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Nitrogen (N) deposition is a pervasive anthropogenic change that can alter the dynamics and storage of carbon (C) in tropical soils by altering the transformation of plant litter C to soil organic C (SOC). The response of fungi may be particularly important for explaining how N deposition affects SOC storage in tropical forests because they are the primary decomposers of plant litter. Fungi can be grouped into different ecological guilds, or functional groups, which often have contrasting responses to the same environmental change. However, little is known if shifts in fungal guilds and their functions explain increased tropical forest SOC storage under N deposition. We addressed this knowledge gap by assessing the effects of long-term (6 years) experimental N addition (+N) on fungal guilds, their functions and SOC fractions in a tropical forest. Total SOC, dissolved organic C (DOC), particulate organic C and fungal necromass C were all higher in +N soils. The fungal community in +N soils was dominated by generalist saprotrophs, and leaf saprotroph abundance increased nearly 93-fold compared to controls. These changes were accompanied by an increase in lignocellulolytic enzymes, indicating accelerated decomposition of recalcitrant litter C compounds. Chitosanase, which catalyses synthesis of glucosamine, was 1.37× higher in +N soils than controls; correspondingly, fungal necromass C contributed 1.42× more to SOC. Greater abundance of saprotrophic enzymes involved in organic acid synthesis in +N soils was associated with higher concentration of Ca2+ and DOC. We propose that N deposition increases SOC storage by enhancing saprotroph abundance and activity, whereby increased organic acid production supports fungal growth both directly and indirectly via the release of nutrients and DOC, resulting in greater inputs of necromass to the soil. Our findings highlight the importance of considering shifts in guild-level fungal abundance and function to investigate changes in SOC dynamics and storage.

In recent years, important processes controlling ecosystem carbon dynamics have been connected to fine-root exudation of soluble carbon compounds. Root exudation patterns may change depending on plant interactions and plant susceptibility to and recovery from drought. Recent investigations suggest that root exudation tends to increase with stress events, but quantification of the amount of carbon released from roots across soil depths with differing water availability and species interactions are missing.We tested if root exudation rates were negatively correlated with soil water content across soil depths during and after drought. We further tested if species in mixture, often considered to be less stressed under drought, exuded less carbon than species in monospecific environments. Exudates were sampled in a mature Fagus sylvatica L. and Picea abies (L.) Karst. forest at the end of a five-year throughfall exclusion period and again one year after the drought ended. We quantified root exudates and their variation with soil depth for both tree species in monospecific and mixed species zones.Carbon exudation significantly increased in fine roots exposed to dry soils (

Forest soils are crucial for many ecosystem services that rely on soil organic matter (SOM) stability. Carbon allocated to roots and released as exudates to the rhizosphere plays a key role in SOM stabilization. Under periodic drought, elevated root exudation and SOM accumulation have been reported. Yet, whether root exudates control SOM formation and stability in mature forests once the drought ends is largely unknown. We examined whether root exudates from P. abies and F. sylvatica trees relate to SOM formation and stability in soil depth profiles one year following five years of experimental drought (Kroof experiment, Germany). We collected root exudates throughout the rooting zone and combined the data with thermogravimetric analysis of SOM in the rhizosphere and non-rooted soil. We found that the rhizosphere of both species was characterized by stable SOM fractions that did not decrease post-drought, suggesting potential protection of SOM due to rhizodeposition and root exudates. In contrast, stable SOM fractions decreased relative to controls in non-rooted topsoil below P. abies, indicating a loss of stabilized SOM from drought-affected and re-wetted soil. Our measurements provide valuable insights into post-drought SOM formation and mechanisms of SOM stabilization in forest ecosystems under climate change.

Human activities pose a major threat to tropical forest biodiversity and ecosystem serv-ices. Although the impacts of deforestation are well studied, multiple land-use andland-cover transitions (LULCTs) occur in tropical landscapes, and we do not knowhow LULCTs differ in their rates or impacts on key ecosystem components. Here, wequantified the impacts of 18 LULCTs on three ecosystem components (biodiversity,carbon, and soil), based on 18 variables collected from 310 sites in the BrazilianAmazon. Across all LULCTs, biodiversity was the most affected ecosystem component,followed by carbon stocks, but the magnitude of change differed widely amongLULCTs and individual variables. Forest clearance for pasture was the most prevalentand high-impact transition, but we also identified other LULCTs with high impact butlower prevalence (e.g., forest to agriculture). Our study demonstrates the importanceof considering multiple ecosystem components and LULCTs to understand the conse-quences of human activities in tropical landscapes Made available in DSpace on 2022-10-06T16:05:22Z (GMT). No. of bitstreams: 1 pnas.2202310119.pdf: 1994033 bytes, checksum: 7b87b3ce0f689008a66ab5755a038d75 (MD5) Previous issue date: 2022

1. Plant diversity supports multiple ecosystem functions, including carbon sequestration. Recent shifts in plant diversity in rangelands due to increased grazing pressure and climate changes have the potential to impact the sequestration of carbon in arid to semi-humid regions worldwide. However, plant diversity, grazing intensity and carbon storage are also influenced by environmental factors such as nutrient availability, climate, and topography. The complexity of these interactions limits our ability to fully assess the impacts of grazing on biodiversity-ecosystem function (BEF) relationships. 2. We assessed how grazing intensity modifies BEF relationships by determining the links between plant diversity and ecosystem carbon stocks (plant and soil carbon) across broad environmental gradients and different plant growth forms. To achieve this, we surveyed 1493 quadrats across 10 rangelands, covering an area of 23,756 ha in northern Iran. 3. We show that aboveground carbon stocks increased with plant diversity across topographic, climatic and soil fertility gradients. The relationship between aboveground carbon stocks and plant diversity was strongest for forbs, followed by shrubs and grasses. Soil carbon stocks increased strongly with soil fertility across sites, but aridity, grazing, plant diversity and topography were also important in explaining variation in soil carbon stocks. 4. Importantly, aboveground and soil carbon stocks declined at high grazing intensity, and grazing modified the relationship between plant diversity and carbon stocks regardless of differences in abiotic conditions across sites. 4. Our study demonstrates that relationships between plant diversity and ecosystem carbon stocks persist across gradients of aridity, topography, and soil fertility, but the relationships are modified by grazing intensity. Our findings suggest that potential losses in plant diversity under grazing intensification could reduce ecosystem carbon storage across wide areas of arid to semi-humid rangelands. We discuss the potential mechanisms underpinning rangeland BEF relationships to stimulate future research., We would like to thank all of those involved in this research and who helped with the fieldwork. A. Sanaei is supported by the Saxon State Ministry for Science, Culture and Tourism (SMWK) – [3-7304/35/6-2021/48880]. Z. Yuan is supported by the National Natural Science Foundation of China (32171581) and the Fundamental Research Funds for the Central Universities. H. Saiz is supported by a María Zambrano fellowship funded by the Ministry of Universities and European Union-Next Generation plan. M. Delgado-Baquerizo is supported by a Ramón y Cajal grant (RYC2018-025483-I), a project from the Spanish Ministry of Science and Innovation (PID2020-115813RA-I00), and a project PAIDI 2020 from the Junta de Andalucía (P20_00879). A. Ali is supported by the faculty start-up research funding program at Hebei University for developing the Forest Ecology Research Group (Special Project No. 521100221033)

1. Carbon (C) loss due to soil erosion is a major issue in semi-arid grasslands. The extent of soil erosion is determined by soil properties and vegetation structure, especially during the non-growing season. In many Inner Mongolian grasslands, intensive land-use, such as overgrazing and mowing, has severely reduced plant cover and damaged soil structure, which has exacerbated soil C loss by erosion. At the same time, increasing winter snowfall due to climate change is stimulating plant growth and altering plant composition. However, we do not know how changes in winter snow cover interact with land-use practices to regulate soil C loss due to erosion. 2. Here, we conducted a 6-year snow manipulation experiment under different land-use practices (control; moderately mowed, MM; heavily mowed, HM) to measure net changes in soil depth, soil C, plant biomass and vegetation structure. 3. After 6 years, soil C loss under ambient snow was three times greater in the MM and four times greater in the HM treatment compared with controls during non-growing season. However, deepened winter snow alleviated erosion-induced soil C loss by 14%, 47%, 16% in the controls, MM and HM treatments respectively. 4. The severity of soil C loss declined with increasing above-ground biomass (AGB), surface root biomass and vegetation structure. Vegetation structure and AGB explained more of the variation in soil C loss than surface root biomass, possibly because a complex canopy and plant cover increases overall surface roughness, thereby reducing soil C loss. Intensified land-use reduced AGB, surface root biomass and vegetation structure, but deepened snow increased overall surface roughness by promoting AGB. Hence, our study demonstrates that deepened snow can alleviate soil C loss due to land-use practices by promoting AGB. A free Plain Language Summary can be found within the Supporting Information of this article.

Here we describe and evaluate the success of a multi-institutional Centre for Doctoral Training (CDT), which was established to address a UK skills shortage in Soil Science. The government-funded ‘STARS’ (Soils Training And Research Studentships) CDT was established in 2015 across a range of universities and research institutes in the UK. It recruited 41 PhD students equitably split across the institutions under four core research themes identified as being central to the national need, namely, (1) Understanding the soil–root interface, (2) Soils and the delivery of ecosystem services, (3) Resilience and response of functions in soil systems and (4) Modelling the soil ecosystem at different spatial and temporal scales. In addition, the STARS CDT provided a diverse skills programme, including: Holistic training in soils, the promotion of collegiality and joint working, strategies to promote science and generate impact, internships with end users (e.g., policymakers, industry), personal wellbeing, and ways to generate a lasting soils training legacy. Overall, both supervisors and students have reported a positive experience of the CDT in comparison to the conventional doctoral training programmes, which have less discipline focus and little chance for students to scientifically interact with their cohorts or to undertake joint training activities. The STARS CDT also allowed students to freely access research infrastructure across the partner institutions (e.g., long-term field trials, specialised analytical facilities, high-performance computing), breaking down traditional institutional barriers and thus maximising the students' potential to undertake high-quality research. The success and legacy of the STARS CDT can be evidenced in many ways; however, it is exemplified by the large number and diversity of journal papers produced, the lasting collaborations, final career destinations, and creation of a web-based legacy portal including new and reflective video material. STARS CDT, Grant/Award Numbers: NE/V017667/1, NE/R010218/1, NE/M009106/1; UKRI

Casuarina equisetifolia plantations have been widely established along tropical and subtropical coasts, where they act as a windbreak to shelter coastal areas. These shelter plantations also fulfill important ecosystem service by sequestering large amounts of carbon (C). However, shelter plantations are usually established on nutrient-poor sand dunes, which may limit tree growth and therefore C sequestration rates. To assess whether fertilization increases the C sequestration of coastal shelter plantations, we conducted a fertilization experiment in a young C. equisetifolia plantation with four treatments: nitrogen addition (+N), phosphorus addition (+P), nitrogen and phosphorus addition (+NP), and controls (CK). We quantified nutrient availability and annual net ecosystem production (NEP) during the key early establishment phase. Fertilization with +N and + P significantly increased ammonium-N and extractable P in the surface soils and the N and P concentrations of branches, respectively. Surprisingly, the mean growth rate of C. equisetifolia plantations (24.59 ± 0.66 Mg ha−1 yr−1) was not affected by fertilization. The mean NEP was 6.18 ± 0.49 Mg C ha−1 yr−1, and there was no difference among fertilization treatments. However, soil organic C significantly increased by 41% and 36% with +N and + P, respectively, but not +NP. C. equisetifolia plantations can sequester large amounts of C in biomass on poor soils without the need for additional nutrients, although fertilization may enhance soil C storage. Considering the potential large planting area, we suggest that coastal C. equisetifolia shelter plantations could be an important nature-based solution for climate change mitigation in coastal regions.

Grazing can alter plant species interactions in natural rangelands, which in turn might influence the productivity of the ecosystem but we do not fully understand how spatial variability in plant diversity-biomass relationships are modulated by grazing intensity. Here, we hypothesized that plant species co-occurrence in rangelands is mainly driven by niche segregation due to grazing and heterogeneity in local resources, and that grazing, therefore, modulates diversity–biomass relationships. We tested our hypothesis across 35 rangeland sites in Iran, using a species co-occurrence index to assess plant spatial aggregation within each site. At each site, we measured aboveground biomass, plant diversity, topography, soil nutrients, and three levels of grazing intensity. High spatial segregation of plant communities (low species co-occurrence) was found at heavily grazed sites, whereas greater spatial aggregation (high species co-occurrence) was found on low and moderate grazed sites, showing varied associational patterns of species with grazing intensity. Soil nutrients increased with grazing intensity and spatial segregation of plant communities was greater at sites with high soil nutrient concentrations, indicating that grazing intensity influences the spatial heterogeneity of plant communities via nutrients deposited in urine and feces. Declining plant biomass with grazing intensity was related to a strong decline in graminoid species diversity, which suggests that the diversity-biomass relationship is influenced by selective grazing of palatable species. The relationships between species co-occurrence and biomass or plant diversity suggest non-random patterns in species co-occurrences with grazing intensity, which could be the result of competition driven by high livestock grazing intensity. We, therefore, suggest that rangeland stocking rates should be managed properly to maintain rangeland production while promoting plant diversity.

Tropical forest biodiversity is being threatened by human activities, and species losses during forest disturbance can compromise important ecosystem functions and services. We assessed how species losses due to tropical forest disturbance affect community functional structure, using Amazonian dung beetles as a model group. We collected empirical data from 106 forest transects and used simulated extinction scenarios to determine how species loss influences community structure at regional and local scales. Although functional and taxonomic community metrics were largely unaffected by primary forest disturbance, they differed markedly between primary and secondary forests. However, our extinction scenarios demonstrated scale-dependence of species losses, whereby functional structure only eroded with species extinction at the local scale. Hence, we extend the spatial insurance hypothesis by demonstrating that landscape-scale functional redundancy offsets the impact of local species losses and confers community-level resistance to primary forest disturbance.

Interactions between plants and soil microbial communities underpin soil processes and forest ecosystem function, but the links between tree diversity and soil microbial diversity are poorly characterized. Differences in both the taxonomic and functional diversity of trees and microbes can shape soil nutrient status and carbon storage, but the stoichiometry of carbon and nutrients in the soil also influences resource availability to plant and microbial communities. Given the key role of resource availability in plant–soil interactions, we hypothesized that relationships between tree diversity metrics and soil bacterial or fungal diversity are mediated by soil stoichiometry. To test our hypothesis, we measured tree diversity metrics (tree species richness, functional trait diversity and functional trait composition) and soil stoichiometry in a temperate forest in China, and we determined soil microbial diversity by Illumina sequencing. We used structural equation models to assess the relationships between tree diversity metrics and soil bacterial or fungal diversity and to evaluate the influence of soil stoichiometry. Overall, microbial diversity was strongly related to soil stoichiometry, whereby fungal diversity was associated with high soil N/P ratios, whereas bacterial diversity was related to high soil C/P ratios. Soil bacterial and fungal diversity were more closely related to tree functional trait diversity and composition than to tree species richness, and the links between tree and soil microbial diversity were mediated by soil stoichiometry. The strong links between tree functional traits, soil stoichiometry and soil bacteria or fungi suggest that resource quality plays a key role in plant–microbial interactions. Our results highlight the importance of nutrient stoichiometry in linkages between tree functional diversity and soil microbial diversity.

Soil organic matter (SOM) in tropical forests is an important store of carbon (C) and nutrients. Although SOM storage could be affected by global changes via altered plant productivity, we know relatively little about SOM stabilisation and turnover in tropical forests compared to temperate systems. Here, we investigated changes in soil C and N within particle size fractions representing particulate organic matter (POM) and mineral-associated organic matter (MAOM) after 13 years of experimental litter removal (L−) and litter addition (L+) treatments in a lowland tropical forest. We hypothesized that reduced nitrogen (N) availability in L− plots would result in N-mining of MAOM, whereas long-term litter addition would increase POM, without altering the C:N ratio of SOM fractions. Overall, SOM-N declined more than SOM-C with litter removal, providing evidence of N-mining in the L− plots, which increased the soil C:N ratio. However, contrary to expectations, the C:N ratio increased most in the largest POM fraction, whereas the C:N ratio of MAOM remained unchanged. We did not observe the expected increases in POM with litter addition, which we attribute to rapid turnover of unprotected SOM. Measurements of ion exchange rates to assess changes in N availability and soil chemistry revealed that litter removal increased the mobility of ammonium-N and aluminium, whereas litter addition increased the mobility of nitrate-N and iron, which could indicate SOM priming in both treatments. Our study suggests that altered litter inputs affect multiple processes contributing to SOM storage and we propose potential mechanisms to inform future work.

The activity of soil microbes is strongly constrained by water availability. However, it is unclear how microbial activity responds to spatial and temporal changes in precipitation, particularly to long-term precipitation changes. To identify the spatiotemporal patterns of microbial responses to precipitation changes of differing durations, we conducted a meta-analysis of data from 95 field studies with drought treatments and 109 field studies with elevated precipitation treatments. Our results indicated that microbial biomass carbon (MBC) decreased by 17% under drought and increased by 18% under elevated precipitation. Across all studies, the phospholipid fatty acid (PLFA) biomarkers for fungi and bacteria decreased significantly under drought but increased under elevated precipitation. In addition, the negative effect of drought on MBC tended to be greater at sites with a high aridity index, but the effect of elevated precipitation on MBC did not differ among sites. More importantly, the responses of MBC, fungal and bacterial PLFA abundance did not vary with treatment duration under drought, but under elevated precipitation, they increased in the first five years of treatment and declined thereafter. These results are important for our prediction of microbial responses to long-term precipitation change, because they imply that microbes acclimate to long-term elevated precipitation.

Soil life supports the functioning and biodiversity of terrestrial ecosystems1, 2. Springtails (Collembola) are among the most abundant soil animals regulating soil fertility and flow of energy through above- and belowground food webs3–5. However, the global distribution of springtail diversity and density, and how these relate to energy fluxes remains unknown. Here, using a global dataset collected from 2,470 sites, we estimate total soil springtail biomass at 29 Mt carbon (threefold higher than wild terrestrial vertebrates6) and record peak densities up to 2 million individuals per m² in the Arctic. Despite a 20-fold biomass difference between tundra and the tropics, springtail energy use (community metabolism) remains similar across the latitudinal gradient, owing to the increase in temperature. Neither springtail density nor community metabolism were predicted by local species richness, which was highest in the tropics, but comparably high in some temperate forests and even tundra. Changes in springtail activity may emerge from latitudinal gradients in temperature, predation7, 8, and resource limitation7, 9, 10 in soil communities. Contrasting temperature responses of biomass, diversity and activity of springtail communities suggest that climate warming will alter fundamental soil biodiversity metrics in different directions, potentially restructuring terrestrial food webs and affecting major soil functions.

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.

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), β-1,4-glucosidase (BG), β-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.

Soils are a fundamental natural resource but intensifying demands and increasing soil degradation necessitate focussed research into the sustainable use of soils. Since soil functioning is critical for the operations and performance of multiple industries, businesses and municipalities, soil scientists need to actively engage with these bodies to orientate research goals towards stakeholder needs. To achieve this, stakeholder views about the current and potential contributions of soil science to different sectors need to be taken into account when setting the future research agenda. Here, we assessed whether the current and future research priorities of soil science match the needs of four major industrial and environmental sectors: agriculture, ecosystem services and natural resources, waste management, and water management. We used an online questionnaire, distributed to 192 organisations and via social media, to compare stakeholders' and scientists' perceptions of (a) the contributions of soil science to date, (b) the areas not currently served by soil science and (c) future research needs in soil science. Stakeholders generally rated the contributions of soil science to date as ‘great’ or ‘fundamental’, but scientists rated the contributions more highly. Respondents identified numerous areas that soil research has not yet sufficiently addressed, which were mostly sector-specific and often overlapped with perceived future research needs. Importantly, stakeholders' and scientists' views of future research priorities differed strongly within sectors, with the notable exception of agriculture, where views were generally consistent. We conclude that soil science may hold unexplored potential in several industrial and environmental sectors. We call for improved research communication and greater stakeholder involvement to shape the future soils research agenda and ensure the sustainable use of soils across multiple areas of society.

Aims: Aboveground plant litter inputs are important sources of soil carbon (C). We aimed to establish how experimentally altered litter inputs affect soil C to 1-m depth across different ecosystems, and over different timeframes. Methods: We performed a meta-analysis of 237 studies across 248 sites worldwide to assess the influence of treatment magnitude, treatment duration, initial soil C content, and climate on the response of soil C to altered aboveground litter inputs. Results: Overall, soil C content was lower under litter removal, but higher under litter addition compared to controls. The effects of litter manipulation were apparent throughout the soil profile and were related to treatment magnitude. Soil C content declined markedly with increasing duration of litter removal, whereas the positive effect of litter addition attenuated over time. Cropland management practices (bare fallow or additional straw incorporation) had similar effects on soil C to litter removal and addition treatments. Conclusions: Our study reveals rapid and consistent changes in soil C content with altered litter inputs and provides important insights into plant residue management to enhance soil C sequestration. We highlight the need for long-term experiments, with a greater focus on the processes underpinning soil C storage in different ecosystems.

Human-modified forests, including plantations and managed forest, will be a major component of tropical landscapes in the near future. To conserve biodiversity across modified tropical landscapes we must first understand what influences diversity in planted areas. We studied dung beetle communities in Eucalyptus plantations to assess the influence of local (canopy openness and soil texture)and landscape factors (surrounding native forest cover)on taxonomic and functional diversity, and to determine whether biodiversity in plantations is affected by timber production. Dung beetle community composition in Eucalyptus plantations was largely explained by the surrounding native forest cover, as Simpson's diversity and functional diversity (Rao's quadratic entropy)increased with the extent of native forest in buffer areas. However, the abundance of dung beetle species associated with native forest was not explained by any of the explanatory variables. The coarse sand content of the soil explained much of the functional similarity between plantations and native forests, as well as variation in dung beetle community structure. The total abundance of dung beetles in plantations increased with coarse sand content, whereas body mass declined, and dung beetle abundance and functional originality decreased with canopy openness. Timber production intensity did not explain the variation in any of the measured diversity parameters. If enhancing biodiversity in plantations is a management goal, then these results highlight the importance of restoring or retaining native forest areas in modified landscapes. They also suggest that integrated management could improve biodiversity in Eucalyptus plantations without reducing timber production. © 2019 Elsevier B.V.

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 mean 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.

Functional diversity is commonly used to assess the conservation value of ecosystems, but we have not yet established whether functional and taxonomic approaches are interchangeable or complementary to evaluate community dynamics over time and in response to disturbances. We used a five-year dataset of dung beetles from undisturbed forest, primary forest corridors, and Eucalyptus plantations to compare the sensitivity of conceptually equivalent metrics to temporal variation in different anthropogenic disturbances. We compared species richness with functional richness, Pielou’s evenness with functional evenness, and Simpson’s diversity with Rao’s quadratic entropy. We assessed the sensitivity of the metrics to anthropogenic changes. The indices showed complex patterns among habitat types, with with similar responses in some cases and not in other, and little incongruence between the pairs within the same year. The influence of disturbance on longer-term temporal variation over the five-year period revealed lower temporal variation in functional than taxonomic metrics. Both approaches showed greater variation in plantations compared to native forests. We evaluated the variation in taxonomic and functional metrics between consecutive years and among habitats. Most metrics showed similar shifts between years in all habitats, except for species and functional richness. We demonstrate that even conceptually similar indices may not provide similar information on dung beetles responses to disturbance. However, the differences between the indices can yield key insights about the drivers of change, especially over the long-term. It is important to use taxonomic and functional diversity in tandem to better understand community responses to environmental and anthropogenic changes.

Tropical forests on upland soils are assumed to be a methane (CH4) sink and a weak source of nitrous oxide (N2O), 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 N2O. Here, we measured CH4 and N2O 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 N2O fluxes were negligible during the dry season but increased markedly after the start of the wet season. By contrast, tree stembases emitted CH4 and N2O 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 N2O 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.

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.

Drought is a severe natural risk that increases drying-rewetting frequencies of soils. Yet, it remains largely unknown how forest ecosystems respond to dry-wet cycles, hampering our ability to evaluate the overall sink and source functionality for this large carbon pool. Recent investigations suggest that the release of soluble carbon via root exudation increases under drought, influencing soil carbon stabilization and mineralization. However, an integration of root exudation into the carbon allocation dynamics of drought stressed trees is missing. We hypothesized that roots in dry soil layers have a higher exudation rate than roots in more moist layers across different soil depths. Further, we tested if higher exudation rates under drought are attenuated by reduced root abundance in dry soils and if the fraction of root exudation from total carbon allocation increases with decreasing photosynthesis rates under drought. At the KROOF experimental site in southern Germany, where mature beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) trees were exposed to artificial drought stress for five consecutive growing seasons, we show that at the root level root exudation rate increases in drier soils. Especially roots in the upper soil profile and roots of spruce trees increased root exudation under drought. When scaled to whole tree level, we did not find differences in total exudation between drought stressed and control trees, indicating sustained root exudation at the tree level under drought. As photosynthesis rates and therefore total carbon assimilation was substantially reduced under drought (by 50 % in beech and almost 70 % in spruce), the fraction of root exudation from total assimilation slightly increased for drought stressed trees. Our results demonstrate that stimulation of root exudation rates with drought exists in natural temperate forest ecosystems but might be mitigated by reduced fine root abundance under drought. Nevertheless, increased exudation per root surface area will have localized impacts on rhizosphere microbial composition and activity especially in the topsoil exposed to more extreme dry-wet cycles. Finally, also the exudate composition can help to determine how priming of soil organic matter relates to belowground carbon allocation dynamics and to disclose processes of complementary species interaction and should be emphasised in future studies.

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., Resorption is a key plant nutrition strategy in old‐growth tropical forests. Trees with high foliar nutrient concentration reabsorb more nutrients. 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.

Plant species diversity affects carbon and nutrient cycling during litter decomposition, yet the generality of the direction of this effect and its magnitude remains uncertain. With a meta-analysis including 65 field studies across the Earth’s major forest ecosystems, we show here that decomposition was faster when litter was composed of more than one species. These positive biodiversity effects were mostly driven by temperate forests but were more variable in other forests. Litter mixture effects emerged most strongly in early decomposition stages and were related to divergence in litter quality. Litter diversity also accelerated nitrogen, but not phosphorus release, potentially indicating a decoupling of nitrogen and phosphorus cycling and perhaps a shift in ecosystem nutrient limitation with changing biodiversity. Our findings demonstrate the importance of litter diversity effects for carbon and nutrient dynamics during decomposition, and show how these effects vary with litter traits, decomposer complexity and forest characteristics.

The crucial role of tropical forests in the global carbon balance is underpinned by their extraordinarily high biomass and productivity, even though the majority of tropical forests grow on nutrient-poor soils. Nutrient cycling by litterfall has long been considered essential for maintaining high primary productivity in lowland tropical forests but few studies have tested this assumption experimentally. We review and synthesise findings from the Gigante Litter Manipulation Project (GLiMP), a long-term experiment in lowland tropical forest in Panama, Central America, in which litter has been removed from or added to large-scale plots for 15 years. We assessed changes in soil and litter nutrient concentrations in response to the experimental treatments and estimated nutrient return and nutrient use efficiency to indicate changes in nutrient cycling. The soil concentrations of most nutrients increased with litter addition and declined with litter removal. Litter removal altered nitrogen, potassium, manganese and zinc cycling, demonstrating the importance of litter inputs for maintaining the availability of these elements to plants. By contrast, litter addition only altered nitrogen cycling and, despite low concentrations of available soil phosphorus, the effects of litter manipulation on phosphorus cycling were inconsistent. We discuss potential mechanisms underlying the observed changes, and we emphasise the importance of decomposition processes in the forest floor for retaining nutrient elements, which partially decouples nutrient cycling from the mineral soil. Finally, by synthesising GLiMP studies conducted during 15 years of litter manipulation, we highlight key knowledge gaps and avenues for future research into tropical forest nutrient cycling.

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.

Regional factors, such as historical and contemporary climate conditions, and local factors, such as vegetation structural attributes, can influence current patterns of plant species richness but their relative roles remain unknown, particularly across forest strata. Here, we used a multi-scale survey of temperate forest plots across a large region of Northeast China to explore the relative importance of environmental factors (paleoclimate, contemporary climate, topography and anthropogenic disturbance) and forest structural attributes (stem abundance, stand basal area and tree size variation) on tree, shrub, and herb species richness. Although environmental and forest structural factors all played a role in explaining plant species richness patterns, their relative roles varied in direction and magnitude depending on forest stratum. Tree species richness increased with the magnitude of change in temperature since the Last Glacial Maximum (Anomaly) but declined with increasing magnitude of change in precipitation (Anomaly). By contrast, herb species richness declined with increasing Anomaly but increased with Anomaly, highlighting contrasting processes for tree and herb species richness driven by paleoclimate. Contemporary climate played a lesser role in explaining species richness, but tree species richness increased with diurnal temperature range, shrub species richness increased with the climatic moisture index, and herb species richness increased with both climatic variables. Herb species richness also increased with disturbance intensity, whereas tree and shrub richness declined. Overall, plant species richness increased significantly with all forest structural attributes, except for stem abundance, which had a negative effect on herb species richness, and forest structure mediated the linkages between plant species richness and disturbance or climate. The pronounced influence of paleoclimate on forest plant species richness highlights the potential threat of current climate change for forest diversity. Together, simultaneous consideration of past and current climate as well as forest structural attributes could improve our understanding of the complex mechanisms shaping patterns of plant species richness across forest strata., This work was supported by the National Natural Science Foundation of China (31961133027) the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB31030000) Key Research Program of Frontier Sciences (ZDBS-LY-DQC019), the LiaoNing Revitalization Talents Program (XLYC1807039) and K.C. Wong Education Foundation.

Disturbance can alter tree species and functional diversity in tropical forests, which in turn could affect carbon and nutrient cycling via the decomposition of plant litter. However, the influence of tropical tree diversity on forest floor organisms and the processes they mediate are far from clear. We investigated the influence of different litter mixtures on arthropod communities and decomposition processes in a 60-year-old lowland tropical forest in Panama, Central America. We used litter mixtures representing pioneer and old growth tree species in experimental mesocosms to assess the links between litter types, decomposition rates, and litter arthropod communities. Overall, pioneer species litter decomposed most rapidly and old growth species litter decomposed the slowest but there were clear non-additive effects of litter mixtures containing both functional groups. We observed distinct arthropod communities in different litter mixtures at 6 mo, with greater arthropod diversity and abundance in litter from old growth forest species. By comparing the decay of different litter mixtures in mesocosms and conventional litterbags, we demonstrated that our mesocosms represent an effective approach to link studies of litter decomposition and arthropod communities. Our results indicate that changes in the functional diversity of litter could have wider implications for arthropod communities and ecosystem functioning in tropical forests.

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 CO 2 (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.

Nitrogen (N) and phosphorus (P) are essential nutrients for plant metabolism, and their availability often limits primary productivity. Whereas the effects of N availability on photosynthetic capacity are well established, we still know relatively little about the effects of P availability at a foliar level, especially in P‐limited tropical forests. We examined photosynthetic capacity, leaf mass per area (LMA) and foliar P fractions in five woody plant species after 6 years of N and P fertilization in a lowland tropical forest. Foliar N:P ratios indicated P limitation of the unfertilized plants; accordingly, photosynthetic P‐use efficiency (PPUE) and LMA decreased with P addition, and foliar N and P concentrations increased, whereas N addition had little effect on measured foliar traits. However, P addition enhanced photosynthetic capacity only in one species and not in other four species. We then assessed plant acclimation to low P availability by quantifying four fractions of foliar P representing different functional pools: structural P, metabolic P (including inorganic P), nucleic acid P, and residual P. We found that P addition enhanced the concentrations of metabolic, structural, and nucleic acid P fractions in all species, but the magnitude of the effect was species‐specific. Our findings indicate that tropical species acclimate to low P availability by altering allocation of foliar P to meet the demand of P for photosynthesis. Importantly, species typical of lowland tropical forests in East Asia maintained their photosynthetic rate under low P availability. We conclude that P limitation of leaf photosynthetic capacity may not be as common as previously assumed due to plant acclimation mechanisms in low‐P tropical forests. Species‐specific strategies to allocate P to different foliar fractions represent a potentially important adaptive mechanism for plants in P‐limited systems.

Tropical reforestation initiatives are widely recognized as a key strategy for mitigating rising atmospheric CO2 concentrations. Although rapid tree growth in young secondary forests and plantations sequesters large amounts of carbon (C) in biomass, the choice of tree species for reforestation projects is crucial, as species identity and diversity affect microbial activity and soil C cycling via plant litter inputs. The decay rate of litter is largely determined by its chemical and physical properties, and trait complementarity of diverse litter mixtures can produce non-additive effects, which facilitate or delay decomposition. Furthermore, microbial communities may preferentially decompose litter from native tree species (homefield advantage). Hence, information on how different tree species influence soil carbon dynamics could inform reforestation efforts to maximize soil C storage. We established a decomposition experiment in Panama, Central America, using mesocosms and litterbags in monoculture plantations of native species (Dalbergia retusa Hemsl. and Terminalia amazonia J.F.Gmel., Exell) or teak (Tectona grandis L.f.) to assess the influence of different litter types and litter mixtures on soil C dynamics. We used reciprocal litter transplant experiments to assess the homefield advantage and litter mixtures to determine facilitative or antagonistic effects on decomposition rates and soil respiration in all plantation types. Although litter properties explained some of the variation in decomposition, the microclimate and soil properties in the plantations also played an important role. Microbial biomass C and litter decomposition were lower in Tectona than in the native plantations. We observed non-additive effects of mixtures with Tectona and Dalbergia litter on both decomposition and soil respiration, but the effect depended on plantation type. Further, there was a homefield disadvantage for soil respiration in Tectona and Terminalia plantations. Our results suggest that tree species diversity plays an important role in the resilience of tropical soils and that plantations with native tree species could help maintain key processes involved in soil carbon sequestration.