Search

Soil fauna plays a key role in regulating biogeochemical cycles, but how multiple global change factors (GCFs) may affect faunal communities remains poorly studied. We conducted a meta-analysis using 1154 observations to evaluate the individual and combined effects of elevated CO

Plant litter is the major source of energy and nutrients in stream ecosystems and its decomposition is vital for ecosystem nutrient cycling and functioning. Invertebrates are key contributors to instream litter decomposition, yet quantification of their effects and drivers at the global scale remains lacking. Here, we systematically synthesized data comprising 2707 observations from 141 studies of stream litter decomposition to assess the contribution and drivers of invertebrates to the decomposition process across the globe. We found that (1) the presence of invertebrates enhanced instream litter decomposition globally by an average of 74%; (2) initial litter quality and stream water physicochemical properties were equal drivers of invertebrate effects on litter decomposition, while invertebrate effects on litter decomposition were not affected by climatic region, mesh size of coarse-mesh bags or mycorrhizal association of plants providing leaf litter; and (3) the contribution of invertebrates to litter decomposition was greatest during the early stages of litter mass loss (0-20%). Our results, besides quantitatively synthesizing the global pattern of invertebrate contribution to instream litter decomposition, highlight the most significant effects of invertebrates on litter decomposition at early rather than middle or late decomposition stages, providing support for the inclusion of invertebrates in global dynamic models of litter decomposition in streams to explore mechanisms and impacts of terrestrial, aquatic, and atmospheric carbon fluxes. ispartof: BIOLOGICAL REVIEWS vol:97 issue:6 pages:2023-2038 ispartof: location:England status: published

Aims Biomass allocation to different organs is a fundamental plant ecophysiological process to better respond to changing environments; yet, it remains poorly understood how patterns of biomass allocation respond to nitrogen (N) additions across terrestrial ecosystems worldwide. Methods We conducted a meta-analysis using 5474 pairwise observations from 333 articles to assess how N addition affected plant biomass and biomass allocation among different organs. We also tested the ‘ratio-based optimal partitioning’ vs. the ‘isometric allocation’ hypotheses to explain potential N addition effects on biomass allocation. Important Findings We found that (i) N addition significantly increased whole plant biomass and the biomass of different organs, but decreased root:shoot ratio (RS) and root mass fraction (RMF) while no effects of N addition on leaf mass fraction and stem mass fraction at the global scale; (ii) the effects of N addition on ratio-based biomass allocation were mediated by individual or interactive effects of moderator variables such as experimental conditions, plant functional types, latitudes and rates of N addition and (iii) N addition did not affect allometric relationships among different organs, suggesting that decreases in RS and RMF may result from isometric allocation patterns following increases in whole plant biomass. Despite alteration of ratio-based biomass allocation between root and shoot by N addition, the unaffected allometric scaling relationships among different organs (including root vs. shoot) suggest that plant biomass allocation patterns are more appropriately explained by the isometric allocation hypothesis rather than the optimal partitioning hypothesis. Our findings contribute to better understand N-induced effects on allometric relationships of terrestrial plants, and suggest that these ecophysiological responses should be incorporated into models that aim to predict how terrestrial ecosystems may respond to enhanced N deposition under future global change scenarios.

Plant litter decomposition is one of the most important ecosystem processes influencing elemental biogeochemistry across terrestrial and aquatic environments. Calcium (Ca), magnesium (Mg), and manganese (Mn) are essential nutrients, which play a key role in plant energy metabolism, photosynthesis, and membrane transport of plants, and the major source of these nutrients is litter decomposition. Although litter decomposition has been subject to extensive research over the last several decades, the dynamics of Ca, Mg, and Mn in decomposing litter has been poorly assessed. Also, very few decomposition studies have been simultaneously carried out across aquatic and terrestrial ecosystems. Here, we conducted a two-year litterbag experiment across alpine forest aquatic (stream and riparian zone) and terrestrial (forest floor) ecosystems to address the dynamics of Ca, Mg, and Mn during litter decomposition. We found that (1) in general concentrations of Ca, Mg, and Mn all increased (that is, high immobilization) during the period of decomposition; (2) Ca and Mn immobilized in decomposing litter regardless of ecosystem type or litter species identity, while Mg showed release (mineralization) or immobilization depending on litter species identity; (3) a common hierarchy of drivers for Ca, Mg, or Mn release rate was found across all ecosystem types; and (4) Ca, Mg, and Mn concentrations all showed positive relationships with litter decomposition. Our results provide evidence of general patterns of Ca, Mg, and Mn dynamics in decomposing litter and significant positive effects of these nutrients on litter decomposition across terrestrial and aquatic environments associated with a forest ecosystem. We suggest that addressing nutrient dynamics across contrasting environments is crucial for predicting how biogeochemical cycling of different nutrients is modulated by litter mass decomposition.

There is growing international interest in better managing soils to increase soil organic carbon (SOC) content to contribute to climate change mitigation, to enhance resilience to climate change and to underpin food security, through initiatives such as international ‘4p1000’ initiative and the FAO's Global assessment of SOC sequestration potential (GSOCseq) programme. Since SOC content of soils cannot be easily measured, a key barrier to implementing programmes to increase SOC at large scale, is the need for credible and reliable measurement/monitoring, reporting and verification (MRV) platforms, both for national reporting and for emissions trading. Without such platforms, investments could be considered risky. In this paper, we review methods and challenges of measuring SOC change directly in soils, before examining some recent novel developments that show promise for quantifying SOC. We describe how repeat soil surveys are used to estimate changes in SOC over time, and how long‐term experiments and space‐for‐time substitution sites can serve as sources of knowledge and can be used to test models, and as potential benchmark sites in global frameworks to estimate SOC change. We briefly consider models that can be used to simulate and project change in SOC and examine the MRV platforms for SOC change already in use in various countries/regions. In the final section, we bring together the various components described in this review, to describe a new vision for a global framework for MRV of SOC change, to support national and international initiatives seeking to effect change in the way we manage our soils., Since soil organic carbon (SOC) changes are difficult to measure, a key barrier to implementing programmes to increase SOC is the need for credible and reliable measurement/monitoring, reporting and verification (MRV) platforms. We review methods for measuring SOC change directly in soils, examine novel developments for quantifying SOC change and describe how surveys, long‐term experiments and chronosequences can be used for testing models and as benchmark sites in global frameworks to estimate SOC change. We review MRV platforms for soil organic carbon change already in use and describe a new vision for a global framework for MRV of SOC change.

Grassland management intensification can greatly influence nitrogen (N) dynamics between aboveground and belowground compartments mainly because of the large amount of available N forms, which are repeatedly added to soils. A better understanding of how chronic fertilisation might affect N use efficiency (NUE) in plants can contribute to reducing N losses from soils and improve the sustainability of managed grasslands. Here we address how NUE might be affected by (1) the addition of key nutrients (e.g. N, P, K, Mg) in different combinations, (2) grazing by rabbits, and (3) liming (i.e. CaCO3 applications) in a 22-year-old permanent grassland experiment established in Berkshire, UK, in 1991. We first calculate seven different NUE indexes, which are known to respond either to changes in soil N availability (i.e. endogenous N inputs from soil N mineralization processes) or to exogenous N inputs (i.e. synthetic N fertiliser). We found that plant NUE calculated as plant biomass produced per unit of N acquired significantly decreased under the chronic addition of multiple nutrients (NPKMg) and was even lower under N-only applications. Most NUE indexes significantly decreased under grazing but greatly increased under liming applications. We found evidence that NUE indexes, which accounted for endogenous N sources decreased at increased rates of soil N mineralization. Finally, we found no significant relationships between any of the NUE indexes and estimates of soil N losses (Mg N ha−1) or N retention in soils (i.e. units of soil N retained per unit of N added) calculated from changes in net soil N budget over 22 years. Our study carried out on relatively acidic sandy soils suggests how liming applications in combination with low levels of multi-nutrient additions (NPKMg) can significantly improve plant biomass production per unit of N added thus contributing to enhance the sustainability of managed grassland ecosystems.

In this study, we show that composition shifts of ammonia oxidizer communities imposed by a 22-year field fertilization regime could be well retained in both fresh and air-dried soils. The abundance and composition of ammonia-oxidizing bacteria (AOB) and archaea (AOA) were measured in fresh soils, which received no fertilization (CK), chemical fertilization (NPK), and chemical plus organic matter fertilization (NPK/OM) for 22 years. The air-drying treatment of fresh soil was also conducted for pairwise analysis. We found that in fresh soils DGGE fingerprints of AOB showed significant changes under both NPK and NPK/OM treatments when compared with control (CK) and that microbial shift was almost identical in air-dried soils. Long-term nutrient fertilization did not affect AOA communities in either air-dried or fresh soils. Compared to CK treatment, real-time PCR indicated that AOB amoA genes increased significantly in fresh soils of NPK (59-fold) and NPK/OM (48-fold) plots and in air-dried NPK and NPK/OM soils by 22-fold and 19-fold respectively. Our results demonstrate that community shifts of AOB in fresh soils under chronic N fertilization could be well preserved in air-dried soils, despite the apparent decline in absolute abundance of ammonia oxidizers. These results suggest that air-dried soil could be a useful resource for deciphering the adaptive strategy of ammonia oxidizers under N enrichment when the significant changes of community composition occurred in fresh soils.

Soil organic carbon (SOC) sequestration across agroecosystems worldwide can contribute to mitigate the effects of climate change by reducing levels of atmospheric CO2. Stabilisation of organic carbon (OC) in the fine soil fraction (< 20 µm) is considered an important long-term store of SOC, and the saturation deficit (difference between measured OC and estimated maximum OC in the fine fraction) is frequently used to assess SOC sequestration potential following the linear regression equation developed by Hassink (1997). However, this approach is often taken without any assessment of the fit of the equation to the soils being studied. The statistical limitations of linear regression have previously been noted, giving rise to the proposed use of boundary line (BL) analysis and quantile regression (QR) to provide more robust estimates of maximum SOC stabilisation. The objectives of this work were to assess the suitability of the Hassink (1997) equation to estimate maximum fine-fraction OC in UK grassland soils of varying sward ages and to evaluate the linear regression, boundary line and quantile regression methods to estimate maximum fine-fraction OC. A chronosequence of 10 grasslands was sampled, in order to assess the relationship between sward age (time since the last reseeding event) and the measured and predicted maximum fine-fraction OC. Significantly different regression equations show that the Hassink (1997) equation does not accurately reflect maximum fine-fraction OC in UK grasslands when determined using the proportion of the fine soil fraction (< 20 µm, %) and measured fine-fraction OC (g C per kg soil). The QR estimate of maximum SOC stabilisation was almost double that of the linear regression and BL analysis (0.89 ± 0.074, 0.43 ± 0.017 and 0.57 ± 0.052 g C per kg soil, respectively). Sward age had an inconsistent effect on the measured variables and potential maximum fine-fraction OC. Fine-fraction OC across the grasslands made up 4.5 % to 55.9 % of total SOC, implying that there may be either high potential for additional C sequestration in the fine fraction of these soils or that protection in aggregates is predominant in these grassland soils. This work highlights the need to ensure that methods used to predict maximum fine-fraction OC reflect the soil in situ, resulting in more accurate assessments of carbon sequestration potential.

The role of soils in the global carbon cycle and in reducing GHG emissions from agriculture has been increasingly acknowledged. The '4 per 1000' (4p1000) initiative has become a prominent action plan for climate change mitigation and achieve food security through an annual increase in soil organic carbon (SOC) stocks by 0.4%, (i.e. 4 parts per thousand per year). However, the feasibility of the 4p1000 scenario and, more generally, the capacity of individual countries to implement soil carbon sequestration (SCS) measures remain highly uncertain. Here, we evaluated country-specific SCS potentials of agricultural land for 24 countries in Europe. Based on a detailed survey of available literature, we estimate that between 0.1% and 27% of the agricultural greenhouse gas (GHG) emissions can potentially be compensated by SCS annually within the next decades. Measures varied widely across countries, indicating differences in country-specific environmental conditions and agricultural practices. None of the countries' SCS potential reached the aspirational goal of the 4p1000 initiative, suggesting that in order to achieve this goal, a wider range of measures and implementation pathways need to be explored. Yet, SCS potentials exceeded those from previous pan-European modelling scenarios, underpinning the general need to include national/regional knowledge and expertise to improve estimates of SCS potentials. The complexity of the chosen SCS measurement approaches between countries ranked from tier 1 to tier 3 and included the effect of different controlling factors, suggesting that methodological improvements and standardization of SCS accounting are urgently required. Standardization should include the assessment of key controlling factors such as realistic areas, technical and practical feasibility, trade-offs with other GHG and climate change. Our analysis suggests that country-specific knowledge and SCS estimates together with improved data sharing and harmonization are crucial to better quantify the role of soils in offsetting anthropogenic GHG emissions at global level.

Soil organic carbon (SOC) sequestration across agroecosystems worldwide can contribute to mitigate the effects of climate change by reducing levels of atmospheric CO2. Mineral associated organic carbon (MAOC) is considered an important long-term store of SOC and the saturation deficit (difference between measured MAOC and estimated maximum MAOC) is frequently used to assess SOC sequestration potential following the linear regression equation developed by Hassink (1997). However, this approach is often taken without any assessment of the fit of the equation to the soils being studied. The statistical limitations of linear regression have previously been noted, giving rise to the proposed use of boundary line (BL) analysis and quantile regression (QR) to provide more robust estimates of maximum SOC stabilisation. The objectives of this work were to assess the suitability of the Hassink (1997) equation to estimate maximum MAOC in UK grassland soils of varying sward ages and to evaluate the linear regression, BL and QR methods to estimate maximum MAOC. A chronosequence of 10 grasslands was sampled, in order to assess the relationship between sward age (time since last reseeding event) and current and predicted maximum MAOC. Significantly different regression equations show that the Hassink (1997) equation does not accurately reflect maximum MAOC in UK grasslands when determined using the proportion of fine soil fraction and current MAOC. The QR estimate of maximum SOC stabilisation was almost double that of linear regression and BL analysis (0.89 ± 0.074, 0.43 ± 0.017 and 0.57 ± 0.052 g C kg−1 soil, respectively). Sward age had an inconsistent effect on the measured variables and potential maximum MAOC. MAOC across the grasslands made up 4.5 to 55.9 % of total SOC, implying that there may be either high potential for additional C sequestration in the mineral fraction of these soils, or stabilisation in aggregates is predominant in these grassland soils. This work highlights the need to ensure that methods used to predict maximum MAOC reflect the soil in situ, resulting in more accurate assessments of carbon sequestration potential.

Our knowledge of root mass decomposition processes has greatly benefited from studies carried out in natural and semi‐natural ecosystems. It is less known, however, how root mass loss might occur in intensively managed ecosystems, such as agricultural grasslands, which receive chronic additions of nitrogen (N) fertilizers. Key questions remain about (1) how repeated N applications either in organic or inorganic forms might influence fine root mass decomposition, and (2) what implications root mass loss might have for soil carbon (C) storage. Here, we use a long‐term grassland experiment established in 1970 at Hillsborough, Northern Ireland (UK), to address how repeated additions of organic (i.e., liquid manures) and inorganic (i.e., NPK) fertilizers over >45 yr might influence fine root mass decomposition and how. We found that after a 10‐month incubation period under field conditions, highest root mass loss (~43%) was associated with highest additions of organic slurries, whereas lowest root mass loss (~29%) occurred in inorganically N‐fertilized and unfertilized soils. Root decomposition was significantly and negatively related to root C:N ratios and fungal‐to‐bacterial (F:B) ratios. Root decomposition, however, was not related to changes in soil C stocks or rates of soil C accumulation. N applications either maintained or increased (up to three times) soil C accumulation compared to unfertilized soils after five decades of intensive management. Our findings suggest that in the short‐term organic fertilization can enhance root detritus decomposition (thus promoting N release in soils for plant uptake) while contributing in the long‐term to soil C accumulation either through extra C inputs from manures or through N‐induced effects on microbial activity or both. Further studies need to address how this dual effect of organic fertilization could contribute making grassland intensification more sustainable by reducing the use of artificial NPK fertilizers.

Multi-functional silvopastoral systems provide a wide range of services to human society including the regulation of nutrients and water in soils and the sequestration of atmospheric carbon dioxide (CO2). Although silvopastoral systems significantly contribute to enhance aboveground carbon (C) sequestration (e.g. C accumulation in woody plant biomass), their long-term effects on soil C pools are less clear. In this study we performed soil physical fractionation analyses to quantify the C pool of different aggregate fractions across three land use types including (1) silvopastoral system with ash trees (Fraxinus excelsior L.), (2) planted woodland with ash trees, and (3) permanent grassland, which were established in 1989 at Loughgall, Northern Ireland, UK. Our results show that 26 years after the conversion of permanent grassland to either silvopastoral or woodland systems, soil C (and N) stocks (0–20 cm depth) did not significantly change between the three land use types. We found, however, that permanent grassland soils were associated with significantly higher C pools (g C kg−1 soil; P 2 mm) whereas soil C pools of the micro-aggregate (53–250 μm) and silt and clay (

Summary Recent studies have revealed many potential benefits of increasing plant diversity in natural ecosystems, as well as in agroecosystems and production forests. Plant diversity potentially provides a partial to complete substitute for many costly agricultural inputs, such as fertilizers, pesticides, imported pollinators and irrigation. Diversification strategies include enhancing crop genetic diversity, mixed plantings, rotating crops, agroforestry and diversifying landscapes surrounding croplands. Here we briefly review studies considering how increasing plant diversity influences the production of crops, forage, and wood, yield stability, and several regulating and supporting agroecosystem services. We also discuss challenges and recommendations for diversifying agroecosystems. There is consistently strong evidence that strategically increasing plant diversity increases crop and forage yield, wood production, yield stability, pollinators, weed suppression and pest suppression, whereas effects of diversification on soil nutrients and carbon remain poorly understood. Synthesis. The benefits of diversifying agroecosystems are expected to be greatest where the aims are to sustainably intensify production while reducing conventional inputs or to optimize both yields and ecosystem services. Over the next few decades, as monoculture yields continue to decelerate or decline for many crops, and as demand for ecosystem services continues to rise, diversification could become an essential tool for sustaining production and ecosystem services in croplands, rangelands and production forests.

The interactive effects of multiple global change drivers on terrestrial carbon (C) storage remain poorly understood. Here, we synthesise data from 633 published studies to show how the interactive effects of multiple drivers are generally additive (i.e. not differing from the sum of their individual effects) rather than synergistic or antagonistic. We further show that (1) elevated CO2 , warming, N addition, P addition and increased rainfall, all exerted positive individual effects on plant C pools at both single-plant and plant-community levels; (2) plant C pool responses to individual or combined effects of multiple drivers are seldom scale-dependent (i.e. not differing from single-plant to plant-community levels) and (3) soil and microbial biomass C pools are significantly less sensitive than plant C pools to individual or combined effects. We provide a quantitative basis for integrating additive effects of multiple global change drivers into future assessments of the C storage ability of terrestrial ecosystems.

Long-term nutrient fertilization of grassland soils greatly increases plant yields but also profoundly alters ecosystem phosphorus (P) dynamics. Here, we addressed how long-term P fertilization may affect ecosystem P budget, P use efficiency (PUE) and the abundance of arbuscular mycorrhizal fungi (AMF), which play a key role in the acquisition of P by plants. We found that 47 years of organic P applications increased soil P availability and total soil P stocks up to 1600% and 400%, respectively, compared to unfertilized-control soils. Grassland soils could retain up to 62% and 48% of P applied since 1970 in organic and inorganic forms, respectively. Nutrient treatments significantly affected rates of AMF root colonization (%), which were higher in control and NPK-fertilized plots when compared to soils receiving increasing applications of organic P. Plant PUE increased with greater AMF root colonization, which remained high (i.e. 50-to-75%) even after ~50 years of continuous ‘normal’ rates of agronomic P inputs (~30 kg P ha−1 year−1). AMF abundance, however, decreased under higher P applications and we found a negative relationship between soil P availability or soil P stocks and rates of AMF root colonization. Our study demonstrates that (1) AMF root colonization is still high in soils, which have received consistent but moderate P inputs for over four decades, and (2) moderate rates of P fertilization are related to a more conservative P ecosystem budget whereby the amount of P retained in soils and up-taken by plants on an annual basis is higher than the amount of P added through fertilization. This is possible only if extra P is ‘mined’ from the soil P ‘bank’ and made available to plant uptake. We suggest that AMF could play a significant role in intensively-managed grasslands contributing to increase P sustainability by reducing the need for extra P fertilizer.

Aim We sought to understand how the individual and combined effects of multiple environmental change drivers differentially influence terrestrial nitrogen (N) concentrations and N pools and whether the interactive effects of these drivers are mainly antagonistic, synergistic or additive. Location Worldwide. Time period Contemporary. Major taxa studied Plants, soil, and soil microbes in terrestrial ecosystems. Methods We synthesized data from manipulative field studies from 758 published articles to estimate the individual, combined and interactive effects of key environmental change drivers (elevated CO2, warming, N addition, phosphorus addition, increased rainfall and drought) on plant, soil, and soil microbe N concentrations and pools using meta‐analyses. We assessed the influences of moderator variables on these effects through structural equation modelling. Results We found that (a) N concentrations and N pools were significantly affected by the individual and combined effects of multiple drivers, with N addition (either alone or in combination with another driver) showing the strongest positive effects; (b) the individual and combined effects of these drivers differed significantly between N concentrations and N pools in plants, but seldom in soils and microbes; (c) additive effects of driver pairs on N concentrations and pools were much more common than synergistic or antagonistic effects across plants, soils and microbes; and (d) environmental and experimental factors were important moderators of the individual, combined and interactive effects of these drivers on terrestrial N. Main conclusions Our results indicate that terrestrial N concentrations and N pools, especially those of plants, can be significantly affected by the individual and combined effects of environmental change drivers, with the interactive effects of these drivers being mostly additive. Our findings are important because they contribute to the development of models to better predict how altered N availability affects ecosystem carbon cycling under future environmental changes. ispartof: GLOBAL ECOLOGY AND BIOGEOGRAPHY vol:28 issue:5 pages:1-35 status: Published online

Summary The productivity of species-diverse plant assemblages strongly depends on the temporal dynamics of nutrient uptake by competing neighbouring plants. Our understanding, however, of how rates of nitrogen (N), phosphorous (P) and potassium (K) uptake might change through time between neighbouring plant species under field conditions is still very limited. Here, we specifically measure the temporal trajectories of N, P and K uptake by staple food plants such as wheat (Triticum aestivum L.), barley (Hordeum vulgare L.) and maize (Zea mays L.) when growing either in monocultures or in intercropping systems. We ask how (i) plant species combinations, (ii) N fertilization and (iii) film mulching might affect key indexes of N, P and K uptake over time. We fit logistic models to characterize the nutrient uptake trajectories. Maximum cumulative N, P and K uptake (kg ha−1) by wheat and barley were significantly greater in wheat–maize or barley–maize intercropping systems than in wheat or barley monocultures. Cumulative nutrient uptake by intercropped maize (either with wheat or with barley) was reduced by interspecific competition at early growth stages, but it increased rapidly after wheat and barley were harvested. Maximum cumulative N and P (but not K) uptake by intercropped maize were significantly higher than the uptake by monoculture maize, particularly when N fertilizer or film mulching was applied. Intercropping induced a significant temporal niche differentiation in maximum daily nutrient uptake rates (kg ha−1 day−1) between intercropped species. Fertilization had much stronger effects on maximum cumulative nutrient uptake of maize than that of wheat and barley. Mulching significantly increased the maximum cumulative nutrient uptake of maize and advanced the time to reach its maximum daily P and K uptake rates. Our study provides evidence of an important temporal niche differentiation mechanism (‘temporal complementarity’) in nutrient uptake rates between neighbouring plant species. A better understanding of temporal trajectories of interspecific nutrient uptake rates remains crucial if we want to maximize the nutrient-use efficiency and sustain overyielding (i.e. high food production) in plant species-diverse systems such as intercropping.

Important biochemical reactions in soils are catalyzed by extracellular enzymes, which are synthesized by microbes and plant roots. Although enzyme activities can significantly affect the decomposition of soil organic matter and thus influence the storage and cycling of carbon (C) and nitrogen (N), it is not clear how enzyme activities relate to changes in the C and N content of different grassland soils. Here we address whether the activity of C-acquiring (b-1,4-glucosidase, BG) and N-acquiring (L-leucine amino-peptidase (LAP) and b-1,4-N-acetyl-glucosaminidase (NAG)) enzymes is linked to changes in the C and N content of a variety of human-managed grassland soils. We selected soils which have a well-documented management history going back at least 19 years in relation to changes in land use (grazing, mowing, ploughing), nutrient fertilization and lime (CaCO 3) applications. Overall we found a positive relationship between BG activity and soil C content as well as between LAP þ NAG activity and soil N. These positive relationships occurred across grasslands with very different soil pH and management history but not in intensively managed grasslands where increases in soil bulk density (i.e. high soil compaction) negatively affected enzyme activity. We also found evidence that chronic nutrient fertilization contributed to increases in soil C content and this was associated with a significant increase in BG activity when compared to unfertilized soils. Our study suggests that while the activities of C-and N-acquiring soil enzymes are positively related to soil C and N content, these activities respond significantly to changes in management (i.e. soil compaction and nutrient fertilization). In particular, the link between BG activity and the C content of long-term fertilized soils deserves further investigation if we wish to improve our understanding of the C sequestration potential of human-managed grassland soils.

Long-term changes in soil organic carbon (SOC) are difficult to quantify experimentally because of measurement errors and high spatial and temporal variability. Modelling can help to provide a more robust assessment by reducing these uncertainties and reproducing greenhouse gas (GHG) and C exchange processes in an ecosystem by identifying key drivers. In this study, the Denitrification-Decomposition (DNDC95) model was used to evaluate SOC density (SOCρ) and its annual changes (ΔSOCρ) in temperate grassland soils, which received different forms of nitrogen (N) (i.e. inorganic and organic) and at different application rates for 45 years. We found that simulated values for SOCρ (0–15 cm depth) in unfertilized (54 t C ha−1) and fertilized soils (55 t C ha−1) were lower than measured values (73 and 77 t C ha−1, respectively). Despite some variations, measured and simulated SOCρ was higher under cattle (88–99 vs. 66–116 t C ha−1) than pig slurry (75–78 vs. 55–69) applications, and increased with increasing rates of added C. Irrespective of nutrient treatment, overall mean sequestration rates were 0.46 ± 0.06 (observed) and 0.37 ± 0.01 (simulated) t C ha−1 yr−1. Simulated values explained 66% of the variability between years and treatments (slope: 1.41; intercept: −34.58 t C ha−1) with reasonably good prediction efficiency The variations in simulated values could be explained by differences in applied N (63%), which were linked to differences in C (62%), rainfall (15%) and air temperature (11%). The model (R2 0.77–0.99/-0.99; p

Grasslands cover more than a third of the European agricultural area and are often intensively managed to support the livestock-farming sector. Despite grassland intensification can greatly influence soil carbon (C) cycling, changes in soil C stocks both across years and at increasing soil depths remain difficult to quantify simultaneously. Here we measured spatial and temporal changes in soil C stocks to assess C stocks’ response to intensive grassland management. We measured soil C stocks (a) in a long-term nutrient fertilization experiment on permanent grassland (established in 1970), and (b) in 126 grassland fields distributed across 11 lowland farms in Northern Ireland (UK), which are associated with different frequencies of soil tillage. Using 45 years of data from the plot-scale grassland experiment we found evidence that significant changes in soil C stocks mainly occurred in the soil top 20 cm (not in deeper soils) and only between ‘extreme’ nutrient treatments (i.e. unfertilized vs. highly fertilized soils). Soil physical fractionation, radiocarbon and stable nitrogen isotope analyses all suggest that new C has accumulated in these soils but perhaps not fast enough to affect C stocks in deeper soil layers. Results at the field-scale from intensively managed grasslands show how the frequency of soil tillage neither affected soil C stocks between 0–20, 20–40 or 40–60 cm depth layers nor the C pool of different soil physical fractions at increasing soil depths. Our findings demonstrate how the response of soil C stocks to grassland intensification (i.e. C stocks response to different rates of nutrient fertilization or frequency of soil tillage) can be very slow both in time and space under cool and humid climate conditions. We suggest how the persistence of these soil C stocks under intensive grassland management offers the unique opportunity to improve nutrient use efficiency and cycling thus promoting the delivery of multiple ecosystem processes.

Sustainable grassland intensification aims to increase plant yields while maintaining the ability of soil to act as a sink rather than sources of atmospheric CO2. High biomass yields from managed grasslands, however, can be only maintained through long-term nutrient fertilization, which can significantly affect soil carbon (C) storage and cycling. Key questions remain about (1) how long-term inorganic vs. organic fertilization influences soil C stocks, and (2) how soil C gains (or losses) contribute to the long-term C balance of managed grasslands. Using 43 years of data from a permanent grassland experiment, we show that soils not only act as significant C sinks but have not yet reached C saturation. Even unfertilized control soils showed C sequestration rates of 0.35 Mg C ha−1 yr−1 (i.e. 35 g C m−2 yr−1; 0–15 cm depth) between 1970 and 2013. High application rates of liquid manure (i.e. cattle slurry) further increased soil C sequestration to 0.86 Mg C ha−1 yr−1 (i.e. 86 g C m−2 yr−1) and a key cause of this C accrual was greater C inputs from cattle slurry. However, average coefficients of slurry-C retention in soils suggest that 85 % of C added yearly through liquid manure is lost possibly via CO2 fluxes and organic C leaching. Inorganically fertilized soils (i.e. NPK) had the lowest C-gain efficiency (i.e. unit of C gained per unit of N added) and lowest C sequestration (similar to control soils). Soils receiving cattle slurry showed higher C-gain and N-retention efficiencies compared to soils receiving NPK or pig slurry. We estimate that net rates of CO2-sequestration in the top 15 cm of the soil can offset 9–25 % of GHG (greenhouse gas) emissions from intensive management. However, because of multiple GHG sources associated with livestock farming, the net C balance of these grasslands remains positive (9–12 Mg CO2-equivalent ha−1 yr−1), thus contributing to climate change. Further C-gain efficiencies (e.g. reduced enteric fermentation and use of feed concentrates, better nutrient management) are required to make grassland intensification more sustainable.

International audience; AbstractThe delivery of multiple ecosystem services from intensively managed cropping systems remains challenging mainly because increases in crop yields are rarely associated with greater soil carbon (C) sequestration or efficient ecosystem nitrogen (N) cycling. The sustainability of intensive cropping systems depends on trade-offs between multiple ecosystem services but experimental evidence from long-term field studies remains limited. The common expectation is that highly productive agroecosystems will be associated with lower soil C and reduced nutrient use efficiency. Here we use data spanning 30 years of a long-term field experiment established in 1985 near Lodi in Northern Italy where four arable systems (i.e., three crop rotations and a grain maize monoculture) and a no-till permanent meadow were all compared across two levels of agronomic inputs. We asked how (i) soil C stocks, (ii) fertilizer N-use efficiency, (iii) productivity, (iv) crude protein, and (v) feed units for lactation might differ across the five traditional cropping systems. We found that soil C sequestration, N-use efficiency, and crude protein production were all significantly higher in the no-till permanent meadow, which was however associated with lower dry matter yields and reduced feed units for lactation when compared with the arable systems. These findings suggest that converting arable soil into no-till permanent meadow is a win-lose solution. The most productive annual rotation (silage maize + Italian ryegrass) offers a complementary win-lose solution whereby high yields and feed units for lactation can be obtained from a smaller cropland area. The other cropping systems are less productive and provide little (or none) soil C storage benefits. Our study is one of the first to show clear trade-offs between multiple ecosystem services and to demonstrate that high fertilization rates might be only justified when the management goal is to spare land for less intensive uses such as permanent meadow.

The functioning of human-managed grassland ecosystems strongly depends on how common management practices will affect grassland ‘belowground compartment’ including soil biogeochemistry and plant roots. Key questions remain about how animal grazing, liming (e.g. the addition of CaCO3 to soils), and nutrient fertilization might affect, in the long-term, soil nutrient cycling and multiple root traits. Here we focus on a mesotrophic grassland located in Berkshire, UK, where contrasting levels of rabbit grazing, liming and different inorganic fertilizers have been applied since 1991. We ask how (1) soil nitrogen (N) availability and cycling, (2) total root mass, (3) root mass decomposition, and (4) arbuscular mycorrhizal fungal (AMF) root colonisation might respond to 22 years of very different management. We found that liming strongly affected total root mass, root decomposition, root AMF colonisation as well as soil N availability and cycling and that these effects were mainly driven by liming-induced increases in soil pH. Increases in soil pH were associated with significant (1) decreases in root mass, (2) increases in root mass decomposability and in the mineralization of N in decomposing root detritus, and (3) increases in AMF infection. Soil pH was also significantly related to greater N availability (i.e. soil NO3 levels) and to lower δ15N natural abundance, which suggests more efficient N uptake by plants in limed soils as we found in our study. The application of multiple nutrients (NPKMg) also reduced total root mass, while N-only fertilization was associated with greater AMF infection. Surprisingly the long-term impact of grazing was generally weak and not significant on most plant and soil parameters. Despite soil pH affecting most belowground variables, changes in soil pH were not associated with any change in soil C and N stocks. Because liming can improve nutrient cycling (and benefits soil pH and grass yields) without negatively affecting soil C sequestration, we suggest that regular liming applications may provide management solutions for increasing the long-term sustainability of permanent grassland. This article is protected by copyright. All rights reserved.

The relationship between plant diversity and productivity in grasslands could depend, partly, on how diversity affects vertical distributions of root biomass in soil; yet, no prior study has evaluated the links among diversity, root depth distributions, and productivity in a long-term experiment. We use data from a 12-year experiment to ask how plant species richness and composition influenced both observed and expected root depth distributions of plant communities. Expected root depth distributions were based on the abundance of species in each community and two traits of species that were measured in monocultures: root depth distributions and root to shoot ratios. The observed proportion of deep root biomass increased more than expected with species richness and was positively correlated with aboveground productivity. Indeed, the proportion of deep root biomass explained variation in productivity even after accounting for legume presence/abundance, and greater nitrogen availability in diverse plots. Diverse plots had root depth distributions that were twice as deep as expected from their species composition and corresponding monoculture traits, partly due to interactions between C4 grasses and legumes. These results suggest the productivity of diverse plant communities was partly dependent on belowground plant interactions that caused roots to be distributed more deeply in soil.

Human-induced increases in nitrogen (N) deposition are common across many terrestrial ecosystems worldwide. Greater N availability not only reduces biological diversity, but also affects the biogeochemical coupling of carbon (C) and N cycles in soil ecosystems. Soils are the largest active terrestrial C pool and N deposition effects on soil C sequestration or release could have global importance. Here, we show that 27 years of chronic N additions to prairie grasslands increased C sequestration in mineral soils and that a potential mechanism responsible for this C accrual was an N-induced increase in root mass. Greater soil C sequestration followed a dramatic shift in plant community composition from native-species-rich C4 grasslands to naturalized-species-rich C3 grasslands, which, despite lower soil C gains per unit of N added, still acted as soil C sinks. Since both high plant diversity and elevated N deposition may increase soil C sequestration, but N deposition also decreases plant diversity, more research is needed to address the long-term implications for soil C storage of these two factors. Finally, because exotic C3 grasses often come to dominate N-enriched grasslands, it is important to determine if such N-dependent soil C sequestration occurs across C3 grasslands in other regions worldwide.

Chronic nitrogen (N) fertilization can greatly affect soil carbon (C) sequestration by altering biochemical interactions between plant detritus and soil microbes. In lignin-rich forest soils, chronic N additions tend to increase soil C content partly by decreasing the activity of lignin-degrading enzymes. In cellulose-rich grassland soils it is not clear whether cellulose-degrading enzymes are also inhibited by N additions and what consequences this might have on changes in soil C content. Here we address whether chronic N fertilization has affected (1) the C content of light versus heavier soil fractions, and (2) the activity of four extracellular enzymes including the C-acquiring enzyme β-1,4-glucosidase (BG; necessary for cellulose hydrolysis). We found that 19 years of chronic N-only addition to permanent grassland have significantly increased soil C sequestration in heavy but not in light soil density fractions, and this C accrual was associated with a significant increase (and not decrease) of BG activity. Chronic N fertilization may increase BG activity because greater N availability reduces root C:N ratios thus increasing microbial demand for C, which is met by C inputs from enhanced root C pools in N-only fertilized soils. However, BG activity and total root mass strongly decreased in high pH soils under the application of lime (i.e. CaCO3), which reduced the ability of these organo-mineral soils to gain more C per units of N added. Our study is the first to show a potential ‘enzyme link’ between (1) long-term additions of inorganic N to grassland soils, and (2) the greater C content of organo-mineral soil fractions. Our new hypothesis is that the ‘enzyme link’ occurs because (a) BG activity is stimulated by increased microbial C demand relative to N under chronic fertilization, and (b) increased BG activity causes more C from roots and from microbial metabolites to accumulate and stabilize into organo-mineral C fractions. We suggest that any combination of management practices that can influence the BG ‘enzyme link’ will have far reaching implications for long-term C sequestration in grassland soils.

Despite the topic of soil nitrogen (N) mineralization being well-studied, very few studies have addressed the relative contribution of different plant and soil variables in influencing soil N mineralization rates, and thus the supply of inorganic N to plants. Here, we used data from a well-studied N-limited grassland to address the relative effects of six plant and soil variables on net and on gross rates of soil N mineralization. We also addressed whether plant effects on soil N mineralization were mediated by changes in C and N concentrations of multiple soil organic matter (SOM) fractions. Regression analyses show that key plant traits (i.e., plant C:N ratios and total root mass) were more important than total C and N concentrations of bulk soil in influencing N mineralization. This was mainly because plant traits influenced the C and N concentration (and C:N ratios) of different SOM fractions, which in turn were significantly associated with changes in net and gross N mineralization. In particular, C:N ratios of a labile soil fraction were negatively related to net soil N mineralization rates, whereas total soil C and N concentrations of more recalcitrant fractions were positively related to gross N mineralization. Our study suggests that changes in belowground N-cycling can be better predicted by simultaneously addressing how plant C:N ratios and root mass affect the composition and distribution of different SOM pools in N-limited grassland systems.

The application of calcium- and magnesium-rich materials to soil, known as liming, has long been a foundation of many agro-ecosystems worldwide because of its role in counteracting soil acidity. Although liming contributes to increased rates of respiration from soil thereby potentially reducing soils ability to act as a CO2 sink, the long-term effects of liming on soil organic carbon (Corg) sequestration are largely unknown. Here, using data spanning 129 years of the Park Grass Experiment at Rothamsted (UK), we show net Corg sequestration measured in the 0–23cm layer at different time intervals since 1876 was 2–20 times greater in limed than in unlimed soils. The main cause of this large Corg accrual was greater biological activity in limed soils, which despite increasing soil respiration rates, led to plant C inputs being processed and incorporated into resistant soil organo-mineral pools. Limed organo-mineral soils showed: (1) greater Corg content for similar plant productivity levels (i.e. hay yields); (2) higher 14C incorporation after 1950s atomic bomb testing and (3) lower C:N ratios than unlimed organo-mineral soils, which also indicate higher microbial processing of plant C. Our results show that greater Corg sequestration in limed soils strongly reduced the global warming potential of long-term liming to permanent grassland suggesting the net contribution of agricultural liming to global warming could be lower than previously estimated. Our study demonstrates that liming might prove to be an effective mitigation strategy, especially because liming applications can be associated with a reduced use of nitrogen fertilizer which is a key cause for increased greenhouse gas emissions from agro-ecosystems.

In a 13-year grassland biodiversity experiment in Minnesota, USA, we addressed two main questions: What set of ecological mechanisms caused aboveground productivity to become approximately 340% greater in highly diverse plant mixtures than in the average monoculture? Why did the effect of diversity on productivity become so much stronger through time? Because our grassland system is N limited, we simultaneously measured critical variables associated with the storage and cycling of this element, such as plant and soil N pools, soil N availability, soil N mineralization rates, and plant N-use efficiency, as well as the initial soil N concentration of each diversity plot when the experiment was established in 1994. We used linear and multiple regression analyses to test for potential effects of these variables on aboveground productivity and to address whether and how such variables were in turn affected by plant species diversity and functional composition across years and also at different time intervals within the same year. We found that seven variables simultaneously controlled productivity: (1) initial total soil nitrogen (N) of each plot, (2) diversity-dependent increases in total soil N through time, (3) soil N mineralization rates, (4) soil nitrate (NO3-) utilization, (5) increases in plant N-use efficiency at greater plant diversity, (6) legume presence, and (7) higher species numbers. The surprising continued significance of higher plant diversity may occur because of its effects on seasonal capture of soil NO3- and moisture and on the accumulation of root-N pools, all of which may have also increased productivity through time at higher species numbers.

Summary 1. Plant functional composition may indirectly affect fine root processes both qualitatively (e.g. by influencing root chemistry) and quantitatively (e.g. by influencing root biomass and thus soil carbon (C) inputs and the soil environment). Despite the potential implications for ecosystem nitrogen (N) cycling, few studies have addressed the linkages between plant functional composition, root decay, root detritus N dynamics and soil N mineralization rates. 2. Here, using data from a large grassland biodiversity experiment, we first show that plant functional composition affected fine root mass loss, root detritus N dynamics and net soil N mineralization rates through its effects on root chemistry rather than on the environment of decomposition. In particular, the presence of legumes and non-leguminous forbs contributed to greater fine root decomposition which in turn enhanced root N release and net soil N mineralization rates compared with C3 and C4 grasses. 3. Second, we show that all fine roots released N immediately during decomposition and showed very little N immobilization regardless of plant composition. As a consequence, there was no evidence of increased root or soil N immobilization rates with increased below-ground plant biomass (i.e. increased soil C inputs) even though root biomass negatively affected root decay. 4. Our results suggest that fine roots represent an active soil N pool that may sustain plant uptake while other soil N forms are being immobilized in microbial biomass and/or sequestered into soil organic matter. However, fine roots may also represent a source of recalcitrant plant detritus that is returned to the soil (i.e. fine roots of C4 and C3 grasses) and that can contribute to an increase in the soil organic matter pool. 5. Synthesis . An important implication of our study is that the simultaneous presence of different plant functional groups (in plant mixtures) with opposite effects on root mass loss, root N release and soil N mineralization rates may be crucial for sustaining multiple ecosystem services such as productivity and soil C and N sequestration in many N-limited grassland systems.

Most explanations for the positive effect of plant species diversity on productivity have focused on the efficiency of resource use, implicitly assuming that resource supply is constant. To test this assumption, we grew seedlings of Echinacea purpurea in soil collected beneath 10-year-old, experimental plant communities containing one, two, four, eight, or 16 native grassland species. The results of this greenhouse bioassay challenge the assumption of constant resource supply; we found that bioassay seedlings grown in soil collected from experimental communities containing 16 plant species produced 70% more biomass than seedlings grown in soil collected beneath monocultures. This increase was likely attributable to greater soil N availability, which had increased in higher diversity communities over the 10-year-duration of the experiment. In a distinction akin to the selection/complementarity partition commonly made in studies of diversity and productivity, we further determined whether the additive effects of functional groups or the interactive effects of functional groups explained the increase in fertility with diversity. The increase in bioassay seedling biomass with diversity was largely explained by a concomitant increase in N-fixer, C4 grass, forb, and C3 grass biomass with diversity, suggesting that the additive effects of these four functional groups at higher diversity contributed to enhance N availability and retention. Nevertheless, diversity still explained a significant amount of the residual variation in bioassay seedling biomass after functional group biomass was included in a multiple regression, suggesting that interactions also increased fertility in diverse communities. Our results suggest a mechanism, the fertility effect, by which increased plant species diversity may increase community productivity over time by increasing the supply of nutrients via both greater inputs and greater retention.

We studied the composition of a savanna woody plant community across a natural herbivory gradient maintained by both browsing and grazing ungulates in an arid part of the Kruger National Park, South Africa. We focused on (1) short-term browsing effects on reproductive and morphological traits of a dominant-palatable woody species, Acacia nigrescens, Miller, (2) the relationship between browsing–grazing intensity and soil parameters, (3) the effects of herbivore–soil interactions on woody species richness and composition, and (4) browser-induced effects on the representation of woody plant functional traits. We show that the number of pods carried by A. nigrescens trees as well as the internode length of external tree branches both decreased significantly at high browsing intensity. Moreover, we found that total soil nitrogen (N) and soil cation (Ca, Na, Mg, and K) concentrations varied significantly according to grazing rather than browsing intensity with soil nutrients decreasing at heavily grazed sites. Although herbivory and soil

Suppressed growth forms of woody species are common where fire and herbivory are major ecosystem drivers, such as in African savannas. Nevertheless, despite their importance in maintaining plant population viability, woody plants in suppressed growth forms have received little attention in ecological studies. We measured a set of morpho-functional traits and investigated plant density variation in suppressed growth forms (hereafter we refer to them as saplings) of two common species, Acacia nigrescens and Acacia tortilis, across sites that had undergone very different histories of attack from large herbivores while fire had been absent for at least 13 y. We show that heavily browsed saplings have higher regrowth abilities, twice the number of stems produced by the main root crown and twice the root diameter at 5 cm soil depth, than lightly browsed saplings. This suggests that Acacia saplings are resilient to chronic herbivory and show high morphological plasticity. However, we show that mammalian herbivores can strongly limit sapling recruitment to mature size classes and possibly affect variation in sapling density between heavily and lightly browsed sites. Further studies should investigate whether the persistence of the sapling bank can be ascribed to the ``storage effect'' by which a plant's reproductive potential is ``stored'' in a suppressed growth form until a window of opportunity allows rapid maturity.

Summary 1. The mechanisms controlling soil carbon (C) and nitrogen (N) accumulation are crucial for explaining why soils are major terrestrial C sinks. Such mechanisms have been mainly addressed by imposing short-term, step-changes in CO 2 , temperature and N fertilization rates on either monocultures or low-diversity plant assemblages. No studies have addressed the long-term effects of plant functional diversity (i.e. plant functional composition) on rates of soil C accumulation in N-limited grasslands where fixation is the main source of N for plants. 2. Here we measure net soil C and N accumulation to 1 m soil-depth during a 12-year-long grassland biodiversity experiment established on agriculturally degraded soils at Cedar Creek, Minnesota, USA. 3. We show that high-diversity mixtures of perennial grassland plant species stored 500% and 600% more soil C and N than, on average, did monoculture plots of the same species. Moreover, the presence of C4 grasses and legumes increased soil C accumulation by 193% and 522%, respectively. Higher soil C and N accrual resulted both from increased C and N inputs via (i) higher root biomass, and (ii) from greater root biomass accumulation to 60 cm soil depth resulting from the presence of highly complementary functional groups (i.e. C4 grasses and legumes). 4. Our results suggest that the joint presence of C4 grass and legume species is a key cause of greater soil C and N accumulation in both higher and lower diversity plant assemblages. This is because legumes have unique access to N, and C4 grasses take up and use N efficiently, increasing below-ground biomass and thus soil C and N inputs. 5. Synthesis. We demonstrate that plant functional complementarity is a key reason why higher plant diversity leads to greater soil C and N accumulation on agriculturally degraded soils. We suggest the combination of key C4 grass‐legume species may greatly increase ecosystem services such as soil C accumulation and biomass (biofuel) production in both high- and low-diversity N-limited grassland systems.

We investigated the linkages between leaf litter quality and decomposability in a savanna plant community dominated by palatable-spinescent tree species. We measured: (1) leaf litter decomposability across five woody species that differ in leaf chemistry; (2) mass decomposition, nitrogen (N); and carbon (C) dynamics in leaf litter of a staple browse species (Acacia nigrescens) as well as (3) variation in litter composition across six sites that experienced very different histories of attack from large herbivores. All decomposition trials included litter bags filled with chopped straw to control for variation in site effects. We found a positive relationship between litter quality and decomposability, but we also found that Acacia and straw litter mass remaining did not significantly vary between heavily and lightly browsed sites. This is despite the fact that both the quality and composition of litter returned to the soil were significantly different across sites. We observed greater N resorption from senescing Acacia leaves at heavily browsed sites, which in turn contributed to increase the C:N ratio of leaf litter and caused greater litter N immobilization over time. This, together with the significantly lower tree- and herb-leaf litter mass beneath heavily browsed trees, should negatively affect decomposition rates. However, estimated dung and urine N deposition from both browsers and grazers was significantly greater at high- than at low-herbivory sites. We hypothesize that N inputs from dung and urine boost litter N mineralization and decomposition (especially following seasonal rainfall events), and thereby offset the effects of poor leaf litter quality at chronically browsed sites.

We measured browsing-induced responses of Acacia trees to investigate "browsing lawns" as an analogy to grazing lawns in a semiarid eutrophic African savanna. During the two-year field study, we measured plant tolerance, resistance, and phenological traits, while comparing variation in leaf nitrogen and specific leaf area (SLA) across stands of Acacia nigrescens, Miller, that had experienced markedly different histories of attack from large herbivores. Trees in heavily browsed stands developed (1) tolerance traits such as high regrowth abilities in shoots and leaves, high annual branch growth rates, extensive tree branching and evidence of internal N translocation, and (2) resistance traits such as close thorn spacing. However, phenological "escape" responses were weak even in heavily browsed stands. Overall, browsing strongly affected plant morpho-functional traits and decreased both the number of trees carrying pods and the number of pods per tree in heavily browsed stands. Hence, there is experimental evidence that tolerance and resistance traits may occur simultaneously at heavily browsed sites, but this comes at the expense of reproductive success. Such tolerance and resistance traits may coexist if browsers trigger and maintain a positive feedback loop in which trees are continually investing in regrowth (tolerance), and if the plant's physical defenses (resistance) are not nutritionally costly and are long-lived. Our results confirm that chronic browsing by ungulates can maintain A. nigrescens trees in a hedged state that is analogous to a grazing lawn. Further research is needed to fully understand the long-term effects of chronic browsing on reproduction within such tree populations, as well as the overall effects on nutrient cycling at the ecosystem level.

Many tropical pioneer species depend on the presence of high seed densities in the soil for successful recruitment following canopy disturbance (Cheke et al. 1979, Dalling & Hubbell 2002, Guevara Sada & Gómez Pompa 1972, Whitmore 1983). However determinants of variation in the composition and abundance of soil seed banks remain poorly understood. Seed bank densities can be affected by rates of seed predation and pathogen infection on the surface and in the soil, by intrinsic rates of loss in viability following dispersal, and by variation in the timing and duration of fruit production (Dalling et al. 1997, Garwood 1983, Murray & Garcia 2002). Here we compare seasonal fluctuations in seed bank density in five Panamanian forests varying in elevation and seasonality of precipitation (Table 1). We predict that lowland forests should show stronger intra-annual fluctuation in seed bank densities than montane forests because seed production and loss rates should be higher under conditions of greater resource availability, and where consistent high temperatures support greater abundance or activity of seed predators and pathogens (Brühl et al. 1999). Secondly, among lowland sites, we predict greater fluctuations in seed bank densities at drier, more seasonal sites where seasonally favourable conditions for seedling recruitment may select for interspecific synchrony in fruit production (Daubenmire 1972, Garwood 1983).

Variation among forests in environmental and biotic conditions may strongly influence seed fate with important consequences for the abundance and distribution of plant species. Here we examine the post-dispersal seed removal rates of six pioneer species (Cecropia peltata, Miconia argentea, Luehea seemannii, Trema micrantha, Apeiba aspera and Jacaranda copaia) from the soil surface at five sites in Panama varying in elevation (0–1100 m) and seasonality (0–4 mo dry season). We compared removal rates of washed seeds placed in vertebrate exclosures, invertebrate exclosures, and unprotected controls in January and June. Overall, removal rates of unprotected seeds were similar among sites. Almost all seed removal could be attributed to litter ants in two subfamilies (Myrmicinae and Ponerinae). Little or no removal was recorded from invertebrate exclosures while vertebrate exclosures had no effect on removal either in lowland and montane forests. Seed removal rates were high for four animal-dispersed species (mean 45% removed over 2 d), whereas two wind-dispersed species were largely untouched (mean 2% removed). These results indicate that seed dispersal characteristics, rather than site characteristics, may be the strongest determinant of the post-dispersal seed fate of pioneers.

The functioning of human-managed grassland ecosystems strongly depends on how common management practices (e.g., animal grazing and the chronic addition of fertilizing materials to soils) interact to influence plant and soil element stoichiometry. Here we use data from a 22-yr-long grassland experiment to address whether and how plant element stoichiometry (i.e., carbon [C], nitrogen [N], phosphorus [P] ratios) might respond to (1) animal grazing, (2) agricultural liming (i.e., CaCO3) applications, and (3) nutrient fertilization. We also ask whether plant C:N:P stoichiometry could predict changes in soil N and P availability and in soil C, N, and P stocks. We found that grassland management significantly affected plant C:N:P ratios as predicted by ecological stoichiometry theory. For example, plant aboveground and belowground C:N and C:P ratios decreased under chronic N and P fertilization, respectively. Plant C:N and C:P ratios were significantly greater in unfertilized (control) soils. Also plant C:N ratios were highest under P-only additions, whereas plant C:P ratios were highest under N-only additions. However, unpredictable changes in C:N:P ratios also occurred, suggesting that plant tissue chemistry may not be a simple reflection of soil nutrient availability. Changes in plant C:nutrient ratios well predicted variation in soil nutrient availability, but not in soil C, N, and P stocks. Contrary to expectations, soil C stocks significantly increased with decreasing plant C:N ratios in the nutrient-fertilized grasslands and not with increasing plant C:N ratios in the unfertilized grasslands. We suggest that a better mechanistic understanding of the negative relationship between plant C:N stoichiometry and soil C accrual will greatly help in improving the sustainability of human-managed grasslands.

Human activities have greatly increased the availability of biologically active forms of nutrients [e.g., nitrogen (N), phosphorous (P), potassium (K), magnesium (Mg)] in many soil ecosystems worldwide. Multi-nutrient fertilization strongly increases plant productivity but may also alter the storage of carbon (C) in soil, which represents the largest terrestrial pool of organic C. Despite this issue is important from a global change perspective, key questions remain on how the single addition of N or the combination of N with other nutrients might affect C sequestration in human-managed soils. Here, we use a 19-year old nutrient addition experiment on a permanent grassland to test for nutrient-induced effects on soil C sequestration. We show that combined NPKMg additions to permanent grassland have 'constrained' soil C sequestration to levels similar to unfertilized plots whereas the single addition of N significantly enhanced soil C stocks (N-only fertilized soils store, on average, 11 t C ha(-1) more than unfertilized soils). These results were consistent across grazing and liming treatments suggesting that whilst multi-nutrient additions increase plant productivity, soil C sequestration is increased by N-only additions. The positive N-only effect on soil C content was not related to changes in plant species diversity or to the functional composition of the plant community. N-only fertilized grasslands show, however, increases in total root mass and the accumulation of organic matter detritus in topsoils. Finally, soils receiving any N addition (N only or N in combination with other nutrients) were associated with high N losses. Overall, our results demonstrate that nutrient fertilization remains an important global change driver of ecosystem functioning, which can strongly affect the long-term sustainability of grassland soil ecosystems (e.g., soils ability to deliver multiple ecosystem services).

Biomass allocation to plant aboveground and belowground compartments may change in response to changes in soil nutrient fertility (e.g., nitrogen (N) and phosphorous (P) availability). It is not clear however whether and how changes in soil fertility could influence carbon(C):nutrient stoichiometry of wild plants distributed along environmental gradients. Here we use ecological stoichiometric theory to test two hypotheses: (1) C:N (or C:P) ratios of different plant compartments (i.e., roots, stems and leaves) increase when soil N (or P) availability decreases, (2) the relative availability of N compared to P in soils predictably influences C:nutrient ratios (e.g., high soil N:P ratios are associated with high plant C:P ratios). Data from 72 wild plant species were collected along a gradient of soil development determined by the temporal progression of four primary ecological successions across Europe. We found significant changes in soil N and P availability and content along the four vegetation successio...

Spatiotemporal redistribution of incident rainfall in vegetated ecosystems results from the partitioning by plants into intercepted, stemflow, and throughfall fractions. However, variation in patterns and drivers of rainfall partitioning across global biomes remains poorly understood, which limited the ability of climate models to improve the predictions of biome hydrological cycle under global climate change scenario. Here, we synthesized and analyzed the partitioning of incident rainfall into interception, stemflow, and throughfall by trees and shrubs at the global scale using 2430 observations from 236 independent publications. We found that (1) globally, median levels of relative interception, stemflow, and throughfall accounted for 21.8%, 3.2%, and 73.0% of total incident rainfall, respectively; (2) rainfall partitioning varied among different biomes, due to variation in plant composition, canopy structure, and macroclimate; (3) relative stemflow tended to be driven by plant traits, such as crown height:width ratio, basal area, and height, while relative interception and throughfall tended to be driven by plant traits as well as meteorological variables. Our global assessment of patterns and drivers of rainfall partitioning underpins the role of meteorological factors and plant traits in biome-specific ecohydrological cycles. We suggest to include these factors in climate models to improve the predictions of local hydrological cycles and associated biodiversity and function responses to changing climate conditions. ispartof: GLOBAL CHANGE BIOLOGY vol:27 issue:14 pages:3350-3357 ispartof: location:England status: published