The structure of maturity: immature trees may drive the productivity of mature forests

Relating forest productivity to local variations in forest structure has been a long-standing challenge. Previous studies often focused on the connection between forest structure and stand-level photosynthesis (GPP). However, biomass production (NPP) and net ecosystem exchange (NEE) are also subject to respiration and other carbon losses, which vary with local conditions and life history traits. Here, we use a simulation approach to study how these losses impact forest productivity and reveal themselves in forest structure. We fit the process-based forest model Formind to a 25ha inventory of an old-growth temperate forest in China and classify trees as "mature" (full-grown) or "immature" based on their intrinsic carbon use efficiency. Our results reveal a strong negative connection between the stand-level carbon use efficiency and the prevalence of mature trees: GPP increases with the total basal area, whereas NPP and NEE are driven by the basal area of immature trees. Accordingly, the basal area entropy - a structural proxy for the prevalence of immature trees - correlated well with NPP and NEE and had higher predictive power than other structural characteristics such as Shannon diversity and height standard deviation. Our results were robust across spatial scales (0.04-1ha) and yield promising hypotheses field studies and new theoretical work

Multiple metrics of diversity have different effects on temperate forest functioning over succession.

Manuscript. Published version available in Oecologia, Dec. 2016, vol. 182, issue 4, pp 1175–1185Biodiversity can be measured by taxonomic, phylogenetic, and functional diversity. How ecosystem functioning depends on these measures of diversity can vary from site to site and depends on successional stage. Here, we measured taxonomic, phylogenetic, and functional diversity, and examined their relationship with biomass in two successional stages of the broad-leaved Korean pine forest in northeastern China. Functional diversity was calculated from six plant traits, and aboveground biomass (AGB) and coarse woody productivity (CWP) were estimated using data from three forest censuses (10 years) in two large fully mapped forest plots (25 and 5 ha). 11 of the 12 regressions between biomass variables (AGB and CWP) and indices of diversity showed significant positive relationships, especially those with phylogenetic diversity. The mean tree diversity-biomass regressions increased from 0.11 in secondary forest to 0.31 in old-growth forest, implying a stronger biodiversity effect in more mature forest. Multi-model selection results showed that models including species richness, phylogenetic diversity, and single functional traits explained more variation in forest biomass than other candidate models. The models with a single functional trait, i.e., leaf area in secondary forest and wood density in mature forest, provided better explanations for forest biomass than models that combined all six functional traits. This finding may reflect different strategies in growth and resource acquisition in secondary and old-growth forests

Local-Scale Drivers of Tree Survival in a Temperate Forest

Tree survival plays a central role in forest ecosystems. Although many factors such as tree size, abiotic and biotic neighborhoods have been proposed as being important in explaining patterns of tree survival, their contributions are still subject to debate. We used generalized linear mixed models to examine the relative importance of tree size, local abiotic conditions and the density and identity of neighbors on tree survival in an old-growth temperate forest in northeastern China at three levels (community, guild and species). Tree size and both abiotic and biotic neighborhood variables influenced tree survival under current forest conditions, but their relative importance varied dramatically within and among the community, guild and species levels. Of the variables tested, tree size was typically the most important predictor of tree survival, followed by biotic and then abiotic variables. The effect of tree size on survival varied from strongly positive for small trees (1–20 cm dbh) and medium trees (20–40 cm dbh), to slightly negative for large trees (>40 cm dbh). Among the biotic factors, we found strong evidence for negative density and frequency dependence in this temperate forest, as indicated by negative effects of both total basal area of neighbors and the frequency of conspecific neighbors. Among the abiotic factors tested, soil nutrients tended to be more important in affecting tree survival than topographic variables. Abiotic factors generally influenced survival for species with relatively high abundance, for individuals in smaller size classes and for shade-tolerant species. Our study demonstrates that the relative importance of variables driving patterns of tree survival differs greatly among size classes, species guilds and abundance classes in temperate forest, which can further understanding of forest dynamics and offer important insights into forest management

Rhizosphere fungal community assembly varied across functional guilds in a temperate forest

Background Rhizosphere fungi play an important role in plant community dynamics and biogeochemical cycling. While the drivers of fungal community assembly have been studied in varied ecosystems, it is still unclear how these processes function for rhizosphere soil fungi in temperate forests. Furthermore, it is unknown whether the relative contributions of important determinants remain consistent or vary across fungal ecological guilds. This study used high-throughput next-generation sequencing to characterize the fungal communities of 247 rhizosphere soil samples from 19 tree species in a temperate forest within Northeast China. We aimed to investigate how three important determinants in temperate forests (host tree species, neighbouring plant communities, and edaphic properties) influence the community assembly of fungal functional guilds in the rhizosphere soil of trees. Results We found that host tree species contributed more to plant pathogens’ community composition than ectomycorrhizal fungi, and plant pathogens consistently showed higher host specialization than ectomycorrhizal fungi. Saprotrophs also showed high host specialization, which was mediated by the tree species’ effect on rhizosphere soil pH. Although neighboring plant communities contributed remarkably to richness of all fungal guilds, this effect on fungal composition varied across functional guilds, with stronger effect for biotrophic guilds (plant pathogens and ectomycorrhizal fungi) than for non-biotrophic guild (saprotrophs). Neighboring plant communities shaped the ectomycorrhizal community composition strongly in all samples regardless of host trees’ mycorrhizal type, whereas edaphic properties were the most important drivers for this guild in samples from only ectomycorrhizal-associated trees. Edaphic properties played an important role in shaping ectomycorrhizal and saprotrophic fungal compositions, indicating the importance of edaphic properties on the fungal functional guilds associated with the absorption and decomposition of nutrients. Conclusions These results demonstrated that rhizosphere soil fungal community assembly determinants varied across fungal guilds, reflecting their different ecological functions in temperate forest ecosystems.This work was supported by the National Key Research and Development Program of China (2022YFF1300501), the Strategic Priority Research Program of the Chinese Academy of Sciences (Grant XDB31030000), the Key Research Program of Frontier Sciences, Chinese Academy of Sciences (Grant ZDBS-LY-DQC019)

El Niño–Southern Oscillation affects the species-level temporal variation in seed and leaf fall in a mixed temperate forest

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.This work was financially supported by the National Natural Science Foundation of China (32171581, 32001121), Natural Science Foundation of Liaoning Province of China (2021-MS-028), Strategic Priority Research Program of the Chinese Academy of Sciences (XDB 31030000), the Key Research Program of Frontier Sciences, Chinese Academy of Sciences (ZDBS-LY-DQC019), and K. C. Wong Education Foundation.Peer reviewe

allodb: An R package for biomass estimation at globally distributed extratropical forest plots

Allometric equations for calculation of tree above-ground biomass (AGB) form the basis for estimates of forest carbon storage and exchange with the atmosphere. While standard models exist to calculate forest biomass across the tropics, we lack a standardized tool for computing AGB across boreal and temperate regions that comprise the global extratropics. Here we present an integrated R package, allodb, containing systematically selected published allometric equations and proposed functions to compute AGB. The data component of the package is based on 701 woody species identified at 24 large Forest Global Earth Observatory (ForestGEO) forest dynamics plots representing a wide diversity of extratropical forests. A total of 570 parsed allometric equations to estimate individual tree biomass were retrieved, checked and combined using a weighting function designed to ensure optimal equation selection over the full tree size range with smooth transitions across equations. The equation dataset can be customized with built-in functions that subset the original dataset and add new equations. Although equations were curated based on a limited set of forest communities and number of species, this resource is appropriate for large portions of the global extratropics and can easily be expanded to cover novel forest types

The role of functional uniqueness and spatial aggregation in explaining rarity in trees

Aim: Determining the drivers of species rarity is fundamental for understanding and conserving biodiversity. Rarity of a given species within its community may arise due to exclusion by other ecologically similar species. Conversely, rare species may occupy habitats that are rare in the landscape or they may be ill-suited to all available habitats. The first mechanism would lead to common and rare species occupying similar ecological space defined by functional traits. The second mechanism would result in common and rare species occupying dissimilar ecological space and spatial aggregation of rare species, either because they are specialists in rare habitats or because rare species tend to be dispersal limited. Here, we quantified the contribution of locally rare species to community functional richness and the spatial aggregation of species across tree communities world-wide to address these hypotheses. Location: Asia and the Americas. Time period: 2002 to 2012 (period that considers the censuses for the plots used). Major taxa studied: Angiosperm and Gymnosperm trees. Methods: We compiled a dataset of functional traits from all the species present in eight tree plots around the world to evaluate the contribution of locally rare species to tree community functional richness using multi- and univariate approaches. We also quantified the spatial aggregation of individuals within species at several spatial scales as it relates to abundance. Results: Locally rare tree species in temperate and tropical forests tended to be functionally unique and are consistently spatially clustered. Furthermore, there is no evidence that this pattern is driven by pioneer species being locally rare. Main conclusions: This evidence shows that locally rare tree species disproportionately contribute to community functional richness, and we can therefore reject the hypothesis that locally rare species are suppressed by ecologically similar, but numerically dominant, species. Rather, locally rare species are likely to be specialists on spatially rare habitats or they may be ill-suited to the locally available environments

Direct and indirect effects of climate on richness drive the latitudinal diversity gradient in forest trees

Data accessibility statement: Full census data are available upon reasonable request from the ForestGEO data portal, http://ctfs.si.edu/datarequest/ We thank Margie Mayfield, three anonymous reviewers and Jacob Weiner for constructive comments on the manuscript. This study was financially supported by the National Key R&D Program of China (2017YFC0506100), the National Natural Science Foundation of China (31622014 and 31570426), and the Fundamental Research Funds for the Central Universities (17lgzd24) to CC. XW was supported by the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB3103). DS was supported by the Czech Science Foundation (grant no. 16-26369S). Yves Rosseel provided us valuable suggestions on using the lavaan package conducting SEM analyses. Funding and citation information for each forest plot is available in the Supplementary Information Text 1.Peer reviewedPostprin

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

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient

Among the local processes that determine species diversity in ecological communities, fluctuation‐dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness