Linking land-use and land-cover transitions to their ecological impact in the Amazon.

Human activities pose a major threat to tropical forest biodiversity and ecosystem services. Although the impacts of deforestation are well studied, multiple land-use and land-cover transitions (LULCTs) occur in tropical landscapes, and we do not know how LULCTs differ in their rates or impacts on key ecosystem components. Here, we quantified the impacts of 18 LULCTs on three ecosystem components (biodiversity, carbon, and soil), based on 18 variables collected from 310 sites in the Brazilian Amazon. Across all LULCTs, biodiversity was the most affected ecosystem component, followed by carbon stocks, but the magnitude of change differed widely among LULCTs and individual variables. Forest clearance for pasture was the most prevalent and high-impact transition, but we also identified other LULCTs with high impact but lower prevalence (e.g., forest to agriculture). Our study demonstrates the importance of considering multiple ecosystem components and LULCTs to understand the consequences of human activities in tropical landscapes.

Carbon allocation to root exudates is maintained in mature temperate tree species under drought.

Carbon (C) exuded via roots is proposed to increase under drought and facilitate important ecosystem functions. However, it is unknown how exudate quantities relate to the total C budget of a drought-stressed tree, that is, how much of net-C assimilation is allocated to exudation at the tree level. We calculated the proportion of daily C assimilation allocated to root exudation during early summer by collecting root exudates from mature Fagus sylvatica and Picea abies exposed to experimental drought, and combining above- and belowground C fluxes with leaf, stem and fine-root surface area. Exudation from individual roots increased exponentially with decreasing soil moisture, with the highest increase at the wilting point. Despite c. 50% reduced C assimilation under drought, exudation from fine-root systems was maintained and trees exuded 1.0% (F. sylvatica) to 2.5% (P. abies) of net C into the rhizosphere, increasing the proportion of C allocation to exudates two- to three-fold. Water-limited P. abies released two-thirds of its exudate C into the surface soil, whereas in droughted F. sylvatica it was only one-third. Across the entire root system, droughted trees maintained exudation similar to controls, suggesting drought-imposed belowground C investment, which could be beneficial for ecosystem resilience.

Distinct storage mechanisms of soil organic carbon in coniferous forest and evergreen broadleaf forest in tropical China.

The impact of human activities on soil carbon (C) storage in tropical forests has aroused wide concern during the past decades, because these ecosystems play a key role in ameliorating global climate change. However, there remain uncertainties about how land-use history alters soil organic carbon (SOC) stability and storage in different forests. In this study, we measured the C content and mass distributions of soil aggregates, density fractions, mineral-bound C and microbial biomass C in the organic horizon, 0-10 cm and 10-20 cm soil layers in coniferous forest and evergreen broadleaf forest at Dinghushan Biosphere Reserve in tropical China. The broadleaf forest had larger SOC stocks than the coniferous forest, but the proportion of SOC stored in different density fractions at 0-10 cm soils was similar between forest types, while a greater proportion of SOC was stored in microaggregates in the coniferous forest. Most of the SOC was held as light fraction C in the organic horizon in the coniferous forest, whereas the concentrations of mineral-bound C were higher in the broadleaf forest. These findings indicate clear differences in the protection of SOC between broadleaf and coniferous forests growing on the same soil type. We propose that historic conversion of broadleaf forest to coniferous forest has reduced soil C sequestration capacity by altering the diversity and quality of plant inputs to the soil, which in turn affected macroaggregate formation, soil chemical properties and microbial biomass. Our results thus demonstrate that changes in forest tree species composition could have long-lasting effects on soil structure and carbon storage, providing crucial evidence for policy decisions on forest carbon sink management.

Deepened winter snow cover enhances net ecosystem exchange and stabilizes plant community composition and productivity in a temperate grassland.

Global warming has greatly altered winter snowfall patterns, and there is a trend towards increasing winter snow in semi-arid regions in China. Winter snowfall is an important source of water during early spring in these water-limited ecosystems, and it can also affect nutrient supply. However, we know little about how changes in winter snowfall will affect ecosystem productivity and plant community structure during the growing season. Here, we conducted a 5-year winter snow manipulation experiment in a temperate grassland in Inner Mongolia. We measured ecosystem carbon flux from 2014 to 2018 and plant biomass and species composition from 2015 to 2018. We found that soil moisture increased under deepened winter snow in early growing season, particularly in deeper soil layers. Deepened snow increased the net ecosystem exchange of CO2 (NEE) and reduced intra- and inter-annual variation in NEE. Deepened snow did not affect aboveground plant biomass (AGB) but significantly increased root biomass. This suggested that the enhanced NEE was allocated to the belowground, which improved water acquisition and thus contributed to greater stability in NEE in deep-snow plots. Interestingly, the AGB of grasses in the control plots declined over time, resulting in a shift towards a forb-dominated system. Similar declines in grass AGB were also observed at three other locations in the region over the same time frame and are attributed to 4 years of below-average precipitation during the growing season. By contrast, grass AGB was stabilized under deepened winter snow and plant community composition remained unchanged. Hence, our study demonstrates that increased winter snowfall may stabilize arid grassland systems by reducing resource competition, promoting coexistence between plant functional groups, which ultimately mitigates the impacts of chronic drought during the growing season.

Tree stem bases are sources of CH4 and N2 O in a tropical forest on upland soil during the dry to wet season transition.

Tropical forests on upland soils are assumed to be a methane (CH4 ) sink and a weak source of nitrous oxide (N2 O), but studies of wetland forests have demonstrated that tree stems can be a substantial source of CH4 , and recent evidence from temperate woodlands suggests that tree stems can also emit N2 O. Here, we measured CH4 and N2 O fluxes from the soil and from tree stems in a semi-evergreen tropical forest on upland soil. To examine the influence of seasonality, soil abiotic conditions and substrate availability (litter inputs) on trace greenhouse gas (GHG) fluxes, we conducted our study during the transition from the dry to the wet season in a long-term litter manipulation experiment in Panama, Central America. Trace GHG fluxes were measured from individual stem bases of two common tree species and from soils beneath the same trees. Soil CH4 fluxes varied from uptake in the dry season to minor emissions in the wet season. Soil N2 O fluxes were negligible during the dry season but increased markedly after the start of the wet season. By contrast, tree stem bases emitted CH4 and N2 O throughout the study. Although we observed no clear effect of litter manipulation on trace GHG fluxes, tree species and litter treatments interacted to influence CH4 fluxes from stems and N2 O fluxes from stems and soil, indicating complex relationships between tree species traits and decomposition processes that can influence trace GHG dynamics. Collectively, our results show that tropical trees can act as conduits for trace GHGs that most likely originate from deeper soil horizons, even when they are growing on upland soils. Coupled with the finding that the soils may be a weaker sink for CH4 than previously thought, our research highlights the need to reappraise trace gas budgets in tropical forests.

Arbuscular mycorrhizal fungal community composition is altered by long-term litter removal but not litter addition in a lowland tropical forest.

Tropical forest productivity is sustained by the cycling of nutrients through decomposing organic matter. Arbuscular mycorrhizal (AM) fungi play a key role in the nutrition of tropical trees, yet there has been little experimental investigation into the role of AM fungi in nutrient cycling via decomposing organic material in tropical forests. We evaluated the responses of AM fungi in a long-term leaf litter addition and removal experiment in a tropical forest in Panama. We described AM fungal communities using 454-pyrosequencing, quantified the proportion of root length colonised by AM fungi using microscopy, and estimated AM fungal biomass using a lipid biomarker. AM fungal community composition was altered by litter removal but not litter addition. Root colonisation was substantially greater in the superficial organic layer compared with the mineral soil. Overall colonisation was lower in the litter removal treatment, which lacked an organic layer. There was no effect of litter manipulation on the concentration of the AM fungal lipid biomarker in the mineral soil. We hypothesise that reductions in organic matter brought about by litter removal may lead to AM fungi obtaining nutrients from recalcitrant organic or mineral sources in the soil, besides increasing fungal competition for progressively limited resources.

The grassland carbon cycle: Mechanisms, responses to global changes, and potential contribution to carbon neutrality.

Grassland is one of the largest terrestrial biomes, providing critical ecosystem services such as food production, biodiversity conservation, and climate change mitigation. Global climate change and land-use intensification have been causing grassland degradation and desertification worldwide. As one of the primary medium for ecosystem energy flow and biogeochemical cycling, grassland carbon (C) cycling is the most fundamental process for maintaining ecosystem services. In this review, we first summarize recent advances in our understanding of the mechanisms underpinning spatial and temporal patterns of the grassland C cycle, discuss the importance of grasslands in regulating inter- and intra-annual variations in global C fluxes, and explore the previously unappreciated complexity in abiotic processes controlling the grassland C balance, including soil inorganic C accumulation, photochemical and thermal degradation, and wind erosion. We also discuss how climate and land-use changes could alter the grassland C balance by modifying the water budget, nutrient cycling and additional plant and soil processes. Further, we examine why and how increasing aridity and improper land use may induce significant losses in grassland C stocks. Finally, we identify several priorities for future grassland C research, including improving understanding of abiotic processes in the grassland C cycle, strengthening monitoring of grassland C dynamics by integrating ground inventory, flux monitoring, and modern remote sensing techniques, and selecting appropriate plant species combinations with suitable traits and strong resistance to climate fluctuations, which would help design sustainable grassland restoration strategies in a changing climate.

Soil carbon storage is related to tree functional composition in naturally regenerating tropical forests.

Regenerating tropical forests are increasingly important for their role in the global carbon cycle. Carbon stocks in above-ground biomass can recover to old-growth forest levels within 60-100 years. However, more than half of all carbon in tropical forests is stored below-ground, and our understanding of carbon storage in soils during tropical forest recovery is limited.Importantly, soil carbon accumulation does not necessarily reflect patterns in above-ground biomass carbon accrual during secondary forest succession, and factors related to past land use, species composition and soil characteristics may influence soil carbon storage during forest regrowth.Using tree census data and a measure of tree community shade tolerance (species-specific light response values), we assessed the relationship between soil organic carbon stocks and tree functional groups during secondary succession along a chronosequence of 40- to 120-year-old naturally regenerating secondary forest and old-growth tropical forest stands in Panama.While previous studies found no evidence for increasing soil C storage with secondary forest age, we found a strong relationship between tree functional composition and soil carbon stocks at 0-10 cm depth, whereby carbon stocks increased with the relative influence of light-demanding tree species. Light demanding trees had higher leaf nitrogen but lower leaf density than shade-tolerant trees, suggesting that rapid decomposition of nutrient-rich plant material in forests with a higher proportion of light-demanding species results in greater accumulation of carbon in the surface layer of soils. Synthesis. We propose that soil carbon storage in secondary tropical forests is more strongly linked to tree functional composition than forest age, and that the persistence of long-lived pioneer trees could enhance soil carbon storage as forests age. Considering shifts in tree functional groups could improve estimates of carbon sequestration potential for climate change mitigation by tropical forest regrowth. Read the free Plain Language Summary for this article on the Journal blog.

Los bosques tropicales en regeneración son cada vez más importantes por su papel en el ciclo global del carbono. Las reservas de carbono en la biomasa aérea pueden recuperarse hasta los niveles de los bosques maduros en un plazo de 60 a 100 años. Sin embargo, más de la mitad de todo el carbono en los bosques tropicales se almacena bajo tierra, y nuestra comprensión del almacenamiento de carbono en los suelos durante la recuperación de los bosques tropicales es limitada.Es importante señalar que la acumulación de carbono en el suelo no refleja necesariamente los patrones de acumulación de carbono en la biomasa aérea durante la sucesión de bosques secundarios y los factores relacionados con el uso pasado del terreno, la composición de especies y las características del suelo pueden influir en el almacenamiento de carbono en el suelo durante la regeneración del bosque.Usando datos del censo de árboles y una medida de la tolerancia a la sombra de la comunidad de árboles (valores de respuesta a la luz específicos de la especie), evaluamos la relación entre las reservas de carbono orgánico del suelo y los grupos funcionales de los árboles durante la sucesión secundaria a lo largo de una cronosecuencia de 40 a 120 años bosques secundarios de regeneración natural y rodales de bosques tropicales primarios en Panamá.Mientras que estudios previos no encontraron evidencia de un aumento del almacenamiento de C en el suelo con la edad del bosque secundario, encontramos una fuerte relación entre la composición funcional de los árboles y las reservas de carbono del suelo a 0­10 cm de profundidad, por lo que las reservas de carbono aumentaron con la influencia relativa de especies de árboles que demanda de luz. Los árboles que requieren luz tenían más nitrógeno en las hojas pero menor densidad de hojas que los árboles tolerantes a la sombra, lo que sugiere que la descomposición rápida del material vegetal rico en nutrientes en los bosques con una mayor proporción de especies que requieren luz da como resultado una mayor acumulación de carbono en la capa superficial de los suelos. Síntesis. Proponemos que el almacenamiento de carbono en el suelo en los bosques tropicales secundarios está más fuertemente relacionado con la composición funcional de los árboles que con la edad del bosque, y que la persistencia de árboles pioneros de larga vida podría mejorar el almacenamiento de carbono en el suelo a medida que los bosques envejecen. La consideración de los cambios en los grupos funcionales de los árboles podría mejorar las estimaciones del potencial de secuestro de carbono para la mitigación del cambio climático mediante la regeneración de los bosques tropicales.

Delayed wet season increases soil net N mineralization in a seasonally dry tropical forest.

Seasonal precipitation regime plays a vital role in regulating nutrient dynamics in seasonally dry tropical forests. Present evidence suggests that not only wet season precipitation is increasing in the tropics of South China, but also that the wet season is occurring later. However, it is unclear how nutrient dynamics will respond to the projected precipitation regime changes. We assessed the impacts of altered seasonal precipitation on soil net N mineralization in a secondary tropical forest. Since 2013, by reducing throughfall and/or irrigating experimental plots, we delayed the wet season by two months from April-September to June-November (DW treatment) or increased annual precipitation by 25% in July and August (WW treatment). We measured soil net N mineralization rates and assessed soil microbial communities in January, April, August and November in 2015 and 2017. We found that a wetter wet season did not significantly affect soil microbes or net N mineralization rates, even in the mid-wet season (August) when soil water content in the WW treatment increased significantly. By contrast, a delayed wet season enhanced soil microbial biomass and altered microbial community structure, resulting in a two-fold increase in net N mineralization rates relative to controls in the early dry season (November). Structural equation modeling showed that the changes in net N mineralization during the early dry season were associated with altered soil microbial communities, dissolved organic N, and litterfall, which were all affected by enhanced soil water content. Our findings suggest that a delayed wet season could have a greater impact on N dynamics than increased precipitation during the wet season. Changes in the seasonal timing of rainfall might therefore influence the functioning of seasonally dry tropical forests.

Seasonal Influence of Biodiversity on Soil Respiration in a Temperate Forest.

Soil respiration in forests contributes to significant carbon dioxide emissions from terrestrial ecosystems but it varies both spatially and seasonally. Both abiotic and biotic factors influence soil respiration but their relative contribution to spatial and seasonal variability remains poorly understood, which leads to uncertainty in models of global C cycling and predictions of future climate change. Here, we hypothesize that tree diversity, soil diversity, and soil properties contribute to local-scale variability of soil respiration but their relative importance changes in different seasons. To test our hypothesis, we conducted seasonal soil respiration measurements along a local-scale environmental gradient in a temperate forest in Northeast China, analyzed spatial variability of soil respiration and tested the relationships between soil respiration and a variety of abiotic and biotic factors including topography, soil chemical properties, and plant and soil diversity. We found that soil respiration varied substantially across the study site, with spatial coefficients of variation (CV) of 29.1%, 27.3% and 30.8% in spring, summer, and autumn, respectively. Soil respiration was consistently lower at high soil water content, but the influence of other factors was seasonal. In spring, soil respiration increased with tree diversity and biomass but decreased with soil fungal diversity. In summer, soil respiration increased with soil temperature, whereas in autumn, soil respiration increased with tree diversity but decreased with increasing soil nutrient content. However, soil nutrient content indirectly enhanced soil respiration via its effect on tree diversity across seasons, and forest stand structure indirectly enhanced soil respiration via tree diversity in spring. Our results highlight that substantial differences in soil respiration at local scales was jointly explained by soil properties (soil water content and soil nutrients), tree diversity, and soil fungal diversity but the relative importance of these drivers varied seasonally in our temperate forest.

Potassium, phosphorus, or nitrogen limit root allocation, tree growth, or litter production in a lowland tropical forest.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 x 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

Long-term fertilization modifies the mineralization of soil organic matter in response to added substrate.

The turnover of SOC in soils is strongly influenced by the availability of substrate and nutrients, especially nitrogen (N) and phosphorus (P). Here, we assessed how long-term fertilization modified SOM mineralization in response to added substrate in a tropical forest. We carried out a 90-day incubation study in which we added two structurally similar compounds which differed in microbial metabolic availability: corn cellulose or corn starch to soils collected from a long-term (11 years) factorial N and P fertilization experiment site in a tropical forest in south China. We measured total soil mineralization rate (CO2 efflux) to characterize SOM mineralization and using 13C isotope signatures to determine the source of the CO2 (original soil C or added substrate) and assessed changes in extracellular enzyme activities: acid phosphomonoesterase (AP), ß-1,4-glucosidase (BG), ß-1,4- N-acetaminophen glucosidase (NAG), phenol oxidase (PHO) and peroxidase (PER), and microbial biomarkers to determine whether nutrient stoichiometry and decomposer communities explain differences in SOM mineralization rates. Total C mineralization increased substantially with substrate addition, particularly cellulose (5.38, 7.13, 5.58 and 5.37 times for N, P, NP fertilization and CK, respectively) compared to no substrate addition, and original soil C mineralization was further enhanced in long-term N (3.40% and 5.18% for cellulose and starch addition, respectively) or NP (35.11% for cellulose addition) fertilized soils compared to control treatment. Enzyme activities were stimulated by the addition of both substrates but suppressed by P-fertilization. Addition of both substrates increased microbial investment in P-acquisition, but only starch addition promoted C investment in N-acquisition. Finally, fungal abundance increased with substrate addition to a greater extent than bacterial abundance, particularly in cellulose-amended soils, and the effect was amplified by long-term fertilization. Our findings indicate that SOM mineralization might be enhanced in N and P enrichment ecosystems, since the litter input can liberate microbes from C limitation and stimulate SOM mineralization if N and P are sufficient. Our study further demonstrates that structurally similar substrates can have distinct effects on SOM mineralization and the extent of SOM mineralization is strongly dependent on elemental stoichiometry, as well as the resource requirements of microbial decomposers.

Adaptation to chronic drought modifies soil microbial community responses to phytohormones.

Drought imposes stress on plants and associated soil microbes, inducing coordinated adaptive responses, which can involve plant-soil signalling via phytohormones. However, we know little about how microbial communities respond to phytohormones, or how these responses are shaped by chronic (long-term) drought. Here, we added three phytohormones (abscisic acid, 1-aminocyclopropane-1-carboxylic acid, and jasmonic acid) to soils from long-term (25-year), field-based climate treatments to test the hypothesis that chronic drought alters soil microbial community responses to plant stress signalling. Phytohormone addition increased soil respiration, but this effect was stronger in irrigated than in droughted soils and increased soil respiration at low phytohormone concentrations could not be explained by their use as substrate. Thus, we show that drought adaptation within soil microbial communities modifies their responses to phytohormone inputs. Furthermore, distinct phytohormone-induced shifts in microbial functional groups in droughted vs. irrigated soils might suggest that drought-adapted soil microorganisms perceive phytohormones as stress-signals, allowing them to anticipate impending drought.

High foliar K and P resorption efficiencies in old-growth tropical forests growing on nutrient-poor soils.

Resorption is the active withdrawal of nutrients before leaf abscission. This mechanism represents an important strategy to maintain efficient nutrient cycling; however, resorption is poorly characterized in old-growth tropical forests growing in nutrient-poor soils. We investigated nutrient resorption from leaves in 39 tree species in two tropical forests on the Guiana Shield, French Guiana, to investigate whether resorption efficiencies varied with soil nutrient, seasonality, and species traits. The stocks of P in leaves, litter, and soil were low at both sites, indicating potential P limitation of the forests. Accordingly, mean resorption efficiencies were higher for P (35.9%) and potassium (K; 44.6%) than for nitrogen (N; 10.3%). K resorption was higher in the wet (70.2%) than in the dry (41.7%) season. P resorption increased slightly with decreasing total soil P; and N and P resorptions were positively related to their foliar concentrations. We conclude that nutrient resorption is a key plant nutrition strategy in these old-growth tropical forests, that trees with high foliar nutrient concentration reabsorb more nutrient, and that nutrients resorption in leaves, except P, are quite decoupled from nutrients in the soil. Seasonality and biochemical limitation played a role in the resorption of nutrients in leaves, but species-specific requirements obscured general tendencies at stand and ecosystem level.

The diversity of decay.

To predict how species loss will affect ecosystems, it is important to consider how biodiversity influences processes such as decomposition.

Author Correction: Tropical forest soil carbon stocks do not increase despite 15 years of doubled litter inputs.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Tropical forest soil carbon stocks do not increase despite 15 years of doubled litter inputs.

Soil organic carbon (SOC) dynamics represent a persisting uncertainty in our understanding of the global carbon cycle. SOC storage is strongly linked to plant inputs via the formation of soil organic matter, but soil geochemistry also plays a critical role. In tropical soils with rapid SOC turnover, the association of organic matter with soil minerals is particularly important for stabilising SOC but projected increases in tropical forest productivity could trigger feedbacks that stimulate the release of stored SOC. Here, we demonstrate limited additional SOC storage after 13-15 years of experimentally doubled aboveground litter inputs in a lowland tropical forest. We combined biological, physical, and chemical methods to characterise SOC along a gradient of bioavailability. After 13 years of monthly litter addition treatments, most of the additional SOC was readily bioavailable and we observed no increase in mineral-associated SOC. Importantly, SOC with weak association to soil minerals declined in response to long-term litter addition, suggesting that increased plant inputs could modify the formation of organo-mineral complexes in tropical soils. Hence, we demonstrate the limited capacity of tropical soils to sequester additional C inputs and provide insights into potential underlying mechanisms.

Distinct responses of soil respiration to experimental litter manipulation in temperate woodland and tropical forest.

Global change is affecting primary productivity in forests worldwide, and this, in turn, will alter long-term carbon (C) sequestration in wooded ecosystems. On one hand, increased primary productivity, for example, in response to elevated atmospheric carbon dioxide (CO 2), can result in greater inputs of organic matter to the soil, which could increase C sequestration belowground. On other hand, many of the interactions between plants and microorganisms that determine soil C dynamics are poorly characterized, and additional inputs of plant material, such as leaf litter, can result in the mineralization of soil organic matter, and the release of soil C as CO 2 during so-called "priming effects". Until now, very few studies made direct comparison of changes in soil C dynamics in response to altered plant inputs in different wooded ecosystems. We addressed this with a cross-continental study with litter removal and addition treatments in a temperate woodland (Wytham Woods) and lowland tropical forest (Gigante forest) to compare the consequences of increased litterfall on soil respiration in two distinct wooded ecosystems. Mean soil respiration was almost twice as high at Gigante (5.0 µmol CO 2 m-2 s-1) than at Wytham (2.7 µmol CO 2 m-2 s-1) but surprisingly, litter manipulation treatments had a greater and more immediate effect on soil respiration at Wytham. We measured a 30% increase in soil respiration in response to litter addition treatments at Wytham, compared to a 10% increase at Gigante. Importantly, despite higher soil respiration rates at Gigante, priming effects were stronger and more consistent at Wytham. Our results suggest that in situ priming effects in wooded ecosystems track seasonality in litterfall and soil respiration but the amount of soil C released by priming is not proportional to rates of soil respiration. Instead, priming effects may be promoted by larger inputs of organic matter combined with slower turnover rates.

Responses of arbuscular mycorrhizal fungi to long-term inorganic and organic nutrient addition in a lowland tropical forest.

Improved understanding of the nutritional ecology of arbuscular mycorrhizal (AM) fungi is important in understanding how tropical forests maintain high productivity on low-fertility soils. Relatively little is known about how AM fungi will respond to changes in nutrient inputs in tropical forests, which hampers our ability to assess how forest productivity will be influenced by anthropogenic change. Here we assessed the influence of long-term inorganic and organic nutrient additions and nutrient depletion on AM fungi, using two adjacent experiments in a lowland tropical forest in Panama. We characterised AM fungal communities in soil and roots using 454-pyrosequencing, and quantified AM fungal abundance using microscopy and a lipid biomarker. Phosphorus and nitrogen addition reduced the abundance of AM fungi to a similar extent, but affected community composition in different ways. Nutrient depletion (removal of leaf litter) had a pronounced effect on AM fungal community composition, affecting nearly as many OTUs as phosphorus addition. The addition of nutrients in organic form (leaf litter) had little effect on any AM fungal parameter. Soil AM fungal communities responded more strongly to changes in nutrient availability than communities in roots. This suggests that the 'dual niches' of AM fungi in soil versus roots are structured to different degrees by abiotic environmental filters, and biotic filters imposed by the plant host. Our findings indicate that AM fungal communities are fine-tuned to nutrient regimes, and support future studies aiming to link AM fungal community dynamics with ecosystem function.

Links between soil microbial communities and plant traits in a species-rich grassland under long-term climate change.

Climate change can influence soil microorganisms directly by altering their growth and activity but also indirectly via effects on the vegetation, which modifies the availability of resources. Direct impacts of climate change on soil microorganisms can occur rapidly, whereas indirect effects mediated by shifts in plant community composition are not immediately apparent and likely to increase over time. We used molecular fingerprinting of bacterial and fungal communities in the soil to investigate the effects of 17 years of temperature and rainfall manipulations in a species-rich grassland near Buxton, UK. We compared shifts in microbial community structure to changes in plant species composition and key plant traits across 78 microsites within plots subjected to winter heating, rainfall supplementation, or summer drought. We observed marked shifts in soil fungal and bacterial community structure in response to chronic summer drought. Importantly, although dominant microbial taxa were largely unaffected by drought, there were substantial changes in the abundances of subordinate fungal and bacterial taxa. In contrast to short-term studies that report high resistance of soil fungi to drought, we observed substantial losses of fungal taxa in the summer drought treatments. There was moderate concordance between soil microbial communities and plant species composition within microsites. Vector fitting of community-weighted mean plant traits to ordinations of soil bacterial and fungal communities showed that shifts in soil microbial community structure were related to plant traits representing the quality of resources available to soil microorganisms: the construction cost of leaf material, foliar carbon-to-nitrogen ratios, and leaf dry matter content. Thus, our study provides evidence that climate change could affect soil microbial communities indirectly via changes in plant inputs and highlights the importance of considering long-term climate change effects, especially in nutrient-poor systems with slow-growing vegetation.