Mycorrhizal feedbacks influence global forest structure and diversity.

One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

Diversity and biogeography of plant phyllosphere bacteria are governed by latitude-dependent mechanisms.

Predicting and managing the structure and function of plant microbiomes requires quantitative understanding of community assembly and predictive models of spatial distributions at broad geographic scales. Here, we quantified the relative contribution of abiotic and biotic factors to the assembly of phyllosphere bacterial communities, and developed spatial distribution models for keystone bacterial taxa along a latitudinal gradient, by analyzing 16S rRNA gene sequences from 1453 leaf samples taken from 329 plant species in China. We demonstrated a latitudinal gradient in phyllosphere bacterial diversity and community composition, which was mostly explained by climate and host plant factors. We found that host-related factors were increasingly important in explaining bacterial assembly at higher latitudes while nonhost factors including abiotic environments, spatial proximity and plant neighbors were more important at lower latitudes. We further showed that local plant-bacteria associations were interconnected by hub bacteria taxa to form metacommunity-level networks, and the spatial distribution of these hub taxa was controlled by hosts and spatial factors with varying importance across latitudes. For the first time, we documented a latitude-dependent importance in the driving factors of phyllosphere bacteria assembly and distribution, serving as a baseline for predicting future changes in plant phyllosphere microbiomes under global change and human activities.

Meta-analysis shows forest soil CO2 effluxes are dependent on the disturbance regime and biome type.

Forest soil CO2 efflux (FCO2 ) is a crucial process in global carbon cycling; however, how FCO2 responds to disturbance regimes in different forest biomes is poorly understood. We quantified the effects of disturbance regimes on FCO2 across boreal, temperate, tropical and Mediterranean forests based on 1240 observations from 380 studies. Globally, climatic perturbations such as elevated CO2 concentration, warming and increased precipitation increase FCO2 by 13% to 25%. FCO2 is increased by forest conversion to grassland and elevated carbon input by forest management practices but reduced by decreased carbon input, fire and acid rain. Disturbance also changes soil temperature and water content, which in turn affect the direction and magnitude of disturbance influences on FCO2 . FCO2 is disturbance- and biome-type dependent and such effects should be incorporated into earth system models to improve the projection of the feedback between the terrestrial C cycle and climate change.

Comment on "Interspecific competition limits bird species' ranges in tropical mountains".

Freeman et al. (Reports, 22 July 2022, p. 416) argue that interspecific competition rather than climate is the leading driver of bird species' elevational ranges. A reanalysis of their data shows no support for the competition hypothesis, but a strong effect of climate seasonality on species ranges. Their results are artifacts arising from a suboptimal model that misses important variables.

Positive interactions of native species melt invasional meltdown over long-term plant succession.

Positive interactions have been hypothesised to influence plant community dynamics and species invasions. However, their prevalence and importance relative to negative interactions remain unclear to understand community change and invasibility. We examined pairwise biotic interactions using over 50 years of successional data to assess the prevalence of positive interactions and their effects on each focal species (either native or exotic). We found that positive interactions were widespread and the relative frequency of positive and negative interactions varied with establishment stage and between native and exotic species. Specifically, positive interactions were more frequent during early establishment and less frequent at later stages. Positive interactions involving native species were more frequent and stronger than those between exotic species, reducing the importance of invasional meltdown on succession. Our study highlights the role of positive native interactions in shielding communities from biological invasion and enhancing the potential for long-term resilience.

Warming shifts soil microbial communities and tropical tree seedling mortality.

Plant-soil feedback (PSF), regulated by both mycorrhizae and soil-borne pathogens, is a primary mechanism maintaining high tree species diversity in the tropics. But how warming actually affects PSF is not well understood. We conducted a field warming experiment to test PSF on seedling mortality of two tree species: a rhizobia-associated tree (Ormosia semicastrata, Fabaceae) suffering from host-specific soil-borne pathogens and an ectomycorrhizal fungi-associated tree (Cyclobalanopsis patelliormis, Fagaceae) with low susceptibility to soil-borne pathogens. Soil fungi from the warming versus control seedling plots were identified by molecular sequencing. Results showed that the elevated temperature lowered seedling mortality of O. semicastrata, but had no effect on C. patelliormis seedlings. This indicates that warming weakened the negative PSF on O. semicastrata, presumably due to the observed decrease of the relative abundance of plant-pathogenic fungi and increase of ectomycorrhizal fungi but did not affect the PSF on C. patelliormis. The differential warming effects on seedling mortality of species with different microbial associations affords an example showcasing how the change in soil-borne microbes in response to global warming would, in turn, through PSF, alter tropical tree species composition and diversity. This study helps shed mechanistic light on the debate of biodiversity change as driven by climate change.

The role of demographic compensation in stabilising marginal tree populations in North America.

Demographic compensation-the opposing responses of vital rates along environmental gradients-potentially delays anticipated species' range contraction under climate change, but no consensus exists on its actual contribution. We calculated population growth rate (λ) and demographic compensation across the distributional ranges of 81 North American tree species and examined their responses to simulated warming and tree competition. We found that 43% of species showed stable population size at both northern and southern edges. Demographic compensation was detected in 25 species, yet 15 of them still showed a potential retraction from southern edges, indicating that compensation alone cannot maintain range stability. Simulated climatic warming caused larger decreases in λ for most species and weakened the effectiveness of demographic compensation in stabilising ranges. These findings suggest that climate stress may surpass the limited capacity of demographic compensation and pose a threat to the viability of North American tree populations.

A mechanistic model for nitrogen-limited plant growth.

BACKGROUND AND AIMS: Nitrogen is often regarded as a limiting factor to plant growth in various ecosystems. Understanding how nitrogen drives plant growth has numerous theoretical and practical applications in agriculture and ecology. In 2004, Göran I. Ågren proposed a mechanistic model of plant growth from a biochemical perspective. However, neglecting respiration and assuming stable and balanced growth made the model unrealistic for plants growing in natural conditions. The aim of the present paper is to extend Ågren's model to overcome these limitations. METHODS: We improved Ågren's model by incorporating the respiratory process and replacing the stable and balanced growth assumption with a three-parameter power function to describe the relationship between nitrogen concentration (Nc) and biomass. The new model was evaluated based on published data from three studies on corn (Zea mays) growth. KEY RESULTS: Remarkably, the mechanistic growth model derived in this study is mathematically equivalent to the classical Richards model, which is the most widely used empirical growth model. The model agrees well with empirical plant growth data. CONCLUSIONS: Our model provides a mechanistic interpretation of how nitrogen drives plant growth. It is very robust in predicting growth curves and the relationship between Nc and relative growth rate.

The patterns of vascular plant discoveries in China.

AIM: (1) To understand geographic patterns of species discovery by examining the effect of growth form, range size, and geographic distribution on discovery probability of vascular plant species in China; (2) to find out which taxa harbor the largest number of undiscovered species and where those species locate; and (3) to find out the determinants of province-level mean discovery time and inventory completeness. LOCATION: China. METHODS: We compiled the discovery time and province-level geographic distributions of ~31,000 vascular plant species described between 1753 and 2013 from Flora of China. We used a Cox proportional hazard model to determine the biological and geographic correlates of discovery probability. Accumulation curves of species discoveries were fitted by a logistic discovery model to estimate inventory completeness of different growth forms and of different provinces. We then used linear regression to identify the determinants of mean discovery time and beta regression to identify the determinants of inventory completeness. RESULTS: We found that species with larger range size and distributed in northeastern part of China have a higher discovery probability. Coastal species were discovered earlier than inland species. Trees and shrubs of seed plants have the highest discovery probability while ferns have the lowest discovery probability. Herbs have the largest number of undiscovered species in China. Most undiscovered species will be found in southwest China, where three global biodiversity hot spots locate. Spatial patterns of mean discovery time and inventory completeness are mainly driven by the total number of species, human population density in an area, and latitude and longitude of a province. MAIN CONCLUSIONS: Socioeconomic factors primarily determine the discovery patterns of vascular plants in China. Undiscovered species are most likely to be narrow-ranged, inconspicuous endemic species such as herbs and ferns, which are prone to extinctions and locate in biodiversity hot spots in southwestern China.

Trait hierarchies are stronger than trait dissimilarities in structuring spatial co-occurrence patterns of common tree species in a subtropical forest.

The dissimilarity and hierarchy of trait values that characterize niche and fitness differences, respectively, have been increasingly applied to infer mechanisms driving community assembly and to explain species co-occurrence patterns. Here, we predict that limiting similarity should result in the spatial segregation of functionally similar species, while functionally similar species will be more likely to co-occur either due to environmental filtering or due to competitive exclusion of inferior competitors (hereafter hierarchical competition).We used a fully mapped 50-ha subtropical forest plot in southern China to explore how pairwise spatial associations between saplings and between adult trees were influenced by trait dissimilarity and hierarchy in order to gain insight into assembly mechanisms. We assessed pairwise spatial associations using two summary statistics of spatial point patterns at different spatial scales and compared the effects of trait dissimilarity and trait hierarchy of different functional traits on the interspecific spatial associations. These comparisons allow us to disentangle the effects of limiting similarity, environmental filtering, and hierarchical competition on species co-occurrence.We found that trait dissimilarity was generally negatively related to interspecific spatial associations for both saplings and adult trees across spatial scales, meaning that species with similar trait values were more likely to co-occur and thus supporting environmental filtering or hierarchical competition. We further found that trait hierarchy outweighed trait dissimilarity in structuring pairwise spatial associations, suggesting that hierarchical competition played a more important role in structuring our forest community than environmental filtering across life stages.This study employed a novel method, by offering the integration of pairwise spatial association and trait dissimilarity as well as trait hierarchy, to disentangle the relative importance of multiple assembly mechanisms in structuring co-occurrence patterns, especially the mechanisms of environmental filtering and hierarchical competition, which lead to indistinguishable co-occurrence patterns. This study also reinforced the importance of trait hierarchy rather than trait dissimilarity in driving neighborhood competition.

Arbuscular mycorrhizal trees influence the latitudinal beta-diversity gradient of tree communities in forests worldwide.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity.

Interactions between all pairs of neighboring trees in 16 forests worldwide reveal details of unique ecological processes in each forest, and provide windows into their evolutionary histories.

When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands' physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees' distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions.

Warming intensifies soil pathogen negative feedback on a temperate tree.

The soil pathogen-induced Janzen-Connell (JC) effect is considered as a primary mechanism regulating plant biodiversity worldwide. As predicted by the framework of the classic plant disease triangle, severity of plant diseases is often influenced by temperature, yet insufficient understanding of how increasing temperatures affect the JC effect contributes uncertainty in predictions about how global warming affects biodiversity. We conducted a 3-yr field warming experiment, combining open-top chambers with pesticide treatment, to test the effect of elevated temperatures on seedling mortality of a temperate tree species, Prunus padus, from a genus with known susceptibility to soil-borne pathogens. Elevated temperature significantly increased the mortality of P. padus seedlings in the immediate vicinity of parent trees, concurrent with increased relative abundance of pathogenic fungi identified to be virulent to Prunus species. Our study offers experimental evidence suggesting that global warming significantly intensifies the JC effect on a temperate tree species due to increased relative abundance of pathogenic fungi. This work advances our understanding about changes in the JC effect linked to ongoing global warming, which has important implications for predicting tree diversity in a warmer future.

Differentiation in stem and leaf traits among sympatric lianas, scandent shrubs and trees in a subalpine cold temperate forest.

The scandent shrub plant form is a variant of liana that has upright and self-supporting stems when young but later becomes a climber. We aimed to explore the associations of stem and leaf traits among sympatric lianas, scandent shrubs and trees, and the effects of growth form and leaf habit on variation in stem or leaf traits. We measured 16 functional traits related to stem xylem anatomy, leaf morphology and nutrient stoichiometry in eight liana, eight scandent shrub and 21 tree species co-occurring in a subalpine cold temperate forest at an elevation of 2600-3200 m in Southwest China. Overall, lianas, scandent shrubs and trees were ordered along a fast-slow continuum of stem and leaf functional traits, with some traits overlapping. We found a consistent pattern of lianas > scandent shrubs > trees for hydraulically weighted vessel diameter, maximum vessel diameter and theoretical hydraulic conductivity. Vessel density and sapwood density showed a pattern of lianas = scandent shrubs < trees, and lianas < scandent shrubs = trees, respectively. Lianas had significantly higher specific leaf area and lower carbon concentration than co-occurring trees, with scandent shrubs showing intermediate values that overlapped with lianas and trees. The differentiation among lianas, scandent shrubs and trees was mainly explained by variation in stem traits. Additionally, deciduous lianas were positioned at the fast end of the trait spectrum, and evergreen trees at the slow end of the spectrum. Our results showed for the first time clear differentiation in stem and leaf traits among sympatric liana, scandent shrub and tree species in a subalpine cold temperate forest. This work will contribute to understanding the mechanisms responsible for variation in ecological strategies of different growth forms of woody plants.

Plant breeding systems influence the seasonal dynamics of plant-pollinator networks in a subtropical forest.

Temporal dynamics of plant-pollinator interactions inform the mechanisms of community assembly and stability. However, most studies on the dynamics of pollination networks do not consider plant reproductive traits thus offering poor understanding of the mechanism of how networks maintain stable structure under seasonal changes in flower community. We studied seasonal dynamics of pollination networks in a subtropical monsoon forest in China with a clear rainy season (April-September) and dry season (October-March) over 2 consecutive years. We constructed dioecy-ignored networks (combining visitations to dioecious male and female plants by ignoring the difference between dioecious and hermaphroditic plants) and dioecy-considered networks (excluding those visitations that only occurred either on dioecious male or female plants) for eight sampling sessions for each season. Although flower richness and flower abundance were higher in the rainy season than in the dry season, no pronounced seasonal difference was found in network specialization, nestedness and modularity for both networks. There were only significant differences in plant community robustness and pollinator specialization between seasons for dioecy-considered networks but not for dioecy-ignored networks. Furthermore, we found the flower abundance of dioecious and hermaphrodite plants mostly showed trade-off variation between rainy and dry seasons. Our results suggest various plant reproductive traits affect the temporal dynamics of pollination networks, which should be considered for conservation of plant-pollinator interactions in forest communities.

Habitat fragmentation changes top-down and bottom-up controls of food webs.

Top-down and bottom-up controls regulate the structure and stability of ecosystems, but their relative roles in terrestrial systems have been debated. Here we studied a hydro-inundated land-bridge system in subtropical China and tested the relative importance of these two controls in determining the rodent-mediated regeneration of a locally dominant tree species. Our results showed that both controls operated in terrestrial habitats and that their relative importance switched as habitat size changed. Habitat loss initially removed predators of rodents that released rodent populations and triggered massive seed predation (top-down control), leading to reduced seedling establishment. A further reduction in habitat size led to decrease in rodent population that was supposed to increase seedling survival of the tree species, but the decline in habitat size deteriorated the abiotic environments (bottom-up control) that severely prevented seedling recruitment. As the ongoing global land use change is creating increasing number of small-sized forest fragments, our findings provide novel insights into the restoration of seriously fragmented forests.

High Temperature can Change Root System Architecture and Intensify Root Interactions of Plant Seedlings.

Climate change could alter plant aboveground and belowground resource allocation. Compared with shoots, we know much less about how roots, especially root system architecture (RSA) and their interactions, may respond to temperature changes. Such responses could have great influence on species'acquisition of resources and their competition with neighbors. We used a gel-based transparent growth system to in situ observe the responses of RSA and root interactions of three common subtropical plant species seedlings in Asia differing in growth forms (herb, shrub, and tree) under a wide growth temperature range of 18-34°C, including low and supra-optimal temperatures. Results showed that the RSA, especially root depth and root width, of the three species varied significantly in response to increased temperature although the response of their aboveground shoot traits was very similar. Increased temperature was also observed to have little impact on shoot/root resource allocation pattern. The variations in RSA responses among species could lead to both the intensity and direction change of root interactions. Under high temperature, negative root interactions could be intensified and species with larger root size and fast early root expansion had competitive advantages. In summary, our findings indicate that greater root resilience play a key role in plant adapting to high temperature. The varied intensity and direction of root interactions suggest changed temperatures could alter plant competition. Seedlings with larger root size and fast early root expansion may better adapt to warmer climates.

Distance-based methods for estimating density of nonrandomly distributed populations.

Population density is the most basic ecological parameter for understanding population dynamics and biological conservation. Distance-based methods (or plotless methods) are considered as a more efficient but less robust approach than quadrat-based counting methods in estimating plant population density. The low robustness of distance-based methods mainly arises from the oversimplistic assumption of completely spatially random (CSR) distribution of a population in the conventional distance-based methods for estimating density of non-CSR populations in natural communities. In this study we derived two methods to improve on density estimation for plant populations of non-CSR distribution. The first method modified an existing composite estimator to correct for the long-recognized bias associated with that estimator. The second method was derived from the negative binomial distribution (NBD) that directly deals with aggregation in the distribution of a species. The performance of these estimators was tested and compared against various distance-based estimators by both simulation and empirical data of three large-scale stem-mapped forests. Results showed that the NBD point-to-tree distance estimator has the best and most consistent performance across populations with vastly different spatial distributions. This estimator offers a simple, efficient and robust method for estimating density for empirical populations of plant species.

Historical records reveal the distinctive associations of human disturbance and extreme climate change with local extinction of mammals.

Accelerated anthropogenic impacts and climatic changes are widely considered to be responsible for unprecedented species extinction. However, determining their effects on extinction is challenging owing to the lack of long-term data with high spatial and temporal resolution. In this study, using historical occurrence records of 11 medium- to large-sized mammal species or groups of species in China from 905 BC to AD 2006, we quantified the distinctive associations of anthropogenic stressors (represented by cropland coverage and human population density) and climatic stressors (represented by air temperature) with the local extinction of these mammals. We found that both intensified human disturbances and extreme climate change were associated with the increased local extinction of the study mammals. In the cold phase (the premodern period of China), climate cooling was positively associated with increased local extinction, while in the warm phase (the modern period) global warming was associated with increased local extinction. Interactive effects between human disturbance and temperature change with the local extinction of elephants, rhinos, pandas, and water deer were found. Large-sized mammals, such as elephants, rhinos, and pandas, showed earlier and larger population declines than small-sized ones. The local extinction sensitivities of these mammals to the human population density and standardized temperature were estimated during 1700 to 2000. The quantitative evidence for anthropogenic and climatic associations with mammalian extinction provided insights into the driving processes of species extinction, which has important implications for biodiversity conservation under accelerating global changes.

Effects of host phylogeny, habitat and spatial proximity on host specificity and diversity of pathogenic and mycorrhizal fungi in a subtropical forest.

Soil plant-pathogenic (PF) and mycorrhizal fungi (MF) are both important in maintaining plant diversity, for example via host-specialized effects. However, empirical knowledge on the degree of host specificity and possible factors affecting the fungal assemblages is lacking. We identified PF and MF in fine roots of 519 individuals across 45 subtropical tree species in southern China in order to quantify the importance of host phylogeny (including via its effects on functional traits), habitat and space in determining fungal communities. We also compared host specificity in PF and MF at different host-phylogenetic scales. In both PF and MF, host phylogeny independently accounted for > 19% of the variation in fungal richness and composition, whereas environmental and spatial factors each explained no more than 4% of the variation. Over 77% of the variation explained by phylogeny was attributable to covariation in plant functional traits. Host specificity was phylogenetically scale-dependent, being stronger in PF than in MF at low host-phylogenetic scales (e.g. within genus) but similar at larger scales. Our study suggests that host-phylogenetic effects dominate the assembly of both PF and MF communities, resulting from phylogenetically clustered plant traits. The scale-dependent host specificity implies that PF were specialized at lower-level and MF at higher-level host taxa.