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Subtropical forests play an important role in global carbon cycle and in mitigating climate change. Understanding the relationship between multiple diversity and ecosystem function is crucial for protecting and managing forests. Here we used forest inventory data from a 6-hectare sample plot in natural evergreen deciduous broad-leaf mixed forest systems for the years 2016 and 2021. We analyzed the effects of multiple aspects of diversity and topographic factors on forest productivity using multiple causal analyses. We found that Fagaceae was the primary contributor to productivity in the forest stand. Elevation, slope, and convexity showed no significant effects on productivity. Structural complexity (stand density, large-diameter trees and tree-size variation) was significantly positively correlated with productivity. Taxonomic and functional diversity indices were weakly correlated with productivity. Specifically, forest productivity was enhanced by traits associated with greater maximum height and lower wood density. Community-weighted mean traits were the most strongest predictor of productivity relative to other variables. Within this forest stand, the mass-ration hypothesis appeared to be more influential on forest productivity compared to the complementarity and selection effects. By integrating multiple drivers of forest ecosystem functioning, our study provides critical system-level insights needed to predict the potential consequences of regional changes in forest diversity, composition, structure and function.

Stand structure affects tree efficiency for a competitive use of resources and largely determines stand productivity and carbon stocks. Consequently, research on individual size and differentiation of stand structures is critical for improving monoculture-stand productivity and carbon stock. Here, we studied the effects of four thinning intensities (CK: 0%, T1: 20%, T2: 30%, and T3: 40%) in an experimental plantation of Cryptomeria japonica var. sinensis, and assessed the individual differentiation characteristics, diameter class-frequency distribution, stand productivity, and carbon stocks over 6 years. The results showed that the Gini coefficient decreased with increasing thinning intensity and stand age. Self-thinning of the C. japonica stands occurred even after thinning, and the self-thinning rate was relatively high at the age of 10-13 years. For T2 and T3 treatments, the self-thinning did not occur in the 6th year after thinning. The mean diameter of each treatment increased with increasing stand age, and the normal distribution curve of diameter class frequency gradually shifted to the right, with small changes in the CK treatment and the larger one in treatment T3. Thinning increased the large-diameter (DBH ≥ 26 cm) timber, especially in T2 and T3 treatments. Stand volume and productivity varied with stand age, with the greatest change in stand volume observed in T3, followed by that in the CK treatment. Stand productivity at different thinning intensities generally decreased and then increased with increasing stand age. Although the carbon stock of individual trees and stand increased with time, the individual trees appeared to have an obviously increasing trend with increasing thinning intensity. The results provided important insights into the implications of designing thinning intensity and timing, and determining the tree-size class removal to meet specific management objectives.

Tree species-specific responses to drought are urgently needed for assessing the impacts of current climate change on forest ecosystems. Here, we characterized the resistance, recovery, resilience, and growth recovery periods in response to different drought events based on tree-ring width index data (>30 years) for three tree species widely distributed in northern China, among which larch (Larix principis-rupprechtii Mayr) and Mongolian pine (Pinus sylvestris L. var. mongolica Litv.) are two major species used for afforestation, and birch (Betula platyphylla Suk.) is one of the most common natural tree species. Despite no significant effects of mild drought on tree growth, severe drought events significantly reduced the growth of all species, with contrasting species-specific responses. Larch trees had the lowest resistance and resilience among the three species, and Mongolian pine trees were more resistant but had a longer recovery period than birch trees. The drought responses varied with tree size. Large Mongolian pine and birch trees were more resistant but large larch trees were much more vulnerable than small trees during severe droughts. Smaller birch trees had higher resilience to severe droughts. Our study shows species-specific differences in drought responses and suggests that drought responses are tree-size dependent and drought-intensity associated, which further provides a guidance for selecting optimal cultivated tree species and designing forest managements in this region.

Forest insect outbreaks cause large changes in ecosystem structure, composition, and function. Humans often respond to insect outbreaks by conducting salvage logging, which can amplify the immediate effects, but it is unclear whether logging will result in lasting differences in forest structure and dynamics when compared with forests affected only by insect outbreaks. We used 15 years of data from an experimental removal of Tsuga canadensis (L.) Carr. (Eastern hemlock), a foundation tree species within eastern North American forests, and contrasted the rate, magnitude, and persistence of response trajectories between girdling (emulating mortality from insect outbreak) and timber harvest treatments. Girdling and logging were equally likely to lead to large changes in forest structure and dynamics, but logging resulted in faster rates of change. Understory light increases and community composition changes were larger and more rapid in the logged plots. Tree seedling and understory vegetation abundance increased more in the girdled plots; this likely occurred because seedlings grew rapidly into the sapling- and tree-size classes after logging and quickly shaded out plants on the forest floor. Downed deadwood pools increased more after logging but standing deadwood pools increased dramatically after girdling. Understory light levels remained elevated for a longer time after girdling. Perhaps because the window of opportunity for understory species to establish was longer in the girdled plots, total species richness increased more in the girdled than logged plots. Despite the potential for greater diversity in the girdled plots, Betula lenta L. (black birch) was the most abundant tree species recruited into the sapling- and tree-size classes in both the girdled and logged plots and is poised to dominate the new forest canopy. The largest difference between the girdling and logging treatments-deadwood structure and quantity-will persist and continue to bolster aboveground carbon storage and structural and habitat diversity in the girdled plots. Human responses to insect outbreaks hasten forest reorganization and remove structural resources that may further alter forest response to ongoing climate stress and future disturbances.

A better understanding of the underlying ecological mechanisms of diversity-biomass relationships in forest layers (i.e., overstory and understory) is critical to understand the importance of vertical stratification to the functioning of forest ecosystems. However, it is not clear how multiple abiotic (i.e., climate and geography) and biological (i.e., biodiversity, functional characteristics, and stand structural complexity) factors simultaneously determine the aboveground biomass (AGB) of each individual forest stratum. We used data on 156,270 trees from 1986 plots in North China to explore the relationships among biological diversity, plant functional traits, stand structure, climate and topography on variation in AGB of each stratum. The results showed that different biological factors determined the AGB of overstory and understory, and thus indicating different underlying ecological mechanisms in temperate forests. The effects of forest biodiversity on AGB were only significant in understory stratum. In the overstory of the forest, forests with high tree-size dimension inequality and high dominant tree height had larger AGB, hence mass ratio effect and stand structure complexity were the main ecological mechanisms for high biomass. In understory, diversity and overstory attributes were the main factors affecting biomass. Tree height and AGB of the overstory reduced the AGB of the understory layer. In consequence overstory attributes and niche complementation were the main ecological mechanisms in the understory. The overstory exerted influence on the understory through resource quantity and resource heterogeneity. Our findings have important implications for carbon management, enhancement of forest functions and sustainable forest management in temperate forests.

Carbon uptake by the land is a key determinant of future climate change. Unfortunately, Dynamic Global Vegetation Models have many unknown internal parameters which leads to significant uncertainty in projections of the future land carbon sink. By contrast, observed forest inventories in both Amazonia and the USA show strikingly common tree-size distributions, pointing to a simpler modelling paradigm. The curvature of these size-distributions is related to the ratio of mortality to growth in Demographic Equilibrium Theory (DET). We extend DET to include recruitment limited by competitive exclusion from existing trees. From this, we find simultaneous maxima of tree density and biomass in terms of respectively the ratio of mortality to growth and the proportion of primary productivity allocated to reproduction, an idea we call Demographic Optimality (DO). Combining DO with the ratio of mortality to growth common to the US and Amazon forests, results in the prediction that about an eighth of productivity should be allocated to reproduction, which is broadly consistent with observations. Another prediction of the model is that seed mortality should decrease with increasing seed size, such that the advantage of having many small seeds is nullified by the higher seed mortality. Demographic Optimality is therefore consistent with the common shape of tree-size distributions seen in very different forests, and an allocation to reproduction that is independent of seed size.

Accurate information on urban forests of tree sizes, health state, community structures, and spatial distribution is still limited in African cities. Using a Google Street View (GSV)-based tree-size measuring method developed by our team, this paper aims to evaluate street trees of four African metropolitan cities using GSV data. The study compiled a large dataset with 46,016 street trees in 3454 sites in Kampala, Nairobi, Bloemfontein, and Johannesburg. The data including tree size (diameter at breast height, DBH; tree height, TH; underbranch height, UBH; canopy size), tree floristic composition (apical dominance types, broadleaf-conifer-palm leaf, flowering or not), tree health (leaf color, diebacks, dead tree, and bracket-supporting percent), streetside development (lane number, roadside shops, parking vehicle, and pedestrian density), and geolocation (latitude, longitude). These data can be spatially visualized with the help of ArcGIS, and the large dataset favors reliable maps from the street-view level. Data statistics showed that four cities were dominated by broad-leaved, apical dominance, and flowering trees, with a low level of unhealthy leaves and a tiny percentage of dead. The arbor-shrubs-herb structure vegetation dominated all four cities. Kampala had the most slender trees (DBH = 23 cm, TH = 8.4 m), while Nairobi and Johannesburg had the thickest trees (DBH = 38 cm, TH = 8.5-8.6 m). Bare land rates were lowest at 23% in Bloemfontein and highest at 33% in Nairobi. Principal analysis and Pearson correlations showed that these tree variations were closely associated with street development and local land use configuration. By comparing the urban tree data in other regions of the world, we found that the trees in African cities are generally giant but have a lower density (the trees within a 100-m street segment). Our findings emphasized that GSV data is feasible enough for urban forest monitoring in Africa, and the database is helpful for urban landscape planning and management.

Understanding the basis of hybrid vigor remains a key question in crop breeding and improvement, especially for rootstock development where F1 hybrids are extensively utilized. Full-sibling UCB-1 F1 seedling rootstocks are widely planted in commercial pistachio orchards that are generated by crossing 2 highly heterozygous outbreeding parental trees of Pistacia atlantica (female) and P. integerrima (male). This results in extensive phenotypic variability, prompting costly removal of low-yielding small trees. To identify the genetic basis of this variability, we assembled chromosome-scale genome assemblies of the parental trees of UCB-1. We genotyped 960 UCB-1 trees in an experimental orchard for which we also collected multiyear phenotypes. We genotyped an additional 1,358 rootstocks in 6 commercial pistachio orchards and collected single-year tree-size data. Genome-wide single marker association tests identified loci associated with tree size and shape, sex, and precocity. In the experimental orchard, we identified multiple trait-associated loci and a strong candidate for ZZ/ZW sex chromosomes. We found significant marker associations unique to different traits and to early vs late phenotypic measures of the same trait. We detected 2 loci strongly associated with rootstock size in commercial orchards. Pseudo-testcross classification of markers demonstrated that the trait-associated alleles for each locus were segregating in the gametes of opposite parents. These 2 loci interact epistatically to generate the bimodal distribution of tree size with undesirable small trees observed by growers. We identified candidate genes within these regions. These findings provide a foundational resource for marker development and genetic selection of vigorous pistachio UCB-1 rootstock.

Increasing evidence suggests that tree growth is sink-limited by environmental and internal controls rather than by carbon availability. However, the mechanisms underlying sink-limitations are not fully understood and thus not represented in large-scale vegetation models. We develop a simple, analytically solved, mechanistic, turgor-driven growth model (TDGM) and a phloem transport model (PTM) to explore the mechanics of phloem transport and evaluate three hypotheses. First, phloem transport must be explicitly considered to accurately predict turgor distributions and thus growth. Second, turgor-limitations can explain growth-scaling with size (metabolic scaling). Third, turgor can explain realistic growth rates and increments. We show that mechanistic, sink-limited growth schemes based on plant turgor limitations are feasible for large-scale model implementations with minimal computational demands. Our PTM predicted nearly uniform sugar concentrations along the phloem transport path regardless of phloem conductance, stem water potential gradients and the strength of sink-demands contrary to our first hypothesis, suggesting that phloem transport is not limited generally by phloem transport capacity per se but rather by carbon demand for growth and respiration. These results enabled TDGM implementation without explicit coupling to the PTM, further simplifying computation. We test the TDGM by comparing predictions of whole-tree growth rate to well-established observations (site indices) and allometric theory. Our simple TDGM predicts realistic tree heights, growth rates and metabolic scaling over decadal to centurial timescales, suggesting that tree growth is generally sink and turgor limited. Like observed trees, our TDGM captures tree-size- and resource-based deviations from the classical ¾ power-law metabolic scaling for which turgor is responsible.

Old-growth forests are valuable sources of ecological, conservation, and management information, yet these ecosystems have received little study in New England, due in large part to their regional scarcity. To increase our understanding of the structures and processes common in these rare forests, we studied the abundance of downed coarse woody debris (CWD) and snags and live-tree size-class distributions in 16 old-growth hemlock forests in western Massachusetts. Old-growth stands were compared with eight adjacent second-growth hemlock forests to gain a better understanding of the structural differences between these two classes of forests resulting from contrasting histories. In addition, we used stand-level dendroecological reconstructions to investigate the linkages between disturbance history and old-growth forest structure using an information-theoretic model selection framework. Old-growth stands exhibit a much higher degree of structural complexity than second-growth forests. In particular, old-growth stands had larger overstory trees and greater volumes of downed coarse woody debris (135.2 vs. 33.2 m3/ha) and snags (21.2 vs. 10.7 m3/ha). Second-growth stands were characterized by either skewed unimodal or reverse-J shaped diameter distributions, while old-growth forests contained bell-shaped, skewed unimodal, rotated sigmoid, and reverse J-shaped distributions. The variation in structural attributes among old-growth stands, particularly the abundance of downed CWD, was closely related to disturbance history. In particular, old-growth stands experiencing moderate levels of canopy disturbance during the last century (1930s and 1980s) had greater accumulations of CWD, highlighting the importance of gap-scale disturbances in shaping the long-term development and structural characteristics of old-growth forests. These findings are important for the development of natural disturbance-based silvicultural systems that may be used to restore important forest characteristics lacking in New England second-growth stands by integrating structural legacies of disturbance (e.g., downed CWD) and resultant tree-size distribution patterns. This silvicultural approach would emulate the often episodic nature of CWD recruitment within old-growth forests.

Stable carbon isotope ratios (δ13C) in tree rings have been widely used to study changes in intrinsic water-use efficiency (iWUE), sometimes with limited consideration of how C-isotope discrimination is affected by tree height and canopy position. Our goals were to quantify the relationships between tree size or tree microenvironment and wood δ13C for eight functionally diverse temperate tree species in northern New England and to better understand the physical and physiological mechanisms underlying these differences. We collected short increment cores in closed-canopy stands and analyzed δ13C in the most recent 5 years of growth. We also sampled saplings in both shaded and sun-exposed environments. In closed-canopy stands, we found strong tree-size effects on δ13C, with 3.7-7.2‰ of difference explained by linear regression vs height (0.11-0.28‰ m-1), which in some cases is substantially stronger than the effect reported in previous studies. However, open-grown saplings were often isotopically more similar to large codominant trees than to shade-grown saplings, indicating that light exposure contributes more to the physiological and isotopic differences between small and large trees than does height. We found that in closed-canopy forests, δ13C correlations with diameter at breast height were nonlinear but also strong, allowing a straightforward procedure to correct tree- or stand-scale δ13C-based iWUE chronologies for changing tree size. We demonstrate how to use such data to correct and interpret multi-decadal composite isotope chronologies in both shade-regenerated and open-grown tree cohorts, and we highlight the importance of understanding site history when interpreting δ13C time series.

The protection of tropical forests is one of the most urgent issues in conservation biology because of the rapid deforestation that has occurred over the last 50 years. Even in protected forests, the anthropogenic effects from newly expanding villages such as harvesting of medicinal plants, pasturing cattle and forest fires can induce environmental modifications, especially on the forest floor. We evaluated the anthropogenic effects of the daily activities of neighboring residents on natural forests in 12 plots extending from the village boundary into a natural forest in Thailand. The basal area per unit land area did not present a significant trend; however, the species diversity of woody plants decreased linearly towards the village boundary, which caused a loss of individual density because of severe declines in small saplings compared with adult trees and large saplings in proximity to the village. An analysis of tree-size categories indicates a lack of small samplings near the village boundary. The current forest appears to be well protected based on the adult tree canopy, but regeneration of the present-day forests is unlikely because of the loss of seedlings.

We investigated the relationships within forest stands between tree size and (a) stem volume growth rate and (b) risk of mortality for individual trees. Values of both x and y variables were plotted relative to the largest value in the stand. We refer to the resultant presentations as relative volume growth and relative survival graphs (VGSs). A pair of VGSs can be produced readily from an individual-tree growth model. It can also be constructed from consecutive sets of field measurements. Comparison of VGSs derived from model and measurement data provides a test of the validity of the components of the growth model. We have analyzed VGSs based on measurement data for Scots pine (Pinus sylvestris L.) in central Finland and for beech (Fagus sylvatica L.) in southern Germany. The graphs based on measurement data varied as a consequence of differences in competition, stand management, and tree species. We analyzed the relationship between VGSs and stand dynamics using a simple growth model. We found that different features of the VGSs imply characteristic tree size distributions in subsequent years. Thus, we conclude that if the VGSs generated by a model do not correspond to those based on field measurements, the model cannot be relied on to reproduce the development of tree-size distribution correctly. Relative growth and survival graphs thus provide a tool for evaluating complicated growth models.

Tropical montane cloud forests (TMCFs) harbour high levels of biodiversity and large carbon stocks. Their location at high elevations make them especially sensitive to climate change, because a warming climate is enhancing upslope species migration, but human disturbance (especially fire) may in many cases be pushing the treeline downslope. TMCFs are increasingly being affected by fire, and the long-term effects of fire are still unknown. Here, we present a 28-year chronosequence to assess the effects of fire and recovery pathways of burned TMCFs, with a detailed analysis of carbon stocks, forest structure and diversity. We assessed rates of change of carbon (C) stock pools, forest structure and tree-size distribution pathways and tested several hypotheses regarding metabolic scaling theory (MST), C recovery and biodiversity. We found four different C stock recovery pathways depending on the selected C pool and time since last fire, with a recovery of total C stocks but not of aboveground C stocks. In terms of forest structure, there was an increase in the number of small stems in the burned forests up to 5-9 years after fire because of regeneration patterns, but no differences on larger trees between burned and unburned plots in the long term. In support of MST, after fire, forest structure appears to approximate steady-state size distribution in less than 30 years. However, our results also provide new evidence that the species recovery of TMCF after fire is idiosyncratic and follows multiple pathways. While fire increased species richness, it also enhanced species dissimilarity with geographical distance. This is the first study to report a long-term chronosequence of recovery pathways to fire suggesting faster recovery rates than previously reported, but at the expense of biodiversity and aboveground C stocks.

Scale-free foraging patterns are widespread among animals. These may be the outcome of an optimal searching strategy to find scarce, randomly distributed resources, but a less explored alternative is that this behaviour may result from the interaction of foraging animals with a particular distribution of resources. We introduce a simple foraging model where individual primates follow mental maps and choose their displacements according to a maximum efficiency criterion, in a spatially disordered environment containing many trees with a heterogeneous size distribution. We show that a particular tree-size frequency distribution induces non-Gaussian movement patterns with multiple spatial scales (Lévy walks). These results are consistent with field observations of tree-size variation and spider monkey (Ateles geoffroyi) foraging patterns. We discuss the consequences that our results may have for the patterns of seed dispersal by foraging primates.

At a newly occupied pond, beavers preferentially felled aspen smaller than 7.5 cm in diameter and selected against larger size classes. After one year of cutting, 10% of the aspen had been cut and 14% of the living aspen exhibited the juvenile growth form. A phenolic compound which may act as a deterrent to beavers was found in low concentrations in aspen bark, and there was no significant regression of relative concentration of this compound on tree diameter. At a pond which had been intermittently occupied by beavers for over 20 years, beavers selected against aspen smaller than 4.5 cm in diameter, and selected in favor of aspen larger than 19.5 cm in diameter. After more than 28 years of cutting at this site, 51% of the aspen had been cut and 49% of the living aspen were juvenileform. The phenolic compound was found in significantly higher concentrations in aspen bark than at the newly occupied site, and there was a significant negative regression of relative concentration on tree diameter. The results of this study show that responses to browsing by trees place constraints on the predictive value of standard energy-based optimal foraging models, and limitations on the use of such models. Future models should attempt to account for inducible responses of plants to damage and increases in concentrations of secondary metabolites through time.

Tree-size distribution is one of the most investigated subjects in plant population biology. The forestry literature reports that tree-size distribution trajectories vary across different stands and/or species, whereas the metabolic scaling theory suggests that the tree number scales universally as -2 power of diameter. Here, we propose a simple functional scaling model in which these two opposing results are reconciled. Basic principles related to crown shape, energy optimization and the finite-size scaling approach were used to define a set of relationships based on a single parameter that allows us to predict the slope of the tree-size distributions in a steady-state condition. We tested the model predictions on four temperate mountain forests. Plots (4 ha each, fully mapped) were selected with different degrees of human disturbance (semi-natural stands versus formerly managed). Results showed that the size distribution range successfully fitted by the model is related to the degree of forest disturbance: in semi-natural forests the range is wide, whereas in formerly managed forests, the agreement with the model is confined to a very restricted range. We argue that simple allometric relationships, at an individual level, shape the structure of the whole forest community.

Variations in crown forms promote canopy space-use and productivity in mixed-species forests. However, we have a limited understanding on how this response is mediated by changes in within-tree biomass allocation. Here, we explored the role of changes in tree allometry, biomass allocation and architecture in shaping diversity-productivity relationships (DPRs) in the oldest tropical tree diversity experiment. We conducted whole-tree destructive biomass measurements and terrestrial laser scanning. Spatially explicit models were built at the tree level to investigate the effects of tree size and local neighbourhood conditions. Results were then upscaled to the stand level, and mixture effects were explored using a bootstrapping procedure. Biomass allocation and architecture substantially changed in mixtures, which resulted from both tree-size effects and neighbourhood-mediated plasticity. Shifts in biomass allocation among branch orders explained substantial shares of the observed overyielding. By contrast, root-to-shoot ratios, as well as the allometric relationships between tree basal area and aboveground biomass, were little affected by the local neighbourhood. Our results suggest that generic allometric equations can be used to estimate forest aboveground biomass overyielding from diameter inventory data. Overall, we demonstrate that shifts in tree biomass allocation are mediated by the local neighbourhood and promote DPRs in tropical forests.

We review the need for scaling effects of ozone (O3) from juvenile to mature forest trees, identify the knowledge presently available, and discuss limitations in scaling efforts. Recent findings on O3/soil nutrient and O3/CO2 interactions from controlled experiments suggest consistent scaling patterns for physiological responses of individual leaves to whole-plant growth, carbon allocation, and water use efficiency of juvenile trees. These findings on juvenile trees are used to develop hypotheses that are relevant to scaling O3 effects to mature trees, and these hypotheses are examined with respect to existing research on differences in response to O3 between juvenile and mature trees. Scaling patterns of leaf-level physiological response to O3 have not been consistent in previous comparisons between juvenile and mature trees. We review and synthesize current understanding of factors that may cause such inconsistent scaling patterns, including tree-size related changes in environment, stomatal conductance, O3 uptake and exposure. carbon allocation to defense, repair, and compensation mechanisms, and leaf production phenology. These factors should be considered in efforts to scale O3 responses during tree ontogeny. Free-air O3 fumigation experiments of forest canopies allow direct assessments of O3 impacts on physiological processes of mature trees, and provide the opportunity to test current hypotheses about ontogenetic variation in O3 sensitivity by comparing O3 responses across tree-internal scales and ontogeny.