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Lowland northern white-cedar (Thuja occidentalis) forests are increasingly exposed to extreme droughts and floods that cause tree mortality. However, it is not clear the extent to which these events may differentially affect regeneration of cedar and its increasingly common associate, balsam fir (Abies balsamea). To test this, we measured how seedlings of cedar and fir were able to avoid, resist, and recover from experimental drought and flood treatments of different lengths (8-66 days). Overall, we found that cedar exhibited a strategy of stress resistance and growth recovery (resilience) from moderate drought and flood stress. Fir, on the other hand, appears to be adapted to avoid drought and flood stress and exhibited overall lower growth resilience. In drought treatments, we found evidence of different stomatal behaviors. Cedar used available water quickly and therefore experienced more drought stress than fir but cedar was able to survive at water potentials > 3 MPa below key hydraulic thresholds. On the other hand, fir employed a more conservative water use strategy and therefore avoided extremely low water potential. In response to flood treatments, cedar survival was higher and only reached 50% if exposed to 23.1 days of flooding in contrast to only 7.4 days to reach 50% mortality for fir. In both droughts and floods, many stressed cedar were able to maintain partially brown canopies and often survived the stress, albeit with reduced growth, suggesting a strategy of resistance and resilience. In contrast, fir that experienced drought or flood stress had a threshold-type responses and they either had full live canopies with little effect on growth or they died suggesting reliance on a strategy of drought avoidance. Combined with increasingly variable precipitation regimes, seasonal flooding, and complex microtopography that can provide safe sites in these forests, these results inform conservation and management of lowland cedar stands.

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Individual modelling is a foundational approach to study the natural forest growth and in this paper, we develop a serial distance-depended individual tree model for some species in natural forest which would provide prediction and characteristics for natural species. The data used to develop individual model for natural mixed forests were collected from 712 remeasured 10-year periodic permanent sample plots of in Baihe Forest Bureau of Changbai Mountains, northeast China. Based on analyzing relationship between diameter increment of individual trees with tree size, competitive status, and site condition and finding out the major independent variables, the growth models for individual trees of 15 species in the natural mixed forests, that have good predicting precision, and easily applicable, were developed using stepwise regression method. The individual growth model developed in this study can reflect the tree increment of 15 species and be generally well suited for simulating tree and stand growth for natural mixed forests in Changbai Mountains. The research results for individual trees growth model of each species showed that main variable to influence on diameter increment of individual trees for natural mixed forests were tree size (D) and then competition index. The site condition was not related with diameter increment. The natural logarithm of DBH (lnD) and square diameter (D2) were included in the predicting models of diameter increment for all 15 species. The diameter increment was directly proportional to tree diameter for each species. For the competitive indexes in growth model, the relative diameter (RD), canopy closure (P), and the ratio of diameter of subject tree with maximum diameter (DDM) were related to diameter increment and the stand density measures were not significantly influenced on diameter increment. As canopy closure increase, tree increment decreases. The site conditions were performance less of factors in increment predictions in the model.

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BACKGROUND AND AIMS: Dioecious plant species, i.e., those in which male and female functions are housed in different individuals, are particularly vulnerable to global environmental changes. For long-lived plant species, such as trees, long-term studies are imperative to understand how growth patterns and their sensitivity to climate variability differentially affect the sexes. METHODS: Here, we explore long-term intersexual differences in wood traits, namely radial growth rates, water use efficiency quantified as stable carbon isotope abundance of wood cellulose, and their climate sensitivity in Ilex aquifolium trees growing in a natural population in NW Spain. KEY RESULTS: We found that sex differences in secondary growth rates were variable over time, with males outperforming females in both radial growth rates and water use efficiency in recent decades. Summer water stress significantly reduced the growth of female trees in the following growing season, while the growth of male trees was primarily favoured by cloudy and rainy conditions the previous fall and winter combined with low cloud cover and warm conditions in summer. Sex-dependent lagged correlations between radial growth and water availability were found, with a strong association between tree growth and cumulative water availability in females at 30 months and in males at 10 months. CONCLUSIONS: Overall, our results point to greater vulnerability of female tress to increasing drought, which could lead to sex-ratio biases threatening population viability in the future.

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Background: Tree ontogeny is the genetic trajectories of regenerative processes in trees, repeating in time and space, including both development and reproduction. Understanding the principles of tree ontogeny is a key priority in emulating natural ecological patterns and processes that fall within the calls for closer-to-nature forest management. By recognizing and respecting the growth and development of individual trees and forest stands, forest managers can implement strategies that align with the inherent dynamics of forest ecosystem. Therefore, this study aims to determine the ontogenetic characteristics of tree regeneration and growth in northern European hemiboreal forests. Methodology: We applied a three-step process to review i) the ontogenetic characteristics of forest trees, ii) ontogenetic strategies of trees for stand-forming species, and iii) summarise the review findings of points i and ii to propose a conceptual framework for transitioning towards closer-to-nature management of hemiboreal forest trees. To achieve this, we applied the super-organism approach to forest development as a holistic progression towards the establishment of natural stand forming ecosystems. Results: The review showed multiple aspects; first, there are unique growth and development characteristics of individual trees at the pre-generative and generative stages of ontogenesis under full and minimal light conditions. Second, there are four main modes of tree establishment, growth and development related to the light requirements of trees; they were described as ontogenetic strategies of stand-forming tree species: gap colonisers, gap successors, gap fillers and gap competitors. Third, the summary of our analysis of the ontogenetic characteristics of tree regeneration and growth in northern European hemiboreal forests shows that stand-forming species occupy multiple niche positions relative to forest dynamics modes. Conclusions: This study demonstrates the importance of understanding tree ontogeny under the pretext of closer-to-nature forest management, and its potential towards formulating sustainable forest management that emulates the natural dynamics of forest structure. We suggest that scientists and foresters can adapt closer-to-nature management strategies, such as assisted natural regeneration of trees, to improve the vitality of tree communities and overall forest health. The presented approach prioritizes ecological integrity and forest resilience, promoting assisted natural regeneration, and fostering adaptability and connectivity among plant populations in hemiboreal tree communities.

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Background: Plants allocate resources to growth, defense, and stress resistance, and resource availability can affect the balance between these allocations. Allocation patterns are well-known to differ among species, but what controls possible intra-specific trade-offs and if variation in growth vs. defense potentially evolves in adaptation to resource availability. Methods: We measured growth and defense in a provenance trial of rubber trees (Hevea brasiliensis) with clones originating from the Amazon basin. To test hypotheses on the allocation to growth vs. defense, we relate biomass growth and latex production to wood and leaf traits, to climate and soil variables from the location of origin, and to the genetic relatedness of the Hevea clones. Results: Contrary to expectations, there was no trade-off between growth and defense, but latex yield and biomass growth were positively correlated, and both increased with tree size. The absence of a trade-off may be attributed to the high resource availability in a plantation, allowing trees to allocate resources to both growth and defense. Growth was weakly correlated with leaf traits, such as leaf mass per area, intrinsic water use efficiency, and leaf nitrogen content, but the relative investment in growth vs. defense was not associated with specific traits or environmental variables. Wood and leaf traits showed clinal correlations to the rainfall and soil variables of the places of origin. These traits exhibited strong phylogenetic signals, highlighting the role of genetic factors in trait variation and adaptation. The study provides insights into the interplay between resource allocation, environmental adaptations, and genetic factors in trees. However, the underlying drivers for the high variation of latex production in one of the commercially most important tree species remains unexplained.

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Understanding the biodiversity-productivity relationship (BPR) is crucial for biodiversity conservation and ecosystem management. While it is known that diversity enhances forest productivity, the underlying mechanisms at the local neighborhood level remain poorly understood. We established a 9.6 ha dynamic forest plot to study how neighborhood diversity, intraspecific competition, and interspecific competition influence tree growth across spatial scales using linear mixed-effects models. Our analysis reveals a significant positive correlation between neighborhood species richness (NSR) and relative growth rate (RGR). Notably, intraspecific competition, measured by conspecific neighborhood density and resource competition, negatively impacts RGR at finer scales, indicating intense competition among conspecifics for limited resources. In contrast, interspecific competition, measured by heterospecific density and resource competition, has a negligible impact on RGR. The relative importance of diversity and intra/interspecific competition in influencing tree growth varies with scale. At fine scales, intraspecific competition dominates negatively, while at larger scales, the positive effect of NSR on RGR increases, contributing to a positive BPR. These findings highlight the intricate interplay between local interactions and spatial scale in modulating tree growth, emphasizing the importance of considering biotic interactions and spatial variability in studying BPR.

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Understanding how different mechanisms act and interact in shaping communities and ecosystems is essential to better predict their future with global change. Disturbance legacy, abiotic conditions, and biotic interactions can simultaneously influence tree growth, but it remains unclear what are their relative contributions and whether they have additive or interactive effects. We examined the separate and joint effects of disturbance intensity, soil conditions, and neighborhood crowding on tree growth in 10 temperate forests in northeast China. We found that disturbance was the strongest driver of tree growth, followed by neighbors and soil. Specifically, trees grew slower with decreasing initial disturbance intensity, but with increasing neighborhood crowding, soil pH and soil total phosphorus. Interestingly, the decrease in tree growth with increasing soil pH and soil phosphorus was steeper with high initial disturbance intensity. Testing the role of species traits, we showed that fast-growing species exhibited greater maximum tree size, but lower wood density and specific leaf area. Species with lower wood density grew faster with increasing initial disturbance intensity, while species with higher specific leaf area suffered less from neighbors in areas with high initial disturbance intensity. Our study suggests that accounting for both individual and interactive effects of multiple drivers is crucial to better predict forest dynamics.

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The climate change scenarios RCP 4.5 and RCP 8.5, with a representative concentration pathway for stabilization of radiative forcing of 4.5 W m-2 and 8.5 W m-2 by 2100, respectively, predict an increase in temperature of 1-4.5° Celsius for Europe and a simultaneous shift in precipitation patterns leading to increased drought frequency and severity. The negative consequences of such changes on tree growth on dry sites or at the dry end of a tree species distribution are well-known, but rarely quantified across large gradients. In this study, the growth of Quercus robur and Quercus petraea (Q. spp.) and Pinus sylvestris in pure and mixed stands was predicted for a historical scenario and the two climate change scenarios RCP 4.5 and RCP 8.5 using the individual tree growth model PrognAus. Predictions were made along an ecological gradient ranging from current mean annual temperatures of 5.5-11.4 °C and with mean annual precipitation sums of 586-929 mm. Initial data for the simulation consisted of 23 triplets established in pure and mixed stands of Q. spp. and P. sylvestris. After doing the simulations until 2100, we fitted a linear mixed model using the predicted volume in the year 2100 as response variable to describe the general trends in the simulation results. Productivity decreased for both Q. spp. and P. sylvestris with increasing temperature, and more so, for the warmer sites of the gradient. P. sylvestris is the more productive tree species in the current climate scenario, but the competitive advantage shifts to Q. spp., which is capable to endure very high negative water potentials, for the more severe climate change scenario. The Q. spp.-P. sylvestris mixture presents an intermediate resilience to increased scenario severity. Enrichment of P. sylvestris stands by creating mixtures with Q. spp., but not the opposite, might be a right silvicultural adaptive strategy, especially at lower latitudes. Tree species mixing can only partly compensate productivity losses due to climate change. This may, however, be possible in combination with other silvicultural adaptation strategies, such as thinning and uneven-aged management.

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It is well established that solar irradiance greatly influences tree metabolism and growth through photosynthesis, but its effects acting through individual climate metrics have not yet been well quantified. Understanding these effects is crucial for assessing the impacts of climate change on forest ecosystems. To describe the effects of solar irradiance on tree growth, we installed 110 automatic dendrometers in two old-growth mountain forest reserves in Central Europe, performed detailed terrestrial and aerial laser scanning to obtain precise tree profiles, and used these to simulate the sum of solar irradiance received by each tree on a daily basis. Generalized linear mixed-effect models were applied to simulate the probability of growth and the growth intensity over seven growing seasons. Our results demonstrated various contrasting effects of solar irradiance on the growth of canopy trees. On the one hand, the highest daily growth rates corresponded with the highest solar irradiance potentials (i.e. the longest photoperiod). Intense solar irradiance significantly decreased tree growth, through an increase in the vapor pressure deficit. These effects were consistent for all species but had different magnitude. Tree growth is the most effective on long rainy/cloudy days with low solar irradiance.

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Tree growth is a key mechanism driving carbon sequestration in forest ecosystems. Environmental conditions are important regulators of tree growth that can vary considerably between nearby urban and rural forests. For example, trees growing in cities often experience hotter and drier conditions than their rural counterparts while also being exposed to higher levels of light, pollution, and nutrient inputs. However, the extent to which these intrinsic differences in the growing conditions of trees in urban versus rural forests influence tree growth response to climate is not well known. In this study, we tested for differences in the climate sensitivity of tree growth between urban and rural forests along a latitudinal transect in the eastern United States that included Boston, Massachusetts, New York City, New York, and Baltimore, Maryland. Using dendrochronology analyses of tree cores from 55 white oak trees (Quercus alba), 55 red maple trees (Acer rubrum), and 41 red oak trees (Quercus rubra) we investigated the impacts of heat stress and water stress on the radial growth of individual trees. Across our three-city study, we found that tree growth was more closely correlated with climate stress in the cooler climate cities of Boston and New York than in Baltimore. Furthermore, heat stress was a significant hindrance to tree growth in higher latitudes while the impacts of water stress appeared to be more evenly distributed across latitudes. We also found that the growth of oak trees, but not red maple trees, in the urban sites of Boston and New York City was more adversely impacted by heat stress than their rural counterparts, but we did not see these urban-rural differences in Maryland. Trees provide a wide range of important ecosystem services and increasing tree canopy cover was typically an important component of urban sustainability strategies. In light of our findings that urbanization can influence how tree growth responds to a warming climate, we suggest that municipalities consider these interactions when developing their tree-planting palettes and when estimating the capacity of urban forests to contribute to broader sustainability goals in the future.

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Chestnut and chestnut byproducts are of worldwide interest, so there is a constant need to develop faster and more accurate monitoring techniques. Recent advances in simultaneous localization and mapping (SLAM) algorithms and user accessibility have led to increased use of handheld mobile laser scanning (HHLS) in precision agriculture. We propose a tree growth monitoring methodology, based on HHLS point cloud processing, that calculates the length of branches through spatial discretization of the point cloud for each tree. The methodology was tested by comparing two point clouds collected almost simultaneously for each of a set of sweet chestnut trees. The results obtained indicated that our HHLS method was reliable and accurate in efficiently monitoring sweet chestnut tree growth. The same methodology was used to calculate the growth of the same set of trees over 37 weeks (from spring to winter). Differences in week 0 and week 37 scans showed an approximate mean growth of 0.22 m, with a standard deviation of around 0.16 m reflecting heterogeneous tree growth.

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Understanding how nutrient addition affects the tree growth is critical for assessing forest ecosystem function and processes, especially in the context of increased nitrogen (N) and phosphorus (P) deposition. Subtropical forests are often considered N-rich and P-poor ecosystems, but few existing studies follow the traditional "P limitation" paradigm, possibly due to differences in nutrient requirements among trees of different size classes. We conducted a three-year fertilization experiment with four treatments (Control, N-treatment, P-treatment, and NP-treatment). We measured soil nutrient availability, leaf stoichiometry, and relative growth rate (RGR) of trees across three size classes (small, medium and large) in 64 plots. We found that N and NP-treatments increased the RGR of large trees. P-treatment increased the RGR of small trees. RGR was mainly affected by N addition, the total effect of P addition was only 10 % of that of N addition. The effect of nutrient addition on RGR was mainly regulated by leaf stoichiometry. This study reveals that nutrient limitation is size dependent, indicating that continuous unbalanced N and P deposition will inhibit the growth of small trees and increase the instability of subtropical forest stand structure, but may improve the carbon sink function of large trees.

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The fraction of photosynthetically assimilated carbon that trees allocate to long-lasting woody biomass pools (biomass production efficiency - BPE), is a key metric of the forest carbon balance. Its apparent simplicity belies the complex interplay between underlying processes of photosynthesis, respiration, litter and fruit production, and tree growth that respond differently to climate variability. Whereas the magnitude of BPE has been routinely quantified in ecological studies, its temporal dynamics and responses to extreme events such as drought remain less well understood. Here, we combine long-term records of aboveground carbon increment (ACI) obtained from tree rings with stand-level gross primary productivity (GPP) from eddy covariance (EC) records to empirically quantify aboveground BPE (= ACI/GPP) and its interannual variability in two European beech forests (Hainich, DE-Hai, Germany; Sorø, DK-Sor, Denmark). We found significant negative correlations between BPE and a daily-resolved drought index at both sites, indicating that woody growth is de-prioritized under water limitation. During identified extreme years, early-season drought reduced same-year BPE by 29 % (Hainich, 2011), 31 % (Sorø, 2006), and 14 % (Sorø, 2013). By contrast, the 2003 late-summer drought resulted in a 17 % reduction of post-drought year BPE at Hainich. Across the entire EC period, the daily-to-seasonal drought response of BPE resembled that of ACI, rather than that of GPP. This indicates that BPE follows sink dynamics more closely than source dynamics, which appear to be decoupled given the distinctive climate response patterns of GPP and ACI. Based on our observations, we caution against estimating the magnitude and variability of the carbon sink in European beech (and likely other temperate forests) based on carbon fluxes alone. We also encourage comparable studies at other long-term EC measurement sites from different ecosystems to further constrain the BPE response to rare climatic events.

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China has experienced history's largest rural-to-urban migration. The social, economic, and environmental challenges brought about by urbanization are diverse and complex. Given China's national goal to achieve carbon neutrality by 2060 and commitment to urban sustainability, large cities have focused on urban greening initiatives. Yet, studies seeking to quantify ecosystem services and disservices only assess healthy, mature trees, rather than those with severe damage, declining health, or lack of vitality due to poor management. In this short communication, we conducted a case-study in one of China's major nursery stock-producing cities, Chengdu, on a common street tree, Ginkgo biloba, to assess the long-term impact of one of the most common yet extreme nursery transplant practices on tree growth (traumatic root-cutting of 'super-large' nursery stock). We used tree-ring data collected in a typical urban greenspace from 23 Ginkgo trees, including 18 trees transplanted as 'super-large' nursery stock and a control group (5 trees) transplanted as small-caliper trees. We found the trees transplanted as 'super-large' nursery stock experienced declining tree growth with decades of lost landscape potential likely due to traumatic root-cutting at the time of transplant from nursery to landscape. The control group allowed contrast between the growth patterns of 'super-large' transplanted trees with those that remained healthy, being transplanted as smaller-caliper trees. For the 'super-large' trees, we found a decrease in carbon sequestration from 7.6 kg C yr-1 on average per tree in 2001 to about 1.5 kg C yr-1 on average per tree in 2021, while no decreasing trends were observed among the control trees. This implies a negative impact on multiple expected ecosystem services including carbon sequestration, shade, canopy coverage, and pollutant mitigation. These results highlight the unrecognized costs of common Chinese nursery and transplant techniques on urban landscape trees, necessitating more research, science-based policies, and better management techniques.

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Perennially frozen soil, also known as permafrost, is important for the functioning and productivity of most of the boreal forest, the world's largest terrestrial biome. A better understanding of complex vegetation-permafrost interrelationships is needed to predict changes in local- to large-scale carbon, nutrient, and water cycle dynamics under future global warming. Here, we analyze tree-ring width and tree-ring stable isotope (C and O) measurements of Gmelin larch (Larix gmelinii (Rupr.) Rupr.) from six permafrost sites in the northern taiga of central Siberia. Our multi-parameter approach shows that changes in tree growth were predominantly controlled by the air and topsoil temperature and moisture content of the active soil and upper permafrost layers. The observed patterns range from strong growth limitations by early summer temperatures at higher elevations to significant growth controls by precipitation at warmer and well-drained lower-elevation sites. Enhanced radial tree growth is mainly found at sites with fast thawing upper mineral soil layers, and the comparison of tree-ring isotopes over five-year periods with different amounts of summer precipitation indicates that trees can prevent drought stress by accessing water from melted snow and seasonally frozen soil. Identifying the active soil and upper permafrost layers as central water resources for boreal tree growth during dry summers demonstrates the complexity of ecosystem responses to climatic changes.

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The dominant paradigm for assessing ecological responses to climate change assumes that future states of individuals and populations can be predicted by current, species-wide performance variation across spatial climatic gradients. However, if the fates of ecological systems are better predicted by past responses to in situ climatic variation through time, this current analytical paradigm may be severely misleading. Empirically testing whether spatial or temporal climate responses better predict how species respond to climate change has been elusive, largely due to restrictive data requirements. Here, we leverage a newly collected network of ponderosa pine tree-ring time series to test whether statistically inferred responses to spatial versus temporal climatic variation better predict how trees have responded to recent climate change. When compared to observed tree growth responses to climate change since 1980, predictions derived from spatial climatic variation were wrong in both magnitude and direction. This was not the case for predictions derived from climatic variation through time, which were able to replicate observed responses well. Future climate scenarios through the end of the 21st century exacerbated these disparities. These results suggest that the currently dominant paradigm of forecasting the ecological impacts of climate change based on spatial climatic variation may be severely misleading over decadal to centennial timescales.

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Agroforestry systems have received a significant attention in recent years and can be considered as a potential strategy in agricultural production to respond to worsening climatic conditions. The decision-making process for farmers to design and implement agroforestry systems is complex due to time-consuming processes of planting, growing and management of trees, as well as the long-term impacts on the field and its productivity. The shading of the arable land by trees is a core issue and should be reduced through a north-south orientation of the tree rows. However, this orientation is often in conflict with other criteria. In order to consider future shading from different tree row orientations into the design process, the modelling tool "ShadOT" was developed. This tool can simulate tree growth and analyses spatial shading over variable time periods by using only a limited number of parameters. This tool was programmed exclusively with open source software and can therefore be easily extended. It offers an ideal platform for testing different agroforestry designs due to its simple approach and minimal parameterization. Two different designs (north-south and west-east orientation) were tested for a field and differences in the temporal and spatial distribution of shaded areas are presented.•Modelling tool for tree growth and shading effects is presented.•The tool is written in Python programming language, uses only open-source software and requires a limited number of inputs.•Identification of spatial-temporal shading patterns of different alley cropping scenarios.

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Scientific consensus is that diverse tree species positively impact forest productivity, especially when species are functionally dissimilar. Under the complementarity hypothesis, differences in species traits reduce competition among neighboring tree species. However, while this relationship has been extensively studied at the community level, there is a lack of understanding regarding how individuals of different species specifically respond to a functionally dissimilar neighborhood. In this study, we used permanent plots from Quebec, Canada, and 19 focal tree species to test whether: (1) tree growth response to neighborhood dissimilarity varies with their identity and competition intensity, and (2) focal tree species' traits explain their response to neighborhood dissimilarity. We demonstrate that: tree growth is primarily influenced by competition, species identity, and their interactions, but that dissimilarity, alone and in interaction with the main drivers of tree growth, explains an additional 1.8% of the variation in species growth. Within this context, (1) most species' respond positively to neighborhood dissimilarity, with magnitude being species and competition dependent, and (2) focal tree traits partly explain these dependencies, with shade-intolerant species benefiting most from dissimilar neighbors under high competition. Our study provides empirical support for the complementarity hypothesis, emphasizing the small but consistent positive effect of functional dissimilarity on tree growth in local neighborhoods. Our findings identify the species with the highest potential of benefiting from dissimilar neighbors but also demonstrate that the positive effect of neighborhood dissimilarity is not limited to a select few species with specific traits; rather, it is observed across a diverse range of species. The cumulative growth responses of individuals to functionally dissimilar neighbors may help explain the commonly observed higher productivity in more diverse communities.

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BACKGROUND: Declines in plant biodiversity often have negative consequences for plant community productivity, and it becomes increasingly acknowledged that this may be driven by shifts in soil microbial communities. So far, the role of fungal communities in driving tree diversity-productivity relationships has been well assessed in forests. However, the role of bacteria and archaea, which are also highly abundant in forest soils and perform pivotal ecosystem functions, has been less investigated in this context. Here, we investigated how tree and shrub richness affects stand-level tree productivity by regulating bacterial and archaeal community diversity and composition. We used a landscape-scale, subtropical tree biodiversity experiment (BEF-China) where tree (1, 2, or 4 species) and shrub richness (0, 2, 4, 8 species) were modified. RESULTS: Our findings indicated a noteworthy decline in soil bacterial α-diversity as tree species richness increased from monoculture to 2- and 4- tree species mixtures, but a significant increase in archaeal α-diversity. Additionally, we observed that the impact of shrub species richness on microbial α-diversity was largely dependent on the level of tree species richness. The increase in tree species richness greatly reduced the variability in bacterial community composition and the complexity of co-occurrence network, but this effect was marginal for archaea. Both tree and shrub species richness increased the stand-level tree productivity by regulating the diversity and composition of bacterial community and archaeal diversity, with the effects being mediated via increases in soil C:N ratios. CONCLUSIONS: Our findings provide insight into the importance of bacterial and archaeal communities in driving the relationship between plant diversity and productivity in subtropical forests and highlight the necessity for a better understanding of prokaryotic communities in forest soils. Video Abstract.

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Stomata have recently been theorized to have evolved strategies that maximize turgor-driven growth over plants' lifetimes, finding support through steady-state solutions in which gas exchange, carbohydrate storage and growth have all reached equilibrium. However, plants do not operate near steady state as plant responses and environmental forcings vary diurnally and seasonally. It remains unclear how gas exchange, carbohydrate storage and growth should be dynamically coordinated for stomata to maximize growth. We simulated the gas exchange, carbohydrate storage and growth that dynamically maximize growth diurnally and annually. Additionally, we test whether the growth-optimization hypothesis explains nocturnal stomatal opening, particularly through diel changes in temperature, carbohydrate storage and demand. Year-long dynamic simulations captured realistic diurnal and seasonal patterns in gas exchange as well as realistic seasonal patterns in carbohydrate storage and growth, improving upon unrealistic carbohydrate responses in steady-state simulations. Diurnal patterns of carbohydrate storage and growth in day-long simulations were hindered by faulty modelling assumptions of cyclic carbohydrate storage over an individual day and synchronization of the expansive and hardening phases of growth, respectively. The growth-optimization hypothesis cannot currently explain nocturnal stomatal opening unless employing corrective 'fitness factors' or reframing the theory in a probabilistic manner, in which stomata adopt an inaccurate statistical 'memory' of night-time temperature. The growth-optimization hypothesis suggests that diurnal and seasonal patterns of stomatal conductance are driven by a dynamic carbon-use strategy that seeks to maintain homeostasis of carbohydrate reserves.