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Trichomes are epidermal outgrowths on plant shoots. Their roles in protecting plants against herbivores and in the biosynthesis of specialized metabolites have long been recognized. Recently, studies are increasingly showing that trichomes also play important roles in water absorption and metal detoxication, with these roles having important implications for ecology, the environment, and agriculture. However, these two functions of trichomes have been largely overlooked and much remains unknown. In this review, we show that the trichomes of 37 plant species belonging to 14 plant families are involved in water absorption, while the trichomes of 33 species from 13 families are capable of sequestering metals within their trichomes. The ability of trichomes to absorb water results from their decreased hydrophobicity compared to the remainder of the leaf surface as well as the presence of special structures for collecting and absorbing water. In contrast, the metal detoxication function of trichomes results not only from the good connection of their basal cells to the underlying vascular tissues, but also from the presence of metal-chelating ligands and transporters within the trichomes themselves. Knowledge gaps and critical future research questions regarding these two trichome functions are highlighted. This review improves our understanding on trichomes.

Tree size shapes forest carbon dynamics and determines how trees interact with their environment, including a changing climate. Here, we conduct the first global analysis of among-site differences in how aboveground biomass stocks and fluxes are distributed with tree size. We analyzed repeat tree censuses from 25 large-scale (4-52 ha) forest plots spanning a broad climatic range over five continents to characterize how aboveground biomass, woody productivity, and woody mortality vary with tree diameter. We examined how the median, dispersion, and skewness of these size-related distributions vary with mean annual temperature and precipitation. In warmer forests, aboveground biomass, woody productivity, and woody mortality were more broadly distributed with respect to tree size. In warmer and wetter forests, aboveground biomass and woody productivity were more right skewed, with a long tail towards large trees. Small trees (1-10 cm diameter) contributed more to productivity and mortality than to biomass, highlighting the importance of including these trees in analyses of forest dynamics. Our findings provide an improved characterization of climate-driven forest differences in the size structure of aboveground biomass and dynamics of that biomass, as well as refined benchmarks for capturing climate influences in vegetation demographic models.

Diversity–biomass relationships (DBRs) often vary with spatial scale in terrestrial ecosystems, but the mechanisms driving these scale-dependent patterns remain unclear, especially for highly heterogeneous forest ecosystems. This study explores how mutualistic associations between trees and different mycorrhizal fungi, i.e., arbuscular mycorrhizal (AM) vs. ectomycorrhizal (EM) association, modulate scale-dependent DBRs. We hypothesized that in soil-heterogeneous forests with a mixture of AM and EM tree species, (i) AM and EM tree species would respond in contrasting ways (i.e., positively vs. negatively, respectively) to increasing soil fertility, (ii) AM tree dominance would contribute to higher tree diversity and EM tree dominance to greater standing biomass, and that as a result (iii) mycorrhizal associations would exert an overall negative effect on DBRs across spatial scales. To empirically test these hypotheses, we collected detailed tree distribution and soil information (e.g., nitrogen, phosphorus, organic matter, pH) from seven temperate and subtropical AM–EM mixed forest megaplots (16–50 ha). Using a spatial codispersion null model and structural equation modeling, we identified the relationships among AM or EM tree dominance, soil fertility, tree species diversity, and biomass and, thus, DBRs across 0.01- to 1-ha scales. We found the first evidence overall supporting the three aforementioned hypotheses in these AM–EM mixed forests: (i) In most forests, with increasing soil fertility, tree communities changed from EM-dominated to AM-dominated; (ii) increasing AM tree dominance had an overall positive effect on tree diversity and a negative effect on biomass, even after controlling for soil fertility and number of trees. Together, (iii) the changes in mycorrhizal dominance along soil fertility gradients weakened the positive DBR observed at 0.01- to 0.04-ha scales in nearly all forests and drove negative DBRs at 0.25- to 1-ha scales in four out of seven forests. Hence, this study highlights a soil-related mycorrhizal dominance mechanism that could partly explain why, in many natural forests, biodiversity–ecosystem functioning (BEF) relationships shift from positive to negative with increasing spatial scale.

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)

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.

International audience; Forest dynamics are shaped by both abiotic and biotic factors. Trees associating with different types of mycorrhizal fungi differ in nutrient use and dominate in contrasting environments, but it remains unclear whether they exhibit differential growth responses to local abiotic and biotic gradients where they co-occur. We used 9-year tree census data in a 25-ha old-growth temperate forest in Northeast China to examine differences in tree growth response to soil nutrients and neighborhood crowding between tree species associating with arbuscular mycorrhizal (AM), ectomycorrhizal (EM), and dual-mycorrhizal (AEM) fungi. In addition, we tested the role of individual-level vs species-level leaf traits in capturing differences in tree growth response to soil nutrients and neighborhood crowding across mycorrhizal types. Across 25 species, soil nutrients decreased AM tree growth, while neighborhood crowding reduced both AM and EM tree growth, and neither soil nor neighbors impacted AEM tree growth. Across mycorrhizal types, individual-level traits were stronger predictors of tree growth than species-level traits. However, most traits poorly mediated tree growth response to soil nutrients and neighborhood crowding. Our findings indicate that mycorrhizal types strongly shape differences in tree growth response to local soil and crowding gradients, and suggest that including plant-mycorrhizae associations in future work offers great potential to improve our understanding of forest dynamics.

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.

Lianas are woody plants that require external support to reach the canopy. They are expanding in forests worldwide, possibly due to climate change and forest disturbance. Most studies on lianas have been conducted in tropical forests. Lianas are less explored in subtropical forests. We aimed to document the density and diversity of lianas, to test how habitat condition and the distribution of tree species affect the distributions of lianas based on data from a fully mapped 20 ha plot in subtropical China. We analyzed habitat association by fitting a generalized linear model with family-level liana abundance as response variable and family identity, and its interaction terms with topographic variables (slope, convexity, elevation, and sin(aspect)), as explanatory variables. We focused on the spatial associations of three liana species and 82 tree species with ≥100 individuals using the pair correlation function and redundancy analysis. We found a total of 1305 lianas, falling into 26 species, and 16 families, in the 20 ha plot. They accounted for 1.5% of individuals, 11.7% of species, and 0.4% of total basal area of woody plants in the plot. There were large variations in distributions of liana with respect to the four topographic variables among families, contrasting with former findings suggesting that lianas favor dry and hot habitats. The three most abundant liana species showed non-random associations with tree species, and they tended to positively associate with similar tree species but negatively associate with different tree species. The distribution of tree species explained 21.8% of variance in liana distribution. Our study suggested that both habitat conditions and tree composition intervene in determining liana distributions and that habitat heterogeneity may be a mechanism for liana diversity maintenance. Our study provides a basic understanding of liana diversity and distribution in this subtropical forest and contributes to future planning of liana studies and diversity conservation in subtropical forests under climate change.

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.

The growth and survival of individual trees determine the physical structure of a forest with important consequences for forest function. However, given the diversity of tree species and forest biomes, quantifying the multitude of demographic strategies within and across forests and the way that they translate into forest structure and function remains a significant challenge. Here, we quantify the demographic rates of 1961 tree species from temperate and tropical forests and evaluate how demographic diversity (DD) and demographic composition (DC) differ across forests, and how these differences in demography relate to species richness, aboveground biomass (AGB), and carbon residence time. We find wide variation in DD and DC across forest plots, patterns that are not explained by species richness or climate variables alone. There is no evidence that DD has an effect on either AGB or carbon residence time. Rather, the DC of forests, specifically the relative abundance of large statured species, predicted both biomass and carbon residence time. Our results demonstrate the distinct DCs of globally distributed forests, reflecting biogeography, recent history, and current plot conditions. Linking the DC of forests to resilience or vulnerability to climate change, will improve the precision and accuracy of predictions of future forest composition, structure, and function.

In temperate forests, autumn leaf phenology signals the end of leaf growing season and shows large variability across tree-crowns, which importantly mediates photosynthetic seasonality, hydrological regulation, and nutrient cycling of forest ecosystems. However, critical challenges remain with the monitoring of autumn leaf phenology at the tree-crown scale due to the lack of spatially explicit information for individual tree-crowns and high (spatial and temporal) resolution observations with nadir view. Recent availability of the PlanetScope constellation with a 3 m spatial resolution and near-daily nadir view coverage might help address these observational challenges, but remains underexplored. Here we developed an integration of PlanetScope with drone observations for improved monitoring of crown-scale autumn leaf phenology in a temperate forest in Northeast China. This integration includes: 1) visual identification of individual tree-crowns (and species) from drone observations; 2) extraction of time series of PlanetScope vegetation indices (VIs) for each identified tree-crown; 3) derivation of three metrics of autumn leaf phenology from the extracted VI time series, including the start of fall (SOF), middle of fall (MOF), and end of fall (EOF); and 4) accuracy assessments of the PlanetScope-derived phenology metrics with reference from local phenocams. Our results show that (1) the PlanetScope-drone integration captures large inter-crown phenological variations, with a range of 28 days, 25 days, and 30 days for SOF, MOF, and EOF, respectively, (2) the extracted crown-level phenology metrics strongly agree with those derived from local phenocams, with a root-mean-square-error (RMSE) of 4.1 days, 3.0 days and 5.4 days for SOF, MOF, and EOF, respectively, and (3) PlanetScope maps large variations in autumn leaf phenology over the entire forest landscape with spatially explicit information. These results demonstrate the ability of our proposed method in monitoring the large spatial heterogeneity of crown-scale autumn leaf phenology in the temperate forest, suggesting the potential of using high-resolution satellites to advance crown-scale phenology studies over large geographical areas.

Congeneric species are critical for understanding the underlying ecological mechanisms of biodiversity maintenance. Ecological mechanisms such as conspecific negative density dependence, species differences in life-history stages related to habitat preference, and limiting similarity are known to influence plant fitness, thereby influencing species coexistence and biodiversity. However, our understanding of these phenomena as they apply to coexistence among coniferous species is limited. We studied two congeneric

Large-diameter trees regulate forest diversity, structure and aboveground biomass (AGB), but the mechanisms whereby they control forest processes remain understudied, especially in early successional forests. We used 1,956 0.16 ha plots from the Korean National Forest Inventory from mostly 20–50 years old stands (biomass accumulation phase) in closed-canopy temperate forests, with 236 species of woody plants and 391,543 individual stems ≥ 6 cm diameter at breast height. Based hereon, we analyzed the effects of large-diameter trees on aboveground biomass (AGB) in the overstory, understory, and the two combined (total vegetation). We also considered the effects of species and functional diversity, functional dominance of traits, stem density, and abiotic drivers (i.e., topographic, climatic and edaphic variables) interacting with large-diameter trees on AGB. We performed model averaging approaches with backward stepwise regressions and piecewise structural equation modeling to quantify and compare the effects of large trees vs. other biotic and abiotic drivers. Overall, large-diameter trees had a dominant effect on AGB compared with remaining tree attributes and abiotic drivers for both the overstory and whole community; however, they did not strongly influence the understory. Large-diameter trees also modulated the strength of the direct effects of abiotic drivers, particularly soil fertility, on AGB, as well as indirect effects via regulating the attributes of smaller-diameter trees. Our study provides new insights into the mechanisms associated with self-thinning and resource availability whereby large-diameter trees drive high AGB in succession forests. We also show that the effect of large-diameter trees is forest stratum-dependent across different types of temperate forests. This study emphasizes the importance of large-diameter trees in determining the structure and function in early successional forests, and the need for conservation and management actions to protect and promote these keystone organisms.

A prominent tree species coexistence mechanism suggests host-specific natural enemies inhibit seedling recruitment at high conspecific density (negative conspecific density dependence). Natural-enemy-mediated conspecific density dependence affects numerous tree populations, but its strength varies substantially among species. Understanding how conspecific density dependence varies with species’ traits and influences the dynamics of whole communities remains a challenge. Using a three-year manipulative community-scale experiment in a temperate forest, we show that plant-associated fungi, and to a lesser extent insect herbivores, reduce seedling recruitment and survival at high adult conspecific density. Plant-associated fungi are primarily responsible for reducing seedling recruitment near conspecific adults in ectomycorrhizal and shade-tolerant species. Insects, in contrast, primarily inhibit seedling recruitment of shade-intolerant species near conspecific adults. Our results suggest that natural enemies drive conspecific density dependence in this temperate forest and that which natural enemies are responsible depends on the mycorrhizal association and shade tolerance of tree species., The Janzen-Connell hypothesis posits that seedlings may be less likely to establish near conspecifics due to shared natural enemies. Here, Jia et al. show that tree species traits determine whether fungal pathogens or insect herbivores inhibit seedling recruitment and survival in a temperate forest.

International audience; The relationships between different aspects of diversity (taxonomic, structural and functional) and the aboveground biomass (AGB) as a major component of global carbon balance have been studied extensively but rarely under the simultaneous influence of forest dieback and management. In this study, we investigate the relationships between taxonomic, functional and structural diversity of woody species (trees and shrubs) and AGB along a gradient of dieback intensity (low, moderate, high and no dieback as control) under two contrasted management conditions (protection by central government vs. traditional management by natives) in a semi-arid oak Quercus brantii Lindl.) forest ecosystem. AGB was estimated and taxonomic diversity, community weighted average (CWM) and functional divergence indices were produced. We found that the aerial biomass was significantly higher in the intensively used area (14.57 (±1.60) t/hm2) than in the protected area (8.70 (±1.05) t/hm2) due to persistence of some large trees but with decreasing values along the dieback intensity gradient in both areas. CWM of height (H), leaf nitrogen content (LNC) and leaf dry matter content (LDMC) were also higher in the traditional managed area than in the protected area. In contrast, in the protected area, the woody species diversity was higher and the inter-specific competition was more intense, explaining a reduced H, biomass and LDMC. Contrary to the results of CWM, none of the functional diversity traits (FDvar) was affected by dieback intensity and only FDvar values of LNC, leaf phosphorus content (LPC) and LDMC were influenced by management. We also found significantly positive linear relationships of AGB with CWM and FDvar indices in the protected area, and with taxonomic and structural diversity indices in the traditional managed area. These results emphasize that along a dieback intensity gradient, the leaf functional traits are efficient predictors in estimating the AGB in protected forests, while taxonomic and structural indices provide better results in forests under a high human pressure. Finally, species identity of the dominant species (i.e., Brant’s oak) proves to be the main driver of AGB, supporting the selection effect hypothesis.

Soil’s water-physical properties support essential soil water retention functions for driving water distribution and availability, which is vital for plant growth and biogeochemical cycling. However, the question concerning how tree compositions and their interactions with other abiotic factors modulate soil’s water-physical properties in disturbed forests remains poorly understood. Based on observational data from nine permanent forest sites (18,747 trees and 210 plots) in the northeast of China, where forests once undergone three different levels of anthropogenic logging disturbance, we evaluated how multiple biotic (i.e., tree diversity and functional trait composition) and abiotic (soil texture and soil organic carbon) factors influence water-physical properties (i.e., in terms of soil capillary water retention (WC) and soil saturated water retention (WS)) in temperate forests. We found that the impacts of logging disturbance on soil water-physical properties were associated with improved tree diversity, acquisitive functional traits, and SOC. These associated attributes were also positively related to WC and WS, while there was no significant effect from soil texture. Moreover, disturbance indirectly affected soil water-physical properties mainly by functional traits and SOC, as acquisitive functional traits significantly mediate the effect from disturbance on WC and SOC mediates the influence from disturbance on WS. Finally, our results emphasize the potential relationships of tree composition with SOC and soil water retention as compared with soil texture and hence suggest that plants can actively modulate their abiotic contexts after disturbance, which is meaningful for understanding forest health and resistance.

El Niño-Southern Oscillation (ENSO), the variation between anomalously cold (La Niña) and warm conditions (El Niño), is one of the most prominent large-scale climate patterns with worldwide effects. Elevated seed and leaf fall has been found at the positive phase of ENSO (El Niño) in tropical forests. However, how seed and leaf fall respond to ENSO at species level is understudied, especially in temperate forests. In this study, we monitored seed and leaf fall at the species-level at 150 points across a 25-ha temperate forest in northeastern China over a span of 12 years. Using time series and wavelet analyses, we assessed three hypotheses: 1) temperate tree species' seed and leaf fall are strongly, but differently, correlated with ENSO and, 2) community synchrony in seed and leaf occurred both at seasonal and ENSO scales; finally, 3) local climatic modulated the effects of ENSO on seed and leaf fall. We found that ENSO was significantly correlated with seed and leaf fall of all species, although correlation strength varied across species (r = 0.206-0.658). Specifically, ENSO indices (ENSO12 or ENSO34) accounted for the most variation in seed and leaf fall of Acer pseudo-sieboldianum (40 % and 34 %, respectively) and ranged 4 %-31 % in all other species. Leaf fall was synchronous with ENSO cycles with a period of 2-7 years, but community synchrony of seed fall was only detected at seasonal scales. ENSO influenced seed fall of Fraxinus mandshurica and Tilla amurensis by mediating rainfall and relative humidity, respectively, highlighting the interactive effects of local climate and ENSO. Our findings highlight the potential effects of ENSO on ecosystems outside of tropical regions and improve our ability to predict regeneration dynamics and nutrient cycling of temperate forests under the context of global change.

Global development has been heavily reliant on the overexploitation of natural resources since the Industrial Revolution. With the extensive use of fossil fuels, deforestation, and other forms of land-use change, anthropogenic activities have contributed to the ever-increasing concentrations of greenhouse gases (GHGs) in the atmosphere, causing global climate change. In response to the worsening global climate change, achieving carbon neutrality by 2050 is the most pressing task on the planet. To this end, it is of utmost importance and a significant challenge to reform the current production systems to reduce GHG emissions and promote the capture of CO2 from the atmosphere. Herein, we review innovative technologies that offer solutions achieving carbon (C) neutrality and sustainable development, including those for renewable energy production, food system transformation, waste valorization, C sink conservation, and C-negative manufacturing. The wealth of knowledge disseminated in this review could inspire the global community and drive the further development of innovative technologies to mitigate climate change and sustainably support human activities., Graphical abstract, Public summary • Carbon neutrality may be achieved by reforming current global development systems to minimize greenhouse gas emissions and increase CO2 capture • Harnessing the power of renewable and carbon-neutral resources to produce energy and other fossil-based alternatives may eliminate our dependence on fossil fuels • Protecting natural carbon sinks and promoting CO2 capture, utilization, and storage are conducive to mitigating climate change • This review presents the current state, opportunities, challenges, and perspectives of technologies related to achieving carbon neutrality

Mechanisms such as conspecific negative density dependence (CNDD) and niche partitioning have been proposed to explain species coexistence and community diversity. However, as a potentially important axis of niche partitioning, the role of interannual climate variability in driving local community dynamics remains largely unknown. Here we used a 15-year monitoring data set of more than 53,000 seedlings in a temperate forest to examine (1) what are the relative effects of interannual climate variability, biotic interactions, and habitat conditions on seedling survival; (2) how the effects of biotic interactions change with interannual climate variability, and habitat conditions; and (3) whether the impacts of interannual climate variability, biotic interactions, and habitat conditions differ with plant traits. Interannual climate variability accounted for the most variation in seedling survival at the community level, followed by biotic interactions, and habitat conditions. Increased snowpack and decreased minimum temperature during the non-growing season had positive effects on seedling survival. Effects of conspecific neighbor density were weakened in higher snowpack, effective accumulated temperature, elevation, and soil-resource gradient, but were intensified with increased ultraviolet radiation, maximum precipitation, minimum temperature, and soil moisture. In addition, the relative importance of interannual climate variability versus biotic interactions differed depending on species-trait groups. Specifically, biotic interactions for gravity-dispersed species had a larger effect size in affecting seedling survival than other trait groups. Also, gravity-dispersed species experienced a stronger CNDD than wind-dispersed, probably because wind-dispersed seedlings rarely had adult conspecifics nearby. We found that interannual climate variability was most strongly associated with seedling survival, but the magnitude of climatic effects varied among species-trait groups. Interannual climate variability may act as an inhibitor or accelerator to density-dependent interactions and should be accounted for in future studies, as both a potential direct and indirect factor in understanding the diversity of forest communities.

Tree-fungal symbioses are increasingly recognized to affect soil nitrogen (N) transformations, yet the role of soil microbes in the process is largely unclear. Soil microbes directly interact with trees and are a primary driver of many N transformation processes. Here, we explored the linkage among tree mycorrhizal associations, soil microbes and N transformation rates in a temperate forest of Northeast China. Across a gradient of increasing ectomycorrhizal (ECM) tree dominance, we measured soil acid-base chemistry, bacterial and fungal abundances, N-hydrolyzing enzyme activities, abundances and community composition of ammonia-oxidizing archaea (AOA) and bacteria, and net N mineralization and net nitrification rates. Results showed that soil pH, exchangeable base cations, inorganic N concentrations and N transformation rates decreased with increasing ECM tree dominance. The ECM tree dominance was negatively related to soil bacterial and AOA amoA gene abundances, and positively to soil fungal abundances and β-N-acetylglucosaminidase activities. These shifts in soil microbial abundances and enzyme activities along the mycorrhizal gradient were linked with the increase in soil acidity with increasing ECM tree dominance. Structural equation models revealed that ECM tree dominance was not directly related to N transformation rates, but indirectly to net N mineralization rates via affecting bacterial and fungal abundances, and indirectly to net nitrification rates via influencing AOA amoA gene abundances. Collectively, our results indicate that soil microbes provide a mechanistic link between mycorrhizal associations and soil N transformations, and suggest that shifts in forest mycorrhizal associations under global change could have profound consequences for biogeochemical cycling of temperate forests.

International audience; The plant-soil interactions may drive the diversity and functioning of forests, but we do not fully understand how interrelationships between plant and soil compartments are underlined by multiple ecological mechanisms. Here, we hypothesize that positive plant-soil interactions enhance biodiversity and functioning in a temperate forest. To do so, we tested the relationships between plant diversity (i.e., tree and herb species richness) and functions (i.e., coarse woody productivity and litterfall productivity), and soil diversity (i.e. bacterial, fungal and nematode) and functions (i.e. soil nutrient and carbon stock), and their interrelationships in a temperate forest in northeast China. The positive relationship between diversity and functioning was predominant within plant and soil compartments, and hence, provide support to the niche complementarity effect. As such, the positive interrelationships between the diversity of soil and plant compartments provide support to the positive plant-soil interactions. Tree species diversity was positively related with herb species diversity and coarse-woody productivity. Importantly, tree species diversity had pronounced positive effect on soil biodiversity resulting in increased soil carbon stocks, indicating that tree species diversity effect matters for linking positive interrelationships between plant and soil compartments of a temperate forest. This study shows that tree diversity effect is the main regulating biotic mechanism for linking the positive connections between plant and soil compartments of a temperate forest, and hence, the niche complementarity effect can enhance forest functioning through positive interactions on resource supply. We argue that linking the multiple key functions and diversity indices of forests can enhance our knowledge on the main influential factors and underlying ecological mechanisms.

Seed distribution and deposition patterns around parent trees are strongly affected by functional traits and therefore influence the development of plant communities. To assess the limitations of seed dispersal and the extent to which diaspore and neighbouring parental traits explain seed rain, we used a 9-year seed data set based on 150 seed traps in a 25-ha area of a temperate forest in the Changbai Mountain. Among 480,598 seeds belonging to 12 families, 17 genera, and 26 species were identified, only 54% of the species with mature trees in the community were represented in seeds collected over the 9 years, indicating a limitation in seed dispersal. Understory species were most limited; overstory species were least limited. Species with wind-dispersed seed had the least limitation, while the lowest similarity in species richness was for animal-dispersed species followed by gravity-dispersed species; fleshy-fruited species had stronger dispersal limitations than dry-fruited species. Generalized linear mixed models showed that relative basal area had a significant positive effect on seed abundance in traps, while the contribution of diaspore traits was low for nearly all groups. These results suggest that tree traits had the strongest contribution to seed dispersal and deposition for all functional groups examined here. These findings strengthen the knowledge that tree traits are key in explaining seed deposition patterns, at least at the primary dispersal stage. This improved knowledge of sources of seeds that are dispersed could facilitate greater understanding of seedling and community dynamics in temperate forests.

In natural communities, species abundance has intrinsic associations with spatial pattern of species and interspecific interaction. However, how these associations contribute to species coexistence remains largely controversial. We expect that common species suffer stronger interspecific competition, which limits their population growth rates. Rare species tend to increase strength of intraspecific competition relative to interspecific competition via stronger aggregation than common species, which facilitates their coexistence with other species. To test this hypothesis, we examined interspecific interaction and conspecific aggregation for adult trees (DBH≥ 10 cm) in a 25-ha forest dynamics plot in Changbaishan, northeast China, and further evaluated their relationships with species abundance. Our results showed that interspecific spatial associations were mostly negative, and common species showed more significant repulsion than rare species, which indicated obvious abundance-asymmetry in interspecific association. In addition, the abundance-dependent conspecific aggregation contributed to abundance-independent local dominance among species via adjusting the proportion of conspecific vs. heterospecific trees at local scale (~5 m), which would facilitate the coexistence between rare and common species. These results suggest that the strength of interspecific interactions in community-level mainly depends on common species, and the prevalent conspecific aggregation in plant communities is crucial to the survival of rare species and long-term species coexistence. Our study uncovers the underlying coexistence mechanisms about the abundance-dependent spatial distribution of species, and provides further insights into the community assembly rules in temperate forests.

Questions: All else being equal, populations of dioecious species with a 50:50 sex ratio have only half the effective reproductive population size of bisexual species of equal abundance. Consequently, there is a need to explain how dioecious and bisexual species coexist. Increased mean individual seed mass, fecundity, and population density have all been proposed as attributes of unisexual individuals or populations that may contribute to the persistence or resilience of dioecious species. To date, no studies have compared sympatric dioecious and cosexual species with respect to all three components of fitness. In this study, we sought evidence for these compensatory advantages (higher seed mass, greater seed production per unit basal area, and higher population density) in dioecious species. Location: Five 20–25ha forest dynamic plots spanning a latitudinal gradient in China, including two temperate, two subtropical, and one tropical forest. Methods: We used a phylogenetically corrected generalized linear modelling approach to assess the phylogenetic dependence and joint evolution of sexual system, seed mass and production, and ecological abundances among 48–333 species and 32,568–136,237 individuals per forest. Results: Across all five forests, we detected no consistent advantage for dioecious relative to sympatric cosexual species with respect to mean individual seed mass, seed production or the density of stems in any size class. Conclusions: Our study suggests that seed traits may provide compensatory mechanisms in some forests, but most often the coexistence of sexual systems cannot be explained by advantages of dioecy related to seed quality and demographic parameters. Future investigations of the factors that promote coexistence may increase our understanding by expanding the search to include attributes such as lifespan and tolerance or resistance to herbivores.

Forest dynamics are shaped by both abiotic and biotic factors. Trees associating with different types of mycorrhizal fungi differ in nutrient use and dominate in contrasting environments, but it remains unclear whether they exhibit differential growth responses to local abiotic and biotic gradients where they co-occur. We used 9-year tree census data in a 25-ha old-growth temperate forest in Northeast China to examine differences in tree growth response to soil nutrients and neighborhood crowding between tree species associating with arbuscular mycorrhizal (AM), ectomycorrhizal (EM), and dual-mycorrhizal (AEM) fungi. In addition, we tested the role of individual-level vs species-level leaf traits in capturing differences in tree growth response to soil nutrients and neighborhood crowding across mycorrhizal types. Across 25 species, soil nutrients decreased AM tree growth, while neighborhood crowding reduced both AM and EM tree growth, and neither soil nor neighbors impacted AEM tree growth. Across mycorrhizal types, individual-level traits were stronger predictors of tree growth than species-level traits. However, most traits poorly mediated tree growth response to soil nutrients and neighborhood crowding. Our findings indicate that mycorrhizal types strongly shape differences in tree growth response to local soil and crowding gradients, and suggest that including plant-mycorrhizae associations in future work offers great potential to improve our understanding of forest dynamics.

International audience; High species diversity is generally thought to be a requirement for sustaining forest multifunctionality. However, the degree to which the relationship between species-, structural-, and trait-diversity of forests and multifunctionality depend on the context (such as stand age or abiotic conditions) is not well studied. Here, we hypothesized that context-dependency of tree species diversity, functional trait composition and stand structural attributes promote temperate forest multifunctionality including above- and below-ground multiple and single functions. To do so, we used repeated forest inventory data, from temperate mixed forests of northeast China, to quantify two above-ground (i.e. coarse woody productivity and wild edible plant biomass), five below-ground (i.e. soil organic carbon, total soil nitrogen, potassium, phosphorus and sulfur) functions, tree species diversity, individual tree size variation (CVDBH) and functional trait composition of specific leaf area (CWMSLA) as well as stand age and abiotic conditions. We found that tree species diversity increased forest multifunctionality and most of the single functions. Below-ground single and multifunctionality were better explained by tree species diversity. In contrast, above-ground single and multifunctionality were better explained by CVDBH. However, CWMSLA was also an additional important driver for maintaining above- and below-ground forest multifunctionality through opposing plant functional strategies. Stand age markedly reduced forest multifunctionality, tree species diversity and CWMSLA but substantially increased CVDBH. Below-ground forest multifunctionality and tree species diversity decreased while above-ground forest multifunctionality increased on steep slopes. These results highlight that context-dependency of forest diversity attributes might regulate forest multifunctionality but may not have a consistent effect on above-ground and below-ground forest multifunctionality due to the fact that those functions were driven by varied functional strategies of different plant species. We argue that maximizing forest complexity could act as a viable strategy to maximizing forest multifunctionality, while also promoting biodiversity conservation to mitigate climate change effects.

Background and aims Soil water storage capacity acts as a vital forest function to intercept rainfall and retain water for plant growth processes. However, whether or how plant functional trait diversity and composition regulate soil water storage capacity remains poorly understood. Methods Structural equation modeling (SEM) was used to detect the direct and indirect effects of multiple biotic (i.e., functional trait composition and functional diversity) and abiotic (topography and soil organic carbon) factors on soil water storage capacity, i.e., in terms of soil capillary water storage content (CW), soil non-capillary water storage content (NCW), and soil saturated water storage content (TSW), in temperate forests recovering from different logging disturbance intensity levels. Results The community-weighted mean of specific leaf area (CWMSLA) increased CW but decreased NCW directly, whereas improved NCW and TSW indirectly via soil organic carbon. Disturbance influenced soil water storage capacity mainly in indirect ways via promoting CWMSLA and soil organic carbon. Elevation increased NCW and TSW but decreased CW directly, and it also had indirect effects on soil water storage capacity via decreasing CWMSLA and soil organic carbon. Moreover, soil organic carbon influenced NCW and TSW directly or mediated the effects of elevation, disturbance, and CWMSLA on soil water storage capacity. Conclusions The quick return on investments trait of CWMSLA shows a positive effect on soil water storage capacity (CW and TSW), supporting the mass ratio mechanism in temperate forests recovering from disturbances. Soil organic carbon also presents additional importance to soil water storage capacity.

International audience; Biodiversity plays a fundamental role in provisioning and regulating forest ecosystem functions and services. Above-ground (plants) and below-ground (soil microbes) biodiversity could have asynchronous change paces to human-driven land-use impacts. Yet, we know very little how they affect the provision of multiple forest functions related to carbon accumulation, water retention capacity and nutrient cycling simultaneously (i.e. ecosystem multifunctionality; EMF). We used a dataset of 22,000 temperate forest trees from 260 plots within 11 permanent forest sites in Northeastern China, which are recovering from three post-logging disturbances. We assessed the direct and mediating effects of multiple attributes of plant biodiversity (taxonomic, phylogenetic, functional and stand structure) and soil biodiversity (bacteria and fungi) on EMF under the three disturbance levels. We found the highest EMF in highly disturbed rather than undisturbed mature forests. Plant taxonomic, phylogenetic, functional and stand structural diversity had both positive and negative effects on EMF, depending on how the EMF index was quantified, whereas soil microbial diversity exhibited a consistent positive impact. Biodiversity indices explained on average 45% (26%–58%) of the variation in EMF, whereas climate and disturbance together explained on average 7% (0.4%–15%). Our result highlighted that the tremendous effect of biodiversity on EMF, largely overpassing those of both climate and disturbance. While above- (β = 0.02–0.19) and below-ground (β = 0.16–0.26) biodiversity had direct positive effects on EMF, their opposite mediating effects (β = −0.22 vs. β = 0.35 respectively) played as divergent pathways to human disturbance impacts on EMF. Our study sheds light on the need for integrative frameworks simultaneously considering above- and below-ground attributes to grasp the global picture of biodiversity effects on ecosystem functioning and services. Suitable management interventions could maintain both plant and soil microbial biodiversity, and thus guarantee a long-term functioning and provisioning of ecosystem services in an increasing disturbance frequency world.

© 2021 The Authors., Aims: Understanding fine-grain diversity patterns across large spatial extents is fundamental for macroecological research and biodiversity conservation. Using the GrassPlot database, we provide benchmarks of fine-grain richness values of Palaearctic open habitats for vascular plants, bryophytes, lichens and complete vegetation (i.e., the sum of the former three groups). Location: Palaearctic biogeographic realm. Methods: We used 126,524 plots of eight standard grain sizes from the GrassPlot database: 0.0001, 0.001, 0.01, 0.1, 1, 10, 100 and 1,000 m and calculated the mean richness and standard deviations, as well as maximum, minimum, median, and first and third quartiles for each combination of grain size, taxonomic group, biome, region, vegetation type and phytosociological class. Results: Patterns of plant diversity in vegetation types and biomes differ across grain sizes and taxonomic groups. Overall, secondary (mostly semi-natural) grasslands and natural grasslands are the richest vegetation type. The open-access file ”GrassPlot Diversity Benchmarks” and the web tool “GrassPlot Diversity Explorer” are now available online (https://edgg.org/databases/GrasslandDiversityExplorer) and provide more insights into species richness patterns in the Palaearctic open habitats. Conclusions: The GrassPlot Diversity Benchmarks provide high-quality data on species richness in open habitat types across the Palaearctic. These benchmark data can be used in vegetation ecology, macroecology, biodiversity conservation and data quality checking. While the amount of data in the underlying GrassPlot database and their spatial coverage are smaller than in other extensive vegetation-plot databases, species recordings in GrassPlot are on average more complete, making it a valuable complementary data source in macroecology., GrassPlot development has been supported by the Bavarian Research Alliance (BayIntAn_UBT_2017_58), the Eurasian Dry Grassland Group (EDGG) and the International Association for Vegetation Science (IAVS); IB, CorM, JAC, IGM, DGM, MHe, DL and MTo were supported by the Basque Government (IT936‐16); CorM, IAx, MCh, JDa, PD, MHá, ZL, ZPr, EŠ and LT were supported by the Czech Science Foundation (19‐28491X); TR was supported by the Estonian Research Council (PUT1173); RJP was funded by the Strategic Research Programme of the Scottish Government’s Rural and Environmental Science and Analytical Services Division”; SBa was supported by the GINOP‐2.3.2‐15‐2016‐00019 project; GFi was partially supported by the MIUR initiative “Department of excellence” (Law 232/2016)"; BJA was funded by the Spanish Research Agency (grant AEI/ 10.13039/501100011033); AK, VB, IM, DS, IV and DV were supported by the National Research Foundation of Ukraine (2020.01/0140); MP and AH were supported by the Estonian Research Council (PRG874, PRG609), and the European Regional Development Fund (Centre of Excellence EcolChange); Data collection of HCP was funded by FORMAS (Swedish Research Council for Environment, Agricultural Science and Spatial Planning) and The Swedish Institute; JR was supported by the Czech Science Foundation (grant No. 20‐09895S) and the long‐term developmental project of the Czech Academy of Sciences (RVO 67985939); ATRA was funded by the Grant of Excellence Departments, MIUR‐Italy (ARTICOLO 1, COMMI 314 – 337 LEGGE 232/2016); JMA was supported by Carl Tryggers stiftelse för vetenskaplig forskning and Qatar Petroleum; AAli was supported by the Jiangsu Science and Technology Special Project (Grant No. BX2019084), and Metasequoia Faculty Research Startup Funding at Nanjing Forestry University (Grant No. 163010230), and he is currently supported by Hebei University through Faculty Research Startup Funding Program; ZB was supported by the NKFI K 124796 grant; The GLORIA‐ Aragón project of JLBA was funded by the Dirección General de Cambio Climático del Gobierno de Aragón (Spain); MCs and LDem were supported by DG Environment through the European Forum on Nature Conservation and Pastoralism and Barbara Knowles Fund, in collaboration with Pogány‐havas Association, Romania; JDa was partially supported by long‐term research development project no. RVO 67985939 of the Czech Academy of Sciences; BD and OV were supported by the NKFI KH 126476, NKFI KH 130338, NKFI FK 124404 and NKFI FK 135329 grants; BD, OV and AKe were supported by the Bolyai János Scholarship of the Hungarian Academy of Sciences; BE was funded by the Environmental Department of the Tyrolean Federal State Government, the MAB Programme of the Austrian Academy of Science, the Mountain Agriculture Research Unit and the Alpine Research Centre Obergurgl of Innsbruck University. The GLORIA projects of BE were funded by the EU project no. EVK2‐CT‐2000‐00056, the Earth System Sciences Program of the Austrian Academy of Sciences (project MEDIALPS), the Amt für Naturparke, Autonome Provinz Bozen‐Südtirol, the Südtiroler Wissenschaftsfonds and the Tiroler Wissenschaftsfonds; RGG was supported by the Spanish Ministry of Research to sample GLORIA sites in central Spain (CGL 2008‐00901/BOS) and present works by the Autonomous Region of Madrid (REMEDINAL TE‐CM, S2018/EMT‐4338); MJ was supporteLatviaed by Latvia Grant No. 194051; NP and SŠ were partly supported by the Slovenian Research Agency, core fundings P1‐0403 and J7‐1822.

Canopy height greatly affects the biomass stock, carbon dynamics, and maintenance of biodiversity in forests. Previous research reported that the maximum forest canopy height (Hmax) at global and regional scales could be explained by variations in water or energy availability, that is, the water- or energy-related hypothesis. However, fundamental gaps remain in our understanding of how different drivers (i.e., water and energy) contribute to the Hmax at the local scale. In this study, we selected eight dynamic forest plots (20–30 ha) across a latitudinal gradient (from 21.6° N to 48.1° N) in China and measured the canopy structure using airborne light detection and ranging (LiDAR) data. Based on the LiDAR point cloud data, we extracted the maximum tree height (Hmax) in a 20 × 20 m quadrat as a proxy for canopy height, and the topographic wetness index (TWI) and digital terrain model-derived insolation (DTMI) were calculated as proxies for water and energy conditions. We used a linear mixed model and spatial simultaneous autoregressive error model to quantify how TWI and DTMI contributed to variations in Hmax at the local scale. We found that the positive effect of TWI was stronger in subtropical and tropical forests, highlighting that water was the main factor that drives the canopy height pattern in these regions. In contrast, although the effects of DTMI can be both positive and negative, its relative contribution was higher in temperate forest plots than in other forest types, supporting the idea that energy input is more critical for Hmax in temperate forests. Overall, our study revealed the directional change from energy to water limitation from temperate to subtropical and tropical forests. Our findings can offer important insights into forest management, especially under global climate change in the Anthropocene.

International audience; Tree diversity has been shown to promote a broad range of ecosystem functions in forests. However, how important these effects are in driving ecosystem multifunctionality in natural forests, relative to other drivers, such as below‐ground biodiversity (e.g. soil microbial diversity), community‐level functional traits and environmental conditions, remains poorly understood. Here, we hypothesize that tree species or phylogenetic diversity (PD), stand structure, functional traits and soil microbial diversity jointly regulate temperate forest multifunctionality (FM) along a local‐scale environmental gradient.Using repeated census data from a 25‐ha old‐growth temperate forest, we first quantified eight ecosystem functions and properties related to above‐ and below‐ground nutrient cycling. We then used these to estimate ecosystem multifunctionality using both averaging and multiple threshold (50%, 75% and 95%) approaches. Finally, we used structural equation models to explore how different facets of tree (tree species, functional and PD) and soil (bacteria, fungi and nematode diversity) biodiversity influence ecosystem multifunctionality, as well as how these relationships are modulated by stand structural attributes and environmental conditions (topography and soil nutrients). Forest multifunctionality was positively related to stand structural complexity but negatively related to acquisitive traits (i.e. community‐weighted mean of specific leaf area). Plant PD had no significant direct effect on FM, but it had a significant indirect effect via increased stand structural complexity. The effect of soil microbial diversity on FM increased with increasing threshold levels of FM and outperformed tree diversity and environmental conditions at the highest threshold level (i.e. 95%). Forests on steep slopes had lower levels of ecosystem multifunctionality due to decreased stand structural complexity. Soil nutrients were responsible for regulating FM via plant trait composition and, to a lesser extent, via tree diversity, stand structure and soil microbial diversity.Synthesis . Plant PD, stand structure and soil microbial diversity jointly regulated FM, and these effects were influenced by local‐scale changes in environmental conditions. Soil microbial diversity was a key driver of highly multifunctional forests, whereas conservation of complex stand structure and conservative trait dominance could enhance mean values of multiple functions.

Although large trees explain the most variation in biomass and structure of forests, but little is known about whether and how large trees can explain variation of forest dynamics (forest productivity and biomass loss). To shed light on the effects of large trees on forest dynamics we analyzed repeated forest inventory data on 74,300 adult trees spread across 700 quadrats of temperate mixed forest in northeastern China. Here, using piecewise structural equation modeling we examined how top 1% high-biomass, 99% remaining-biomass trees, functional trait diversity (FD) and composition (CWM), climate and soil conditions influence forest dynamics in temperate old growth forests. We found that top 1% high-biomass and top 1% large-diameter trees highly increased forset dynamics and biomass stock rather their relevant 99% remaining trees, FD and CWM supporting big-size trees hypothesis. In addition, climate is more important determinant of forest productivity rates than soil nutrients. Moreover, forest productivity and biomass stock declined with CWM traits. Hence, we highlight top 1% trees overruled the effect of 99% remaining trees, FD and CWM on forest dynamics and biomass stock. We argue that including site variation and the big trees to the integrative ecological modeling help better understanding of ecological mechanisms and drivers of forest dynamics.

Plant sexual systems appear to play an important role in community assembly: Dioecious species are found to tend to have a higher propensity to colonize communities in early successional stages. Here, we test two demographic hypotheses to explain this pattern in temperate forests. First, we test demographic differences between hermaphrodite and dioecious species in stressful younger successional stages: Previous theory predicts that hermaphrodite seed production is more harmed in stressful environments than that of dioecious populations leading to an advantage for females of dioecious species. Second, in primary forest, we hypothesized that dioecious species would show demographic advantage over monomorphic ones. We used data from two temperate forest plots in Northeast China surveyed over 10 years to compare the rates of growth and mortality of tree species with contrasting breeding systems in both secondary and primary forests. We assessed the effect of breeding system on the growth‐mortality trade‐off, while controlling for other traits usually considered as correlates of growth and mortality rates. We show that in the secondary forest, dioecious species showed weak advantage in demographic rates compared with monomorphic species; dioecious species showed considerably both lower relative growth and mortality rates compared to the hermaphrodites in the primary forest over 10 years, consistent with a priori predictions. Hermaphrodites showed strong growth‐mortality trade‐offs across forest stages, even when possibly confounding factors had been accounted for. These results suggest that sexual system influences community succession and assembly by acting on the rates of growth and mortality, and the trade‐off between them. As vegetation develops, the demographic differences between breeding systems are much larger. Our results demonstrate the association between breeding system, succession, and community assembly and that this relationship is succession‐stage dependent. Our findings support the suggestion that the demographic advantage of dioecious species facilitates the coexistence of sexual systems in primary forest.

Forest ecosystem productivity is crucial for regulating global climate change and carbon budget. Previous studies have shown that community diversity (such as species, functional and structural diversity) promotes forest productivity. However, species, functional, and structural diversity are often affected by canopy gap disturbance. In the context of global change, increases in canopy gap disturbance can augment the uncertainty in these relationships. Here, we used forest inventory, functional traits, and environmental data from a 20-ha subtropical forest plot located in the Zhejiang Province of China. We applied spatial synchronous autoregressive error modeling and structural equation models (SEMs) to analyze the effects of environmental factors (topography and soil properties), canopy gap disturbance (with and without the inclusion of the effect), community diversity variables on productivity. Results showed that stand structural diversity explained the largest amount of variation in productivity, followed by initial above-ground biomass and stand density. For canopy disturbed quadrats, stand structural explained the largest amount of variation in productivity, followed by stand density and species richness. For canopy non-disturbed quadrats, initial above-ground biomass explained the largest amount of variation in productivity, followed by stand density and stand structural diversity. Canopy gap disturbance enhanced the effect of stand structural diversity on productivity and reduced the effect of initial above-ground biomass. Our results provide strong evidence for the effect of canopy gap disturbance on the relationship between biodiversity and productivity in a subtropical forest. We argue that canopy gap disturbance reduces competition and promotes complementary utilization of resources, thus enhancing the impact of niche complementarity on productivity through species richness, stand density, and tree DBH diversity in forests.

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.

Questions Mast seeding, synchronous highly variable seed production among years, occurs in many perennial plant species and across diverse plant communities. Two predominant ultimate hypotheses for mast seeding are pollination efficiency and predator satiation, with weather conditions as a proximate cause. Little consensus has been achieved regarding the relative impacts of ultimate selection and proximate weather on variation in seed production. Moreover, mast-seeding studies often focus on a single species or phylogenetically closely related species, while studies on plant communities consisting of tree species with a diversity of reproductive strategies are less common. Location The 25-ha Changbaishan temperate forest dynamic plot, Northeast China. Methods We used eight-years of seed rain data to characterize the patterns and evaluate underlying selective drivers of mast seeding. We employed generalized linear mixed-effects models to analyze the relationships between interannual variability of mast seeding and weather conditions. Results All 20 species in this forest community exhibited high variability in annual seed production, but the magnitude of seed production among species was generally asynchronous across years. Wind-pollinated species had higher interannual variation of seed production than animal-pollinated species, while species dispersed by seed predators and abiotic modes (e.g., wind and gravity) showed little variation. Species responded individually to weather conditions for both temperature and precipitation, and spring phenology at the same year as seed production had a larger effect among species than both the weather conditions of the same summer and season-long lags. Conclusions Our findings suggest that pollination efficiency hypothesis hada much stronger effect than predation satiation hypothesis on mast seeding, and weather conditions showed the proximate role of weather drivers in producing the community-wide mast seeding pattern. We emphasize the necessity to simultaneously assess drivers of mast seeding for multiple species within a plant community. This article is protected by copyright. All rights reserved.

Ectomycorrhizal (ECM) fungi are closely related to vegetation compositions, edaphic properties, and site-specific processes. However, the coevolutionary mechanisms underlying the spatial distributions in floristic and ECM fungal composition in the context of biotic adaptations and abiotic variances remain unclear. We combine a total of 25 ECM fungus-associated environmental variables to impose three types of composite scores and then quantify the environmental gradients of geographical site, soil chemical property and vegetation functional trait across 122 grids of 20 m × 20 m in a 25-hm2 forest plot. Significant dissimilarities in vegetational and ECM fungal abundance and composition existed along the above environmental gradients. Specifically, a contrasting floristic distribution (e.g., Betula platyphylla vs. Tilia mandshurica) existed between the northeastern and southwestern areas and was closely related to the nutrient and moisture gradients (with high levels in the west and low levels in the east). Furthermore, the ECM fungal communities were more abundant in the nutrient-poor and low-moisture environments than in the nutrient-rich and high-moisture environments, and the mixed-forest in the middle-gradient sites between the northeastern and southwestern areas harbored the highest ECM fungal diversity. These findings suggest that predictable within-site vegetation succession is closely related to ECM-associated determinants and the natural spatial heterogeneity of edaphic properties at a local scale.

International audience; Forests play a key role in regulating the global carbon cycle, and yet the abiotic and biotic conditions that drive the demographic processes that underpin forest carbon dynamics remain poorly understood in natural ecosystems. To address this knowledge gap, we used repeat forest inventory data from 92,285 trees across four large permanent plots (4-25 ha in size) in temperate mixed forests in northeast China to ask the following questions: (1) How do soil conditions and stand age drive biomass demographic processes? (2) How do vegetation quality (i.e., functional trait diversity and composition) and quantity (i.e., initial biomass stocks) influence biomass demographic processes independently from soil conditions and stand age? (3) What is the relative contribution of growth, recruitment, and mortality to net biomass change? Using structural equation modeling, we showed that all three demographic processes were jointly constrained by multiple abiotic and biotic factors and that mortality was the strongest determinant on net biomass change over time. Growth and mortality, as well as functional trait diversity and the community-weighted mean of specific leaf area (CWM SLA), declined with stand age. By contrast, high soil phosphorous concentrations were associated with greater functional diversity and faster dynamics (i.e., high growth and mortality rates), but associated with lower CWM SLA and initial biomass stock. More functionally diverse communities also had higher recruitment rates, but did not exhibit faster growth and mortality. Instead, initial biomass stocks and CWM SLA were stronger predictors of biomass growth and mortality, respectively. By integrating the full spectrum of abiotic and biotic drivers of forest biomass dynamics, our study provides critical system-level insights needed to predict the possible consequences of regional changes in forest diversity, composition, structure and function in the context of global change.

International audience; AbstractKey MessageDeterministic processes drive functional and phylogenetic temporal changes of woody species in temperate forest, depending on successional stage and tree size classes. The dominant influential factors changed from abiotic (especially topographic factors) to biotic (e.g., basal area), with both increasing successional stage and tree size, suggesting that deterministic processes shifted from habitat filtering to biotic interactions.ContextDisentangling how deterministic and stochastic processes govern the temporal change of community composition is critical for understanding community assembly and predicting long-term ecosystem change. However, rare studies have examined the distinctive imprint of these two processes on functional and phylogenetic temporal changes of woody species in temperate forests.AimsThe objectives of this study are to detect (1) the relative importance of deterministic vs. stochastic process in driving the functional and phylogenetic temporal changes at different successional stages and tree size classes in temperate forests and (2) the relative influence of abiotic vs. biotic factors on temporal change.MethodsWe analyzed 10 years of detailed species composition, phylogenetic information, and 14 functional traits from 66 woody species in young-growth (5-ha) and old-growth (25-ha) forest plots in Northeast China. We devised a null model approach to determine the relative importance of deterministic and stochastic processes in driving functional and phylogenetic temporal changes. Then, we investigated the influence of abiotic (soil and topography) and biotic (basal area, stem density, and species richness) factors on temporal change using boosted regression tree (BRT) models. All analyses were conducted for different tree sizes (all trees, small trees [diameter at breast height (DBH) < 10 cm], and large trees [DBH ≥ 10 cm]).ResultsIn young-growth forest, functional and phylogenetic temporal changes were basically higher than expected for all trees, small trees, and large trees. Conversely, in the old-growth forest, functional and phylogenetic temporal changes were lower than expected for all trees, but opposite patterns were found for two size classes; i.e., change was higher than expected for large trees and lower than expected for small trees. BRT models showed that the most influential factors underlying the temporal change of all trees shifted from abiotic (e.g., topography) to biotic (e.g., basal area) factors with increasing succession stage. Regardless of successional stage, the relevant factors changed from abiotic factors for small trees to biotic factors for large trees.ConclusionDeterministic processes govern tree functional and phylogenetic temporal changes, and the underlying deterministic processes change from habitat filtering to biotic interaction over succession and with increasing tree size.

Large trees, as the versatile provider of ecosystem services, are the most susceptible to episodic disturbance regimes such as extreme climate and biotic agents across the globe, and hence, largely determine forest demographic processes. Despite the recognized fundamental ecological significance of large-diameter trees to above-ground biomass (AGB) stock, relatively little is known about their distinctive role, interplaying with abiotic and biotic drivers, in shaping forest AGB dynamics (i.e., gain and loss) at regional or larger extents. Here, we hypothesized that the AGB of few large trees drives AGB stock and dynamics (gain and loss) better than species diversity and trait composition in temperate forests along topography and soil gradients. We analyzed two repeated forest inventories data from 700 plots (including 74,300 adult trees) across 15 plots in temperate forests of northeastern China, including old-growth and disturbed forests. A steady net AGB change increase was found for 13 of the 15 plots. Despite disturbed forests were recovering from post-logging disturbance with a lower density of large-diameter trees and AGB stock than old-growth plots, disturbed forests had higher AGB gain, net AGB change and lower AGB loss than old-growth forests. The largest trees (i.e. top 1% AGB or diameter) were consistently more important for explaining AGB stock, gain and loss compared to plant diversity and trait composition. Besides, the elevation-related topographic factors were more direct determinants of forest AGB accumulation than soil nutrients, and also exerted the indirect impacts via increasing the dominance of resource-conservative trees, stand structural diversity and top 1% large trees. Overall, we highlight the key overruling effect of top 1% large trees on AGB stock and dynamics in response to local environmental factors. This study could improve our understanding of the multiple ecological mechanisms underlying the global change drivers of forest AGB stock and dynamics.

Forest ecosystems play a fundamental role in the global carbon cycle. However, how stand-level changes in tree age and structure influence biomass stock and dynamics in old-growth forests is a question that remains unclear. In this study, we quantified the aboveground biomass (AGB) standing stock, the coarse woody productivity (CWP), and the change in biomass over ten years (2004–2014) in a 25 ha unmanaged broad-leaved Korean pine mixed forest in northeastern China. In addition, we quantified how AGB stock and change (tree growth, recruitment and mortality) estimations are influenced by the variation in habitat heterogeneity, tree size structure and subplot size. Our analysis indicated that Changbai forest had AGB of 265.4 Mg ha −1 in 2004, and gained1.36 Mg ha −1 y −1 between 2004 and 2014. Despite recruitment having better performance in nutrient rich habitat, we found that there is a directional tree growth trend independent of habitat heterogeneity for available nutrients in this old growth forest. The observed increases in AGB stock (∼70%) are mainly attributed to the growth of intermediate size trees (30–70 cm DBH), indicating that this forest is still reaching its mature stage. Meanwhile, we indicated that biomass loss due to mortality reduces living biomass, not increment, may be the primary factor to affect forest biomass dynamics in this area. Also, spatial variation in forest dynamics is large for small sizes (i.e. coefficient of variation in 20 × 20 m subplots is 53.2%), and more than 90 percent of the inherent variability of these coefficients was predicted by a simple model including plot size. Our result provides a mean by which to estimate within-plot variability at a local scale before inferring any directional change in forest dynamics at a regional scale, and information about the variability of forest structure and dynamics are fundamental to design effective sampling strategies in future study.

Elemental stoichiometry of soil is a useful indicator of nutrient constraints. The aim of this study is to assess the relative contributions of local-scale determinants to the soil carbon/nitrogen ratio (C:N) and nitrogen/phosphorus ratio (N:P) in a 25 ha temperate forest plot. Leaf litter biomass at 967 soil sampling points was predicted using mapped tree distribution and leaf litter data (150 traps) for seven tree species. Boosted regression trees (BRT) were used to examine the relative contribution of local-scale variables to key elemental stoichiometry of soil. These variables included leaf litter input, neighbor species composition, soil pH and moisture, and topography. Both biotic and abiotic variables influenced soil C:N and N:P ratios, but abiotic factors had a greater influence. Soil pH was the most important predictor of soil C:N and N:P with a strong positive correlation. Topography and soil moisture explained more of the local-scale variability of soil C:N and N:P ratios than tree composition and leaf litter inputs. Our results suggest soil pH was a better predictor of local-scale spatial variability of soil elemental stoichiometry than tree composition and leaf litter inputs in temperate forests. These results improve our understanding of the mechanisms underlying the heterogeneity of soil nutrients.

Advances in forest carbon mapping have the potential to greatly reduce uncertainties in the global carbon budget and to facilitate effective emissions mitigation strategies such as REDD+ (Reducing Emissions from Deforestation and Forest Degradation). Though broad-scale mapping is based primarily on remote sensing data, the accuracy of resulting forest carbon stock estimates depends critically on the quality of field measurements and calibration procedures. The mismatch in spatial scales between field inventory plots and larger pixels of current and planned remote sensing products for forest biomass mapping is of particular concern, as it has the potential to introduce errors, especially if forest biomass shows strong local spatial variation. Here, we used 30 large (8–50 ha) globally distributed permanent forest plots to quantify the spatial variability in aboveground biomass density (AGBD in Mg ha–1) at spatial scales ranging from 5 to 250 m (0.025–6.25 ha), and to evaluate the implications of this variability for calibrating remote sensing products using simulated remote sensing footprints. We found that local spatial variability in AGBD is large for standard plot sizes, averaging 46.3% for replicate 0.1 ha subplots within a single large plot, and 16.6% for 1 ha subplots. AGBD showed weak spatial autocorrelation at distances of 20–400 m, with autocorrelation higher in sites with higher topographic variability and statistically significant in half of the sites. We further show that when field calibration plots are smaller than the remote sensing pixels, the high local spatial variability in AGBD leads to a substantial "dilution" bias in calibration parameters, a bias that cannot be removed with standard statistical methods. Our results suggest that topography should be explicitly accounted for in future sampling strategies and that much care must be taken in designing calibration schemes if remote sensing of forest carbon is to achieve its promise.