotic plant species is affecting growth and performance of natives more strongly than exotics. However, the question is how uniform these responses are across contrasting life forms. Here, we test the hypothesis that life form matters for effects on soil and plant–soil feedback. In a meta-analysis we show that exotics enhanced C cycling, numbers of meso-invertebrates and nematodes, while having variable effects on other soil biota and processes. Plant effects on soil biota and processes were not dependent on life form, but patterns in feedback effects of natives and exotics were dependent on life form. Native grasses and forbs caused changes in soil that subsequently negatively affected their biomass, whereas native trees caused changes in soil that subsequently positively affected their biomass. Most exotics had neutral feedback effects, although exotic forbs had positive feedback effects. Effects of exotics on natives differed among plant life forms. Native trees were inhibited in soils conditioned by exotics, whereas native grasses were positively influenced in soil conditioned by exotics. We conclude that plant life form matters when comparing plant–soil feedback effects both within and between natives and exotics. We propose that impact analyses of exotic plant species on the performance of native plant species can be improved by comparing responses within plant life form.

The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance.

The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance., The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance.

The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance., The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance.

The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance., The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance.