Plant diversity effects on soil food webs are stronger than those of elevated CO2 and N deposition in a long-term grassland experiment

Recent metaanalyses suggest biodiversity loss affects the functioning of ecosystems to a similar extent as other global environmental change agents. However, the abundance and functioning of soil organisms have been hypothesized to be much less responsive to such changes, particularly in plant diversity, than aboveground variables, although tests of this hypothesis are extremely rare. We examined the responses of soil food webs (soil microorganisms, nematodes, microarthropods) to 13-y manipulation of multiple environmental factors that are changing at global scales-specifically plant species richness, atmospheric CO2, and N deposition-in a grassland experiment in Minnesota. Plant diversity was a strong driver of the structure and functioning of soil food webs through several bottom-up (resource control) effects, whereas CO2 and N only had modest effects. We found few interactions between plant diversity and CO2 and N, likely because of weak interactive effects of those factors on resource availability (e.g., root bio-mass). Plant diversity effects likely were large because high plant diversity promoted the accumulation of soil organic matter in the site's sandy, organic matter-poor soils. Plant diversity effects were not explained by the presence of certain plant functional groups. Our results underline the prime importance of plant diversity loss cascading to soil food webs (density and diversity of soil organisms) and functions. Because the present results suggest prevailing plant diversity effects and few interactions with other global change drivers, protecting plant diversity may be of high priority to maintain the biodiversity and functioning of soils in a changing world

Nematode community shifts in response to experimental warming and canopy conditions are associated with plant community changes in the temperate-boreal forest ecotone

Global climate warming is one of the key forces driving plant community shifts, such as range shifts of temperate species into boreal forests. As plant community shifts are slow to observe, ecotones, boundaries between two ecosystems, are target areas for providing early evidence of ecological responses to warming. The role of soil fauna is poorly explored in ecotones, although their positive and negative effects on plant species can influence plant community structure. We studied nematode communities in response to experimental warming (ambient, +1.7, +3.4 °C) in soils of closed and open canopy forest in the temperate-boreal ecotone of Minnesota, USA and calculated various established nematode indices. We estimated species-specific coverage of understory herbaceous and shrub plant species from the same experimental plots and tested if changes in the nematode community are associated with plant cover and composition. Individual nematode trophic groups did not differ among warming treatments, but the ratio between microbial-feeding and plant-feeding nematodes increased significantly and consistently with warming in both closed and open canopy areas and at both experimental field sites. The increase in this ratio was positively correlated with total cover of understory plant species, perhaps due to increased predation pressure on soil microorganisms causing higher nutrient availability for plants. Multivariate analyses revealed that temperature treatment, canopy conditions and nematode density consistently shaped understory plant communities across experimental sites. Our findings suggest that warming-induced changes in nematode community structure are associated with shifts in plant community composition and productivity in the temperate-boreal forest ecotones

Light, earthworms, and soil resources as predictors of diversity of 10 soil invertebrate groups across monocultures of 14 tree species

Management of biodiversity and ecosystem services requires a better understanding of the factors that influence soil biodiversity. We characterized the species (or genera) richness of 10 taxonomic groups of invertebrate soil animals in replicated monocultures of 14 temperate tree species. The focal invertebrate groups ranged from microfauna to macrofauna: Lumbricidae, Nematoda, Oribatida, Gamasida, Opilionida, Araneida, Collembola, Formicidae, Carabidae, and Staphylinidae. Measurement of invertebrate richness and ancillary variables occurred ~34 years after the monocultures were planted. The richness within each taxonomic group was largely independent of richness of other groups; therefore a broad understanding of soil invertebrate diversity requires analyses that are integrated across many taxa. Using a regression-based approach and ~125 factors related to the abundance and diversity of resources, we identified a subset of predictors that were correlated with the richness of each invertebrate group and richness integrated across 9 of the groups (excluding earthworms). At least 50% of the variability in integrated richness and richness of each invertebrate group was explained by six or fewer predictors. The key predictors of soil invertebrate richness were light availability in the understory, the abundance of an epigeic earthworm species, the amount of phosphorus, nitrogen, and calcium in soil, soil acidity, and the diversity or mass of fungi, plant litter, and roots. The results are consistent with the hypothesis that resource abundance and diversity strongly regulate soil biodiversity, with increase in resources (up to a point) likely to increase the total diversity of soil invertebrates. However, the relationships between various resources and soil invertebrate diversity were taxon-specific. Similarly, diversity of all 10 invertebrate taxa was not high beneath any of the 14 tree species. Thus, changes to tree species composition and resource availability in temperate forests will likely increase the richness of some soil invertebrates while decreasing the richness of others

Effects of litter traits, soil biota, and soil chemistry on soil carbon stocks at a common garden with 14 tree species

Tree species interact with soil biota to impact soil organic carbon (C) pools, but it is unclear how this interaction is shaped by various ecological factors. We used multiple regression to describe how ~100 variables were related to soil organic C pools in a common garden experiment with 14 temperate tree species. Potential predictor variables included: (i) the abundance, chemical composition, and decomposition rates of leaf litter and fine roots, (ii) species richness and abundance of bacteria, fungi, and invertebrate animals in soil, and (iii) measures of soil acidity and texture. The amount of organic C in the organic horizon and upper 20 cm of mineral soil (i.e. the combined C pool) was strongly negatively correlated with earthworm abundance and strongly positively correlated with the abundance of aluminum, iron, and protons in mineral soils. After accounting for these factors, we identified additional correlations with soil biota and with litter traits. Rates of leaf litter decomposition, measured as litter mass loss, were negatively correlated with size of the combined soil organic C pool. Somewhat paradoxically, the combined soil organic C pool was also negatively related to the ratio of recalcitrant compounds to nitrogen in leaf litter. These apparent effects of litter traits probably arose because two independent components of litter “quality” were controlling different aspects of decomposition. Leaf litter mass loss rates were positively related with leaf litter calcium concentrations, reflecting greater utilization and depolymerization of calcium-rich leaf litter by earthworms and other soil biota, which presumably led to greater proportional losses of litter C as CO2 or dissolved organic C. The fraction of depolymerized and metabolized litter that is ultimately lost as CO2 is an inverse function of microbial C use efficiency, which increases with litter nutrient concentrations but decreases with concentrations of recalcitrant compounds (e.g. lignin); thus, high ratios of recalcitrant compounds to nitrogen in leaf litter likely caused a greater fraction of depolymerized litter to be lost as CO2. Existing conceptual models of soil C stabilization need to reconcile the effects of litter quality on these two potentially counteracting factors: rates of litter depolymerization and microbial C use efficiency

Light, earthworms, and soil resources as predictors of diversity of 10 soil invertebrate groups across monocultures of 14 tree species

Management of biodiversity and ecosystem services requires a better understanding of the factors that influence soil biodiversity. We characterized the species (or genera) richness of 10 taxonomic groups of invertebrate soil animals in replicated monocultures of 14 temperate tree species. The focal invertebrate groups ranged from microfauna to macrofauna: Lumbricidae, Nematoda, Oribatida, Gamasida, Opilionida, Araneida, Collembola, Formicidae, Carabidae, and Staphylinidae. Measurement of invertebrate richness and ancillary variables occurred ~34 years after the monocultures were planted. The richness within each taxonomic group was largely independent of richness of other groups; therefore a broad understanding of soil invertebrate diversity requires analyses that are integrated across many taxa. Using a regression-based approach and ~125 factors related to the abundance and diversity of resources, we identified a subset of predictors that were correlated with the richness of each invertebrate group and richness integrated across 9 of the groups (excluding earthworms). At least 50% of the variability in integrated richness and richness of each invertebrate group was explained by six or fewer predictors. The key predictors of soil invertebrate richness were light availability in the understory, the abundance of an epigeic earthworm species, the amount of phosphorus, nitrogen, and calcium in soil, soil acidity, and the diversity or mass of fungi, plant litter, and roots. The results are consistent with the hypothesis that resource abundance and diversity strongly regulate soil biodiversity, with increases in resources (up to a point) likely to increase the total diversity of soil invertebrates. However, the relationships between various resources and soil invertebrate diversity were taxon-specific. Similarly, diversity of all 10 invertebrate taxa was not high beneath any of the 14 tree species. Thus, changes to tree species composition and resource availability in temperate forests will likely increase the richness of some soil invertebrates while decreasing the richness of others