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Diverse plant communities are often more productive than mono-specific ones. Several possible mechanisms underlie this phenomenon but their relative importance remains unknown. Here we investigated whether light interception alone or in combination with light use efficiency (LUE) of dominant and subordinate species explained greater productivity of mixtures relative to monocultures (i.e. overyielding) in 108 young experimental tree communities. We found mixed-species communities that intercepted more light than their corresponding monocultures had 84% probability of overyielding. Enhanced LUE, which arose via several pathways, also mattered: the probability of overyielding was 71% when, in a mixture, species with higher 'inherent' LUE (i.e. LUE in monoculture) intercepted more light than species with lower LUE; 94% when dominant species increased their LUE in mixture; and 79% when subordinate species increased their LUE. Our results suggest that greater light interception and greater LUE, generated by inter and intraspecific variation, together drive overyielding in mixed-species forests.

Reduction pruning of the main stem is commonly used during the maintenance of power lines to encourage the establishment and development of scaffold limbs away from wires. Understanding the physiology of epicormic branch initiation and growth as well as wound compartmentalization following reduction pruning are important for optimizing the pruning cycle and maintaining healthy and safe trees. In this study, the influence of both intensity and time of year of pruning on epicormic branch response and wound compartmentalization was investigated on 56 11-year-old Pennsylvania ash trees (Fraxinus pennsylvanica Marsh.) about 5 to 7 m in height within a controlled nursery environment. During the second growing season following reduction of the main stem, the number, height, and volume of epicormic branches, as well as tallest epicormic branches and the area of discolored wood, increased with pruning intensity. Pruning during the leaf-on season compared to the leaf-off season limited the establishment and development of epicormic branches without affecting wound-closure rate or the area of wood discoloration at the cutting point. Results are consistent with the known seasonal fluctuation of carbohydrates reserves. In the context of the electrical distribution network, where trees are subjected to pruning throughout the year, trees pruned in summer during a maintenance cycle could be pruned during the next cycle, in winter, and so on, to optimize the return interval of the pruning cycle.

Habitat trees, which provide roosting, foraging and nesting for multiple taxa, are retained in managed forests to support biodiversity conservation. To what extent their spatial distribution influences provisioning of habitats has rarely been addressed. In this study, we investigated whether abundance and richness of tree-related microhabitats (TreMs) differ between habitat trees in clumped and dispersed distributions and whether the abundance of fifteen groups of TreMs is related to tree distribution patterns. To identify habitat trees, we quantified TreMs in temperate mountain forests of Germany. We determined clumping (the Clark–Evans index), size of the convex hull, diameter at breast height, as well as altitude, slope and aspect of sites for their possible influence on TreMs. We additionally determined the difference in TreM abundance and richness among four options of selecting five habitat trees per ha from 15 candidates: (a) the most clumped trees, (b) five randomly selected and dispersed trees, (c) the single tree with highest abundance or richness of TreMs and its four closest neighbors and (d) a “reference selection” of five trees with known highest abundance or richness of TreMs irrespective of their distribution. The degree of clumping and the size of the convex hull influenced neither the abundance nor richness of TreMs. The reference selection, option (d), contained more than twice the number of TreMs compared to the most clumped, (a), or random distributions, (b), of five habitat trees, while option (c) assumed an intermediate position. If the goal of habitat tree retention is to maximize stand-level abundance and richness of TreMs, then it is clearly more important to select habitat trees irrespective of their spatial pattern.

Niche complementarity, mass-ratio, and vegetation quantity effects have been identified as major drivers of the biodiversity-ecosystem functioning (BEF) relationships. However, their relative contribution to biomass and productivity is not yet clear in temperate secondary forests. Based on the observations from a 21.12-ha temperate secondary forest plot in northeastern China, we assessed how these mechanisms regulate forest biomass and productivity. The niche complementarity effect was quantified using a functional diversity metric that was calculated from six locally collected functional traits. The mass-ratio effect was described as functional trait composition using community-weighted mean trait values. Vegetation quantity effect was evaluated using vegetation biomass. We performed structural equation modeling to test the alternative mechanisms. Our results provide evidence for all three mechanisms. Functional diversity increased forest productivity, in line with the niche complementarity hypothesis. Acquisitive traits (e.g., greater specific leaf area and leaf nitrogen concentration) enhance productivity, while conservative traits (e.g., greater wood density) enhance the long-term accumulation of biomass, demonstrating the mass-ratio hypothesis. Furthermore, we observed a significant positive relationship between biomass and productivity, confirming the vegetation quantity hypothesis. We conclude that functional traits drive biomass and productivity through multiple mechanisms. Both niche complementarity and the mass-ratio effects play roles in this temperate secondary forest. In addition, we emphasize the importance of preserving sufficient biomass stock to ensure maximum productivity in secondary forests. Our study contributes to the identification of the mechanisms underlying BEF relationships and has practical significance for guiding temperate secondary forest management and conservation.

Trees that grow in urban areas are confronted with a wide variety of stresses that undermine their long-term survival. These include mechanical damage to the crown, root reduction and stem injury, all of which remove significant parts of plant tissues. The single or combined effects of these stresses generate a complex array of growth and ecophysiological responses that are hard to predict. Here we evaluated the effects of different individual and combined damage on the dynamics of non-structural carbohydrates (NSC, low weight sugars plus starch) concentration and new tissue growth (diameter increment) in young trees. We hypothesized that (i) tissue damage will induce larger reductions in diameter growth than in NSC concentrations and (ii) combinations of stress treatments that minimally alter the “functional equilibrium” (e.g., similar reductions of leaf and root area) would have the least impact on NSC concentrations (although not on growth) helping to maintain tree health and integrity. To test these hypotheses, we set up a manipulative field experiment with 10-year-old trees of common urban species (Celtis occidentalis, Fraxinus pennsylvanica, and Tilia cordata). These trees were treated with a complete array of mechanical damage combinations at different levels of intensity (i.e., three levels of defoliation and root reduction, and two levels of stem damage). We found that tree growth declined in relation to the total amount of stress inflicted on the trees, i.e., when the combined highest level of stress was applied, but NSC concentrations were either not affected or, in some cases, increased with an increasing level of stress. We did not find a consistent response in concentration of reserves in relation to the combined stress treatments. Therefore, trees appear to reach a new “functional equilibrium” that allows them to adjust their levels of carbohydrate reserves, especially in stems and roots, to meet their metabolic demand under stressful situations. Our results provide a unique insight into the carbon economy of trees facing multiple urban stress conditions in order to better predict long-term tree performance and vitality.

Trees display contrasting specialized annual shoots during their life-span and along with ontogeny-driven modifications in their branching pattern, they can fulfill different combinations of light exploitation and space exploration functions Tree ontogeny is related to major changes in tree structure and function at different scales, from individual organs to the whole tree. Yet, little is known about the direct effects of tree ontogeny on shoot specialization and branching patterns. Such specific architectural changes occurring with tree growth and aging are of critical importance for understanding the response of trees to their environment. The uppermost branching system of 0.1- to 23-m-tall sugar maple trees was sampled at the end of the growing season. Measurements were made at both the branching system (n = 40) and annual shoot scales (n = 803). An algorithm for automated shoot typology was developed to characterize branching pattern variations. Sugar maple shoots were divided into four types with contrasting sizes and levels of foliage (i.e., relative biomass allocation into leaves, LMF). These morphological differences were interpreted as functional specializations for light exploitation (high LMF) or space exploration and support (low LMF). Only annual trunk shoots exhibited trait value changes during ontogeny such as a minimum allocation to foliage in the current-year shoots for the 5-m-tall trees, which is related to lower light interception capabilities but higher space exploration abilities. However, this relative loss of light interception function is compensated by ontogenetic changes at the branching system scale, which are associated with higher rates of ramification to produce lateral shoots. This study reveals how branching system and annual shoot traits change simultaneously during tree ontogeny to fulfill different functions, particularly light exploitation and space exploration.

SummaryBackground and AimsCarbohydrate reserves play a vital role in plant survival during periods of negative carbon balance. Considering active storage of reserves, there is a trade-off between carbon allocation to growth and to reserves and defense. A resulting hypothesis is that allocation to reserves exhibits a coordinated variation with functional traits associated with the ‘fast-slow’ plant economics spectrum.MethodsWe tested the relationship between non-structural carbohydrates (NSC) of tree organs and functional traits using 61 angiosperm tree species from temperate and tropical forests with phylogenetic hierarchical Bayesian models.Key ResultsOur results provide evidence that NSC concentrations in woody organs and plant functional traits are largely decoupled, meaning that species’ resilience is unrelated to their position on the ‘fast-slow’ plant economics spectrum. In contrast, we found that variation between NSC concentrations in leaves and the fast-slow continuum was coordinated, as species with higher leaf NSC had traits values associated with resource conservative species such as lower SLA, lower Amax, and high wood density. We did not detect an influence of leaf habit on the variation of NSC concentrations in tree organs.ConclusionsEfforts to predict the response of ecosystems to global change will need to integrate a suite of plant traits, such as NSC concentrations in woody organs, that are independent of the ‘fast-slow’ spectrum and that capture how species respond to a broad range of global change factors.

An extreme summer drought caused unprecedented tree dieback across Central Europe in 2018, highlighting the need for a better mechanistic understanding of drought-induced tree mortality. While numerous physiological risk factors have been identified, the principal mechanisms, hydraulic failure and carbon starvation, are still debated. We studied 9,435 trees from 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighbourhood competition influence individual mortality risk. We observed a reduced mortality risk for trees with wider hydraulic safety margins, while a rising sugar fraction of the non-structural carbohydrate pool and bark beetle infestation were associated with higher risk. Taller trees had a lower mortality risk. The sign and magnitude of neighbourhood effects on mortality risk depended on the species-identity of the involved trees, with most species having beneficial and some having deleterious effects on their neighbours. While severe tissue dehydration causing hydraulic failure precedes drought-induced tree mortality, we show that the probability of this event depends on a series of mutually inclusive processes including pest infestation and starch depletion for osmotic adjustment, and is modulated by the size and species identity of a tree and its neighbours.

Salix species are widely used for wood production, but the interactive effects of soil types and atmospheric conditions on Salix foliar nutrients and aboveground biomass productivity have not yet been elucidated. The objectives of this study were fourfold: (1) to assess the variation in foliar nutrients and leaf mass of Salix miyabeana ‘SX67’ grown as short rotation coppice (SRC) across three growing seasons and nine locations in Quebec with different permanent site characteristics and atmospheric conditions, (2) to test if atmospheric conditions and permanent site characteristics could explain the variation in foliar nutrients and leaf mass, (3) to develop models that consider foliar traits and nutrient interactions to produce more robust predictions of annual aboveground biomass yields, and (4) to compare nutritional requirements of ‘SX67’ to other Salix cultivars used for SRC. Leaf samples were collected over three growing seasons at all sites. For each site, atmospheric conditions were simulated and foliar nutrient levels were measured to perform centered log ratio (clr) transformations for each foliar nutrient. This approach considered foliar nutrient interactions and dealt with clr scores as linearly independent. The clr scores were more largely influenced by permanent site characteristics than by atmospheric conditions, despite large variations in degree-days. However, some foliar nutrients and leaf mass were linearly related to atmospheric variables within sites. Strong relationships between annual aboveground biomass yields and leaf mass were computed (e.g. adjusted R2 = 0.62), likely due to a proportional allocation between foliage and wood. Although significant linear relationships between clr scores (i.e. N, Ca and Mn) and annual aboveground biomass yields were detected, yields were more robustly explained non-linearly by thresholds (i.e. N, Ca and P) (e.g. R2 = 0.85), likely due to permanent characteristics specific to each of the sites and climatic limitations during the growing seasons studied. The thresholds detected by non-linear models suggested high N and P use efficiencies and a large Ca requirement of ‘SX67’.

The ability of plants to tolerate stress is determined in part by the carbon allocated to their reserves. We studied two common urban tree species in northeastern North America, Acer saccharinum (Silver maple, native) and Acer platanoides (Norway maple, exotic), to assess the dynamics of non-structural carbohydrate (NSC) concentrations immediately following a maintenance pruning of 30% of the tree crown. NSC concentrations were measured by high-performance liquid chromatography in branches, main stems, and root tissues for both pruned and un-pruned trees at three intervals during the growing season. NSC concentrations in tree organs of A. platanoides were 75% higher than in A. saccharinum. Maintenance pruning did not have any significant depletion effect on carbohydrate concentrations in the tissues of either species. Yet, there was a significant increase in the NSC concentrations in un-pruned branches of pruned trees of A. platanoides at the end of the growing season, but no effect was observed in A. saccharinum. Higher levels of carbohydrates after pruning in woody plant tissues suggested that A. platanoides may have compensatory mechanisms that allow this species to respond better to maintenance pruning than A. saccharinum. © 2018 Elsevier GmbH

The increasing dominance of American beech regeneration (Fagus grandifolia Ehrh.) to the expense of sugar maple (Acer saccharum Marsh.) challenges the long-term economic viability of forest management in northern hardwoods. Based on earlier studies showing that more light in the understory favours sugar maple over beech, the irregular shelterwood system has shifted from experimental to operational over the last decade. In this paper, we evaluate the success of this shift toward irregular shelterwood for promoting sugar maple over beech by measuring the frequency, abundance of sugar maple and beech seedlings and saplings, and growth of seedlings three to six years after logging in northern hardwood forests of western Quebec, Canada. Results showed a dominance of beech regeneration regardless of the harvest intensity, particularly among tall seedlings and saplings. However, we found that the transition probability indices (projected relative combined abundance and growth) of sugar maple could be favoured, albeit to a limited extent, by an increased basal area removal, particularly where the initial occurrence of sugar maple in the advance seedling regeneration is lower than 60% and the initial beech basal area is low in the overstory (i.e.

Reduction pruning of main tree stems is commonly performed in the maintenance of an electricity distribution network to encourage the growth of scaffold limbs away from the wires. Understanding both the initiation and growth of epicormic branch after reduction pruning interventions, as well as the area of discolored wood at the cutting point are important for optimizing the pruning cycle and maintaining safe trees. In this study, we investigated post-pruning traumatic responses in relation to tree vitality before pruning in 116 hackberries (Celtis occidentalis L.) and 86 green ash (Fraxinus pennsylvanica Marsh.) growing in a controlled environment of a nursery. Four years before pruning, each tree of both species was assigned a degree of damage commonly occurring in urban areas, including root injury, defoliation, and stem injury, for a total of 18 treatment combinations arranged in a random block design. In both species, trees that had lower vitality prior to pruning had a reduced number, height, and volume of the epicormic branches at the cutting point two years after main stem pruning. Interestingly and contrary to expectations, a positive relationship was found between the area of discolored wood at the cutting point and tree vitality prior to pruning. However, below a vitality threshold, the area of wood discoloration was exacerbated by crown dieback and retrenchment. Celtis showed a less efficient ability to compartmentalize and produced fewer epicormic branches compared to Fraxinus, and epicormic branches were of reduced height and volume when low growth was observed. From a management perspective, although both species can survive after several damage events and after reduction of the main stem, the results suggest to conduct an assessment of tree vitality before pruning to identify trees with lower vitality and better predict the return pruning time.

Plantations of fast-growing hybrid trees, such as hybrid poplars and hybrid larch, are increasingly used for wood and timber production, but they are also believed to impair forest biodiversity. Most studies that have assessed how such plantations may alter the diversity and composition of understorey plants were established in agricultural landscapes or have compared tree plantations with old-growth natural forests. Moreover, many important aspects of biodiversity have been overlooked in previous studies, such as functional and beta-diversity. Here, we present results from a study that was aimed at quantifying alpha- and beta-diversity of understorey plant species and functional groups in hybrid poplar (9–10 years) and hybrid larch plantations (16 years) located within a forested landscape of Quebec, Canada. These hybrid plantations were compared to naturally regenerated secondary forests and to native plantations of black spruce of the same origin (clear cut) and similar age. Our results indicate that fast-growing hybrid plantations do not present lower taxonomic and functional alpha-biodiversity indices, but may harbour more diverse communities, in part through the introduction of plant species that are associated with open habitats. We provide further evidence that planted forests may be as heterogeneous as naturally regenerated forests in terms of understorey plant composition. Plant species and functional composition differed slightly between stand types (naturally regenerated forests, native and fast-growing hybrid plantations), with plantations offering a greater potential for colonisation by ruderal species, while being detrimental to species of closed forest habitats. Lastly, plantations of fast-growing hybrids do not induce greater changes in understorey vegetation relative to native plantations of black spruce, at least during the first stand rotation.

Studies of biodiversity–ecosystem function in treed ecosystems have generally focused on aboveground functions. This study investigates intertrophic links between tree diversity and soil microbial community function and composition. We examined how microbial communities in surface mineral soil responded to experimental gradients of tree species richness (SR), functional diversity (FD), community‐weighted mean trait value (CWM), and tree identity. The site was a 4‐year‐old common garden experiment near Montreal, Canada, consisting of deciduous and evergreen tree species mixtures. Microbial community composition, community‐level physiological profiles, and respiration were evaluated using phospholipid fatty acid (PLFA) analysis and the MicroResp™ system, respectively. The relationship between tree species richness and glucose‐induced respiration (GIR), basal respiration (BR), metabolic quotient (qCO 2) followed a positive but saturating shape. Microbial communities associated with species mixtures were more active (basal respiration [BR]), with higher biomass (glucose‐induced respiration [GIR]), and used a greater number of carbon sources than monocultures. Communities associated with deciduous tree species used a greater number of carbon sources than those associated with evergreen species, suggesting a greater soil carbon storage capacity. There were no differences in microbial composition (PLFA) between monocultures and SR mixtures. The FD and the CWM of several functional traits affected both BR and GIR. In general, the CWM of traits had stronger effects than did FD, suggesting that certain traits of dominant species have more effect on ecosystem processes than does FD. Both the functions of GIR and BR were positively related to aboveground tree community productivity. Both tree diversity (SR) and identity (species and functional identity—leaf habit) affected soil microbial community respiration, biomass, and composition. For the first time, we identified functional traits related to life‐history strategy, as well as root traits that influence another trophic level, soil microbial community function, via effects on BR and GIR.

Soil and climatic conditions for optimizing aboveground biomass yields of bioenergy short rotation coppices (SRCs) of Salix are not well elucidated. The objective of this study was to identify and quantify the limitations induced by soil and climate, and compare the magnitude of their effects, on annual aboveground yields across ten SRCs of Salix miyabeana SX67 in Quebec, Canada. The effects of weather variation between years on yields were also tested within locations. In five plots per SRC, soil bulk density, particle size, exchangeable cations and bulk composition were analysed, and moisture deficits were estimated using leaf δ13C. For each location, numerous weather variables were simulated for spring, summer and the whole growing season. Climate was calculated by averaging weather variables for growing seasons for which annual yields were available. Annual aboveground biomass yields were modelled using linear regression, partitioning of the variance and mixed models with soil, weather and climate variables as predictors. Across SRCs, silt content, soil organic matter, pH, exchangeable Ca and Mg, and total N and Zn were significantly and positively related to aboveground yields (adj. R 2 ranging from 0.38 to 0.79). Generally, annual yields were negatively related to summer temperature within SRCs (adj. R 2 = 0.92) and drought across SRCs (adj. R 2 = 0.54). Partitioning of the variance revealed that soil variables (~80%) had a greater effect on productivity than did climate variables (~10%). In fact, soil properties buffered or exacerbated water shortages and, thus, had a preponderant effect on yield.

Tree plantations are commonly used to restore abandoned agricultural fields with varying degrees of success. Agricultural soils differ from forest soils in nutrient availability and microbial communities. The objective of this study was to test the effect of adding small amounts of forest soil on the survival, growth and rates of mycorrhizal fungal colonization of trees planted in an abandoned agricultural field over the crucial first three growing seasons. Seedlings of two arbuscular mycorrhizal (AM) and two ectomycorrhizal (EM) tree species were planted in an abandoned agricultural field. Soil inocula were taken from four forest stands, each dominated by one of the planted species. Half of the soil samples were sterilized before inoculation to distinguish microbial from nutrient effects. The effect of the quantity of soil inoculum added was tested using 300 and 1500 ml of forest soil. Tree mortality was low and did not vary between treatments. The growth of EM tree species responded, positively or negatively, to forest soil inoculation. A negative feedback was detected on the growth of red oak seedlings inoculated with red oak soil. Seedlings inoculated with EM sterilized soils were smaller than control seedlings, presumably due to lower nutrient availability of EM forest soils compared to agricultural field soil. The majority of the effects, either positive or negative, were observed the first year. After three seasons of growth, only yellow birch seedlings that had received 1500 ml of non-sterilized red oak soil still benefited from soil inoculation. More research is needed in nutrient-limited soils to determine whether inoculation would have greater or longer term benefits on tree survival and growth.

The decomposition of plant material is an important ecosystem process influencing both carbon cycling and soil nutrient availability. Quantifying how plant diversity affects decomposition is thus crucial for predicting the effect of the global decline in plant diversity on ecosystem functioning. Plant diversity could affect the decomposition process both directly through the diversity of the litter, and/or indirectly through the diversity of the host plant community and its affect on the decomposition environment. Using a biodiversity experiment with trees in which both functional and taxonomic diversity were explicitly manipulated independently, we tested the effects of the functional diversity and identity of the living trees separately and in combination with the functional diversity and identity of the decomposing litter on rates of litter decomposition and soil respiration. Plant traits, predominantly leaf chemical and physical traits, were correlated with both litter decomposition and soil respiration rates. Surface litter decomposition, quantified by mass loss in litterbags, was best explained by abundance-weighted mean trait values of tree species from which the litter was assembled (functional identity). In contrast, soil respiration, which includes decomposition of dissolved organic carbon and root respiration, was best explained by the variance in trait values of the host trees (functional diversity). This research provides insight into the effect of loss of tree diversity in forests on soil processes. Such understanding is essential to predicting changes in the global carbon budget brought on by biodiversity loss.

Urban trees are subjected to numerous biotic and mechanical damage, which can affect their growth rates and health. However, for most species, a systematic analysis of tree above- and below-ground growth reactions to a variety of damages is still lacking. Under a fully factorial experimental setup, using two common urban trees (Celtis occidentalis, Fraxinus pennsylvanica), we tested the effects of various degrees of frequently occurring damage as defoliation, root reduction and stem injuries for a total of 18 treatments. We hypothesized that i) an increasing amount of damage would proportionally negatively affect both root and stem growth, ii) there would be a lag or lasting effect on growth, and iii) both species would react similarly to the treatments. Contrary to our expectation, increasing levels of single or combined damage did not have an incremental effect on either stem or root growth. Although Celtis was significantly less vigorous than Fraxinus, it did not react strongly to damage treatments compared to the control. Interestingly, Celtis that experienced stem damage alone or in combination with other damages showed higher growth rates than the control. For Celtis, root injury was the treatment having the most impact, decreasing both root and stem growth consistently throughout the 5 years following treatments, whereas defoliation decreased growth only in the first two years. All damage treatments negatively affected stem and root growth of Fraxinus trees. Stem growth was affected the most by defoliation in the first year following the treatment, while root injury became the driving factor in subsequent years. For both species, stem injury showed the least influence on growth rates. The control and low-level damage treatments often affected growth rates in a similar way, suggesting that low-intensity stress triggers compensatory reactions stimulating photosynthetic rates and nutrient utilization. The slower-growing tree species, Celtis, showed a less negative reaction to all damage treatments compared to Fraxinus. This study illustrates that various types of above- and below-ground injuries do not have a simple additive effect on tree growth and that trees are capable of compensating for the loss of foliage, roots, or phloem to meet their metabolic demand.

Considering intraspecific trait variability (ITV) in ecological studies has improved our understanding of species persistence and coexistence. These advances are based on the growing number of leaf ITV studies over local gradients, but logistical constraints have prevented a solid examination of ITV in root traits or at scales reflecting species’ geographic ranges. We compared the magnitude of ITV in above‐ and below‐ground plant organs across three spatial scales (biophysical region, locality and plot). We focused on six understorey species (four herbs and two shrubs) that occur both in disturbed and undisturbed habitats across boreal and temperate Canadian forests. We aimed to document ITV structure over broad ecological and geographical scales by asking: (a) What is the breadth of ITV across species range‐scale? (b) What proportion of ITV is captured at different spatial scales, particularly when local scale disturbances are considered? and (c) Is the variance structure consistent between analogous leaf and root traits, and between morphological and chemical traits? Following standardized methods, we sampled 818 populations across 79 forest plots simultaneously, including disturbed and undisturbed stands, spanning four biophysical regions (~5,200 km). Traits measured included specific leaf area (SLA), specific root length (SRL) and leaf and root nutrient concentrations (N, P, K, Mg, Ca). We used variance decomposition techniques to characterize ITV structure across scales. Our results show that an important proportion of ITV occurred at the local scale when sampling included contrasting environmental conditions resulting from local disturbance. A certain proportion of the variability in both leaf and root traits remained unaccounted for by the three sampling scales included in the design (36% on average), with the largest amount for SRL (54%). Substantial differences in magnitude of ITV were found among the six species, and between analogous traits, suggesting that trait distribution was influenced by species strategy and reflects the extent of understorey environment heterogeneity. Even for species with broad geographical distributions, a large proportion of within‐species trait variability can be captured by sampling locally across ecological gradients. This has practical implications for sampling design and trait selection for both local studies and continental‐scale modelling. A free Plain Language Summary can be found within the Supporting Information of this article.

The discipline of ecology suffers from a lack of knowledge of non-climatic factors (for example, plant–soil, plant–plant and plant–insect interactions) to predict tree species range shifts under climate change. The next generation of simulation models of forest response to climate change must build upon local observations of species interactions and growth along climatic gradients. We examined whether sugar maple (Acer saccharum) seedlings were disadvantaged with respect to soil nutrient uptake under coniferous canopies, as this species would need to migrate northward into conifer-dominated forests in response to climate change. An experimental design was applied to 3 sites, forming the largest possible latitudinal/climatic gradient for sugar maple in Quebec (Canada) and isolating the effect of conifer presence on its seedling’s nutritional status. We tested whether: (1) both soil and climate and (2) presence of conifers affected foliar nutrient levels of sugar maple seedlings. Climate and soil (through pH) strongly affected nutrient availability for sugar maple seedlings and predicted 63.7% of their foliar nutrient variability. When controlling for site effects, we found a significant negative effect of conifers on foliar Ca and Mg levels of maple seedlings, which can adversely affect their overall health and vigour. When considering projected modifications of the forest environment due to climate change, we suggest that northward migration of sugar maple will be negatively affected by the presence of conifers through reduced foliar nutrition.

We studied three hybrid poplar plantations in Quebec (Canada) established on sites with varying soil and environmental characteristics to investigate the effects of logging residues on the water potential, carbon isotope ratio and foliar nutrients of planted trees. On each site, four treatments representing different residue loads, as well as treatments aimed at manipulating specific factors of the environment (Herbicide, Geotextile) were applied to test their effects on seedling water potential, carbon isotope ratio and foliar nutrients. Along with analyses of variance, we used structural equation modelling to infer causal relationships of logging residues on height, basal diameter and foliar nutrition of trees through their effects on soil temperature, soil water content and competing vegetation cover. Logging residues decreased soil temperature at all sites and woody plants cover at one site out of three. Height, basal diameter and unit leaf mass were strongly related to each other. Foliar δ13C, N concentration and unit leaf mass increased with decreasing cover of woody plants suggesting an important role of competition for resources. Overall, logging residues had no direct influence on hybrid poplar dimensions after two growing seasons: their effects on the microenvironment of this resource demanding tree species were either cancelling out each other, or were not significant enough to have a significant impact on the growth drivers measured. For example, presence of logging residues might reduce soil temperature, impeding overall seedling performance. Our study highlights the fact that any given silvicultural method aimed at manipulating logging residues has a complex influence involving the interaction of multiple environmental drivers and that the net effect on tree productivity will depend on species and site specific conditions.

Exploring the link between above- and belowground biodiversity has been a major theme of recent ecological research, due in large part to the increasingly well-recognized role that soil microorganisms play in driving plant community processes. In this study, we utilized a field-based tree experiment in Minnesota, USA, to assess the effect of changes in plant species richness and phylogenetic diversity on the richness and composition of both ectomycorrhizal and saprotrophic fungal communities. We found that ectomycorrhizal fungal species richness was significantly positively influenced by increasing plant phylogenetic diversity, while saprotrophic fungal species richness was significantly affected by plant leaf nitrogen content, specific root length and standing biomass. The increasing ectomycorrhizal fungal richness associated with increasing plant phylogenetic diversity was driven by the combined presence of ectomycorrhizal fungal specialists in plots with both gymnosperm and angiosperm hosts. Although the species composition of both the ectomycorrhizal and saprotrophic fungal communities changed significantly in response to changes in plant species composition, the effect was much greater for ectomycorrhizal fungi. In addition, ectomycorrhizal but not saprotrophic fungal species composition was significantly influenced by both plant phylum (angiosperm, gymnosperm, both) and origin (Europe, America, both). The phylum effect was caused by differences in ectomycorrhizal fungal community composition, while the origin effect was attributable to differences in community heterogeneity. Taken together, this study emphasizes that plant-associated effects on soil fungal communities are largely guild-specific and provides a mechanistic basis for the positive link between plant phylogenetic diversity and ectomycorrhizal fungal richness.

International audience; Background and Aims: Despite a longstanding interest in variation in tree species vulnerability to ice storm damage, quantitative analyses of the influence of crown structure on within-crown variation in ice accretion are rare. In particular, the effect of prior interception by higher branches on lower branch accumulation remains unstudied. The aim of this study was to test the hypothesis that intra-crown ice accretion can be predicted by a measure of the degree of sheltering by neighbouring branches.Methods: Freezing rain was artificially applied to Acer platanoides L., and in situ branch-ice thickness was measured directly and from LiDAR point clouds. Two models of freezing rain interception were developed: ‘IceCube’, which uses point clouds to relate ice accretion to a voxel-based index (sheltering factor; SF) of the sheltering effect of branch elements above a measurement point; and ‘IceTree’, a simulation model for in silico evaluation of the interception pattern of freezing rain in virtual tree crowns.Key Results: Intra-crown radial ice accretion varied strongly, declining from the tips to the bases of branches and from the top to the base of the crown. SF for branches varied strongly within the crown, and differences among branches were consistent for a range of model parameters. Intra-crown variation in ice accretion on branches was related to SF (R2 = 0·46), with in silico results from IceTree supporting empirical relationships from IceCube.Conclusions: Empirical results and simulations confirmed a key role for crown architecture in determining intra-crown patterns of ice accretion. As suspected, the concentration of freezing rain droplets is attenuated by passage through the upper crown, and thus higher branches accumulate more ice than lower branches. This is the first step in developing a model that can provide a quantitative basis for investigating intra-crown and inter-specific variation in freezing rain damage.

Two main effects are proposed to explain biodiversity-ecosystem functioning relationships: niche complementarity and selection effects. Both can be functionally defined using the functional diversity (FD) and functional identity (FI) of the community respectively. Herein, we present results from the first tree diversity experiment that separated the effect of selection from that of complementarity by varying community composition in high-density plots along a gradient of FD, independent of species richness and testing for the effects of FD and community weighted means of traits (a proxy for FI) on stem biomass increment (a proxy for productivity). After 4 years of growth, most mixtures did not differ in productivity from the averages of their respective monocultures, but some did overyield significantly. Those positive diversity effects resulted mostly from selection effects, primarily driven by fast-growing deciduous species and associated traits. Net diversity effect did not increase with time over 4 years.

A climate warming of 2–5°C by the end of the century will impact the likelihood of seed germination of sugar maple (Acer saccharum), a dominant tree species which possesses a restricted temperature range to ensure successful reproduction. We hypothesize that seed origin affects germination due to the species' local adaptation to temperature. We tested this by experimentally investigating the effect of incubation temperature and temperature shifting on sugar maple seed germination from seven different seed sources representing the current species range. Survival analysis showed that seeds from the northern range had the highest germination percentage, while the southern range had the lowest. The mean germination percentage under constant temperatures was best when temperatures were ≤5°C, whereas germination percentages plummeted at temperatures ≥11°C (5.8%). Cool shifting increased germination by 19.1% over constant temperature treatments and by 29.3% over warm shifting treatments. Both shifting treatments caused earlier germination relative to the constant temperature treatments. A climate warming of up to +5°C is shown to severely reduce germination of seeds from the southern range. However, under a more pronounced warming of 7°C, seed germination at the northern range become more affected and now comparable to those found from the southern range. This study states that the high seed germination percentage found in sugar maple at the northern range makes it fairly resilient to the warmest projected temperature increase for the next century. These findings provide forest managers with the necessary information to make accurate projections when considering strategies for future regeneration while also considering climate warming.

Research on biodiversity and ecosystem functioning has demonstrated links between plant diversity and ecosystem functions such as productivity. At other trophic levels, the plant microbiome has been shown to influence host plant fitness and function, and host-associated microbes have been proposed to influence ecosystem function through their role in defining the extended phenotype of host organisms However, the importance of the plant microbiome for ecosystem function has not been quantified in the context of the known importance of plant diversity and traits. Here, using a tree biodiversity-ecosystem functioning experiment, we provide strong support for the hypothesis that leaf bacterial diversity is positively linked to ecosystem productivity, even after accounting for the role of plant diversity. Our results also show that host species identity, functional identity and functional diversity are the main determinants of leaf bacterial community structure and diversity. Our study provides evidence of a positive correlation between plant-associated microbial diversity and terrestrial ecosystem productivity, and a new mechanism by which models of biodiversity-ecosystem functioning relationships can be improved.

Tree seedlings planted in abandoned agricultural fields interact with herb communities through competition, tolerance, and facilitation. In addition, they are subject to herbivory by small mammals, deer or invertebrates. To increase the success of forest restoration in abandoned fields and reduce management costs, we should determine which species are tolerant to or facilitated by herbaceous vegetation and those which require protection from competition and predation. Eight native tree species were planted in plots covered by herbaceous vegetation, plots where herbaceous vegetation was removed, and plots where seedlings were surrounded by an organic mulch mat. Half of the seedlings were protected against small mammal damage. Results showed that two non-pioneer and moderately shade-tolerant species (yellow birch and red oak) were inhibited by herbaceous vegetation. Birch species were particularly affected by small mammal predation. No effects of predation or herbaceous competition were observed for conifer species. Rather, herbaceous vegetation had a positive effect on the survival and the height growth of tamarack (Larix laricina). None of the tested herb communities had a stronger competitive effect on tree growth than another. Restoration of abandoned fields using multi-tree species should be designed at the seedling scale rather than at the site scale to account for different tree responses to predation and competition as well as variable site conditions. An approach resembling precision agriculture is proposed to lower costs and any potential negative impact of more intensive vegetation management interventions.

Recent studies have reported positive net diversity effects on aboveground tree growth. However, whether similar effects occur belowground through root investment, and whether such effects are related to evergreenness of tree communities, is less clear. Here we studied vertical distribution of standing fine root biomass of twelve North American temperate tree species planted in a common garden tree diversity experiment of varying species richness and evergreenness to test whether belowground niche complementarity of trees could explain positive diversity effects reported aboveground. We tested two alternative hypotheses: trees in mixtures increase uptake of soil resources (1) by increasing vertical root stratification and/or producing a greater fine root density (mg cm−3) or (2) by producing similar or fewer fine roots that are potentially more efficient. Additionally, we hypothesized that proportional allocation to belowground biomass increases with evergreenness of tree communities. Fine roots were sampled in six layers of 5–10 cm, from 0 to 40 cm depth in single-, two- and four-species mixtures. We did not observe an effect of species richness on rooting depth or root density, refuting the hypothesis that aboveground overyielding in tree mixtures is linked to fine root overyielding. Rather, we observed a significant negative diversity effect (− 7.6%) on total fine root density, suggesting overall less investment to fine roots with increasing diversity. The strong positive effect of evergreeness on proportional allocation to fine roots over aboveground parts suggests that deciduous tree roots may be generally more efficient at absorbing soil resources, at least in the early years after tree establishment.

15 pages, 6 figures, 3 tables, supplementary data https://doi.org/10.1093/femsec/fiy115, Freshwater bacterioplankton communities are influenced by the inputs of material and bacteria from the surrounding landscape, yet few studies have investigated how different terrestrial inputs affect bacterioplankton. We examined whether the addition of soils collected under various tree species combinations differentially influences lake bacterial communities. Lake water was incubated for 6 days following addition of five different soils. We assessed the taxonomic composition (16S rRNA gene sequencing) and metabolic activity (Biolog Ecoplates) of lake bacteria with and without soil addition, and compared these to initial soil communities. Soil bacterial assemblages showed a strong influence of tree composition, but such community differences were not reflected in the structure of lake communities that developed during the experiment. Bacterial taxa showing the largest abundance increases during incubation were initially present in both lake water and across most soils, and were related to Cytophagales, Burkholderiales and Rhizobiales. No clear metabolic profiles based on inoculum source were found, yet soil-amended communities used 60% more substrate than non-inoculated communities. Overall, we show that terrestrial inputs influence aquatic communities by stimulating the growth and activity of certain ubiquitous taxa distributed across the terrestrial-aquatic continuum, yet different forest soils did not cause predictable changes in lake bacterioplankton assemblages, This work was supported by the Natural Science and Engineering Research Council of Canada (NSERC) and Hydro-Quebec, as well as by the Fons de Reserche Nature et Technologies (FRQNT) and the Canada Research Chairs program

International audience; Despite considerable research demonstrating that biodiversity increases productivity in forests and regulates herbivory and pathogen damage, there remain gaps in our understanding of the shape, magnitude, and generality of these biodiversity-ecosystem functioning (BEF) relationships. Here, we review findings from TreeDivNet, a global network of 25 tree diversity experiments, on relationships between levels of biodiversity and (a) tree growth and survival and (b) damage to trees from pests and pathogens. Tree diversity often improved the survival and above- and belowground growth of young trees. The mechanistic bases of the diversity effects on tree growth and survival include both selection effects (i.e., an increasing impact of particular species in more species-rich communities) and complementary effects (e.g. related to resource differentiation and facilitation). Plant traits and abiotic stressors may mediate these relationships. Studies of the responses of invertebrate and vertebrate herbivory and pathogen damage have demonstrated that trees in more diverse experimental plots may experience more, less, or similar damage compared to conspecific trees in less diverse plots. Documented mechanisms producing these patterns include changes in concentration, frequency, and apparency of hosts; herbivore and pathogen diet breadth; the spatial scale of interactions; and herbivore and pathogen regulation by natural enemies. Our review of findings from TreeDivNet indicates that tree diversity experiments are extending BEF research across systems and scales, complementing previous BEF work in grasslands by providing opportunities to use remote sensing and spectral approaches to study BEF dynamics, integrate belowground and aboveground approaches, and trace the consequences of tree physiology for ecosystem functioning. This extension of BEF research into tree-dominated systems is improving ecologists’ capacity to understand the mechanistic bases behind BEF relationships. Tree diversity experiments also present opportunities for novel research. Since experimental tree diversity plantations enable measurements at tree, neighbourhood and plot level, they allow for explicit consideration of temporal and spatial scales in BEF dynamics. Presently, most TreeDivNet experiments have run for less than ten years. Given the longevity of trees, exciting results on BEF relationships are expected in the future.

Understanding tree recruitment dynamics in various growth environments is essential for a better assessment of tree species’ adaptive capacity to climate change. We investigated the microsite factors influencing survival, growth, and foliar nutrition of natural and planted sugar maple seedlings (Acer saccharum) along a gradient of tree species that reflect the change in composition from temperate hardwoods to boreal forests of eastern Canada. We specifically tested whether the increasing abundance of conifers in the forest and its modifications on soil properties negatively affects foliar nutrition of natural seedlings as well as the survival and growth of seedlings planted directly in the natural soil and in pots filled with enriched soil. Results of natural seedlings indicate that under conifer‐dominated stands, lower soil pH, accelerated dissolution of some minerals, lower temperature and moisture, and higher levels of phenolic compounds have created microsites that are less suitable for sugar maple foliar nutrition and regeneration. These conditions were omnipresent under hemlock. The growth of seedlings planted in the natural soil was negatively impacted by the overall low soil quality under all forest types (as compared to seedlings planted in pots with enriched soil). However, survival and growth of the seedlings were not negatively affected by conifers, regardless of planting type, likely because of stored nutrients from the nursery. Also, lower survival was found under maple–birch stands for seedlings planted both in the natural soil and in pots with enriched soil due to higher shading. This study has identified key microsite factors created by specific conifers that may impede or benefit the potential of sugar maple to maintain its current range or expand its range northward under climate change.

In natural forests, tree leaf surfaces host diverse bacterial communities whose structure and composition are primarily driven by host species identity. Tree leaf bacterial diversity has also been shown to influence tree community productivity, a key function of terrestrial ecosystems. However, most urban microbiome studies have focused on the built environment, improving our understanding of indoor microbial communities but leaving much to be understood, especially in the nonbuilt microbiome. Here, we provide the first multiple-species comparison of tree phyllosphere bacterial structures and diversity along a gradient of urban intensity. We demonstrate that urban trees possess characteristic bacterial communities that differ from those seen with trees in nonurban environments, with microbial community structure on trees influenced by host species identity but also by the gradient of urban intensity and by the degree of isolation from other trees. Our results suggest that feedback between human activity and plant microbiomes could shape urban microbiomes., Tree leaf-associated microbiota have been studied in natural ecosystems but less so in urban settings, where anthropogenic pressures on trees could impact microbial communities and modify their interaction with their hosts. Additionally, trees act as vectors spreading bacterial cells in the air in urban environments due to the density of microbial cells on aerial plant surfaces. Characterizing tree leaf bacterial communities along an urban gradient is thus key to understand the impact of anthropogenic pressures on urban tree-bacterium interactions and on the overall urban microbiome. In this study, we aimed (i) to characterize phyllosphere bacterial communities of seven tree species in urban environments and (ii) to describe the changes in tree phyllosphere bacterial community structure and diversity along a gradient of increasing urban intensity and at two degrees of tree isolation. Our results indicate that, as anthropogenic pressures increase, urban leaf bacterial communities show a reduction in the abundance of the dominant class in the natural plant microbiome, the Alphaproteobacteria. Our work in the urban environment here reveals that the structures of leaf bacterial communities differ along the gradient of urban intensity. The diversity of phyllosphere microbial communities increases at higher urban intensity, also displaying a greater number and variety of associated indicator taxa than the low and medium urban gradient sites. In conclusion, we find that urban environments influence tree bacterial community composition, and our results suggest that feedback between human activity and plant microbiomes could shape urban microbiomes. IMPORTANCE In natural forests, tree leaf surfaces host diverse bacterial communities whose structure and composition are primarily driven by host species identity. Tree leaf bacterial diversity has also been shown to influence tree community productivity, a key function of terrestrial ecosystems. However, most urban microbiome studies have focused on the built environment, improving our understanding of indoor microbial communities but leaving much to be understood, especially in the nonbuilt microbiome. Here, we provide the first multiple-species comparison of tree phyllosphere bacterial structures and diversity along a gradient of urban intensity. We demonstrate that urban trees possess characteristic bacterial communities that differ from those seen with trees in nonurban environments, with microbial community structure on trees influenced by host species identity but also by the gradient of urban intensity and by the degree of isolation from other trees. Our results suggest that feedback between human activity and plant microbiomes could shape urban microbiomes.

Sugar maple (SM, Acer saccharum Marsh.) dominated forests of North America are increasingly affected by many human-induced modifications in environmental conditions. As a remedy, adapted silvicultural treatments are needed. Even though it is generally accepted that SM health is related to soil fertility and even though there is extensive literature on SM – American beech (AB, Fagus grandifolia Ehrh.) regeneration stand dynamics related to light availability, the interaction between these two factors has rarely been studied. Our main objective was thus to verify the possible role of a light–soil interaction on SM–AB stand dynamics. We used a factorial design with three factors (harvest intensity, liming, and AB sapling elimination) to test this interaction. Our results showed that the radial growth of SM and AB tree and sapling growth was positively affected by canopy opening but not by liming. Liming did not favour AB seedlings, whereas it favoured SM seedlings in specific canopy opening situations, confirming, albeit partially, the light–soil interaction hypothesis. Overall, liming had very limited effects on SM–AB stand dynamics compared with canopy opening and AB sapling elimination treatments. We do not advocate the extensive use of liming, as other silvicultural strategies tested provided more promising results to favour SM over AB.

Studies have reported divergent results on the effect of soil fertility and canopy opening on understory density and growth of sugar maple (AS; Acer saccharum Marsh.) and American beech (FG; Fagus grandifolia Ehrh.). The main objective of this study was to evaluate the effect of a gradient of canopy opening and soil fertility on the density and growth of AS and FG saplings in southwestern Quebec, Canada. We investigated 56 stands containing both AS and FG that were subjected to different disturbance history types (DHTs) (UF, unmanaged forest; PC, partial cut; and CC, clearcut) on various soil types. AS and FG absolute and relative sapling density varied greatly among the 56 stands; however, no significant effects of DHT, soil nutrient availability, or their interaction were found. Both species responded positively in terms of radial growth to canopy openings, with FG growth being slightly better than AS growth in PC stands compared with other canopy treatments. Contrary to our hypothesis, AS did not show significantly higher growth than FG following clear-cutting. These results do not support the idea that AS abundance and growth could be promoted by increasing the intensity of the canopy opening during harvest, at least on the generally acidic and base-poor soils that were investigated.

The allocation of non-structural carbohydrates (NSCs) to reserves constitutes an important physiological mechanism associated with tree growth and survival. However, procedures for measuring NSC in plant tissue are expensive and time-consuming. Near-infrared spectroscopy (NIRS) is a high-throughput technology that has the potential to infer the concentration of organic constituents for a large number of samples in a rapid and inexpensive way based on empirical calibrations with chemical analysis. The main objectives of this study were (i) to develop a general NSC concentration calibration that integrates various forms of variation such as tree species and tissue types and (ii) to identify characteristic spectral regions associated with NSC molecules. In total, 180 samples from different tree organs (root, stem, branch, leaf) belonging to 73 tree species from tropical and temperate biomes were analysed. Statistical relationships between NSC concentration and NIRS spectra were assessed using partial least squares regression (PLSR) and a variable selection procedure (competitive adaptive reweighted sampling, CARS), in order to identify key wavelengths. Parsimonious and accurate calibration models were obtained for total NSC (r2 of 0·91, RMSE of 1·34% in external validation), followed by starch (r2 = 0·85 and RMSE = 1·20%) and sugars (r2 = 0·82 and RMSE = 1·10%). Key wavelengths coincided among these models and were mainly located in the 1740-1800, 2100-2300 and 2410-2490 nm spectral regions. This study demonstrates the ability of general calibration model to infer NSC concentrations across species and tissue types in a rapid and cost-effective way. The estimation of NSC in plants using NIRS therefore serves as a tool for functional biodiversity research, in particular for the study of the growth-survival trade-off and its implications in response to changing environmental conditions, including growth limitation and mortality. © 2015 British Ecological Society.

Given evidences that diverse ecosystems provide more services than depauperate ones, much attention has now turned toward finding meaningful and operational diversity indices. We ask two questions: (1) Does phylogenetic diversity contain additional information not explained by functional traits? And (2) What are the strength and nature of the correlation between phylogeny and functional traits according to the evolutionary scale considered? We used data from permanent forest plots of northeastern Canada for which these links have been demonstrated and important functional traits identified. We show that the nature of the relationship between traits and phylogeny varies dramatically among traits, but also according to the evolutionary distance considered. The demonstration that different characters show phylogenetic autocorrelation at different evolutionary depths suggests that phylogenetic content of traits may be too crude to determine whether phylogenies contain relevant information. However, our study provides support for the use of phylogenies to assess ecosystem functioning when key functional traits are unavailable. We also highlight a potentially important contribution of phylogenetics for conservation and the study of the impact of biodiversity loss on ecosystem functioning and the provision of services, given the accumulating evidence that mechanisms promoting diversity effects shift over time to involve different traits.

Summary The ability of soil microbial communities to withstand and recover from disturbance or stress is important for the functional stability of forest ecosystems. However, the relationship between the community responses of soil microbes and variation in tree mixtures vs functional composition remains poorly understood. We investigated soil biochemical properties and soil microbial resistance and resilience to drought in three 4-year-old tree monocultures (Acer saccharum Marsh, Larix laricina (Duroi) K. Koch and Pinus strobus L.) and two tree species combinations (L. laricina/A. saccharum and L. laricina/P. strobus) planted in a high-density tree field experiment located in southern Quebec, Canada. The experimentally imposed drought stress consisted of maintaining soil material for 30 days at 25% of water-holding capacity (WHC). Microbial biomass was assessed immediately after the water stress (resistance) and 15 and 30 days following drought (resilience). Results showed that tree communities influenced soil chemistry, soil respirometry properties and microbial resistance and resilience. We measured significant non-additive (i.e. both synergistic and antagonistic) effects of mixing tree species in some of the soil biochemical properties measured, mostly in the L. laricina/A. saccharum mixture. However, we did not find non-additive effects of tree mixtures on microbial resistance and resilience. A structural equation modelling analysis revealed that resistance and resilience were mostly modulated by direct effects of community-weighted means (CWM) of leaf litter lignin content and mineralizable N, and by indirect links from tree density and CWM of leaf litter N content via mineralizable N. This study suggests that tree species identity surpassed species mixtures as a key driver of soil microbial resistance and resilience. We showed a trade-off between microbial resistance and resilience in soil food webs, which is consistent with ecological theory. Our results indicate that differences in functional traits between tree species may rapidly be reflected in divergent soil biochemical properties and that these differences can in turn drive soil microbial resistance and resilience to drought.

Several recent studies have reported a marked increase in American beech dominance (Fagus grandifolia Ehrh.) relative to sugar maple (Acer saccharum Marsh.) in late successional forests of North America. However, many factors have been proposed to explain this sudden shift in tree species composition. We investigated the microsite factors responsible for maple regeneration failure under maple-beech stands, focusing on both light availability and soil conditions. The survival and growth of maple seedlings planted in the natural soil and in pots with enriched soil were monitored for two years, as well as foliar nutrition and herbivory damages of natural seedlings. The results indicate that low light availability associated with the presence of beech is the primary factor leading to maple regeneration failures. Soil nutrient availability and foliar nutrition of natural seedlings did not differ between forest types. Yet, the results indicate that factors such as allelopathy and preferential herbivory on maple seedlings under beech could be superimposed effects that hinder maple regeneration. Under similar forests, a control of beech sapling abundance in the understory followed by selection cutting could be one way to promote and maintain maple populations in the longer term.

BACKGROUND: Interest in tree form assessments using the terrestrial laser scanner (TLS) has increased in recent years. Yet many existing methods are limited to small-sized trees, principally due to noise and occlusion phenomena. In this paper, a novel voxel-based program that is dedicated to the analyses of large tree structures is presented. The method is based on the assumption that architectural trait variations (i.e. branching angle, bifurcation ratio, biomass allocation, etc.) influence the way a tree explores space. This method uses the concept of space exploration that considers a voxel as a portion of space explored by the tree. Once the TLS scene is voxelized, the program provides tools that extract qualitative (geometrical) and quantitative (volumetric) metrics. These tools measure (1) voxel dispersion in three dimensions (3-D), (2) projections of the voxel cloud in 2-D and (3) multi-temporal changes within a single tree crown. SCOPE: To test algorithm capabilities of measuring larger tree architectural traits, two application studies were conducted using point clouds that were either generated by a tree growth simulation model, thereby allowing algorithm application in a perfectly controlled environment, or acquired in the field with a TLS device. The space exploration concept makes it possible to take advantage of the volumetric nature of voxels to compensate for occlusion. The hypothesis that large-sized voxels can be used to reduce occlusion in the original point cloud was tested, as well as the consequences of voxel size on quantification of tree volume and on precision of derived metrics. CONCLUSIONS: Results show that space exploration is well adapted to highlight architectural differences among trees. They also suggest that large-sized voxels are efficient for occlusion compensation at the expense of metrics precision in some cases. The best resolution to choose depending on the research objectives and quality of the TLS scan is discussed.

Despite being instrumental in forest ecology, the definition and nature of shade tolerance are complex and not beyond controversy. Moreover, the role it plays in the trait-demography relationship remains unclear. Here, we hypothesize that shade tolerance can be achieved by alternative combinations of traits depending on the species' functional group (evergreen vs. deciduous species) and that its ability to explain the array of traits involved in demography will also vary between these two groups. We used dimension reduction to identify the main trait spectra for 48 tree species, including 23 evergreens and 25 deciduous - dispersed across 21 genera and 13 families. We assessed the relationship between functional traits, shade tolerance, and demographic performance at high and low light using structural equation modelling. The dimensions found corresponded to the trait spectra previously observed in the literature and were significantly related to measures of demography. However, our results support the existence of a divergence between evergreen and deciduous species in the way shade tolerance relates to the demography of species along light gradients. We show that shade tolerance can be attained through different combination of traits depending on the functional and geographical context, and thus, its utilization as a predictor of forest dynamics and species coexistence requires previous knowledge on the role it plays in the demographic performance of the species under study. The authors acknowledge Dr. Yoshida and Dr. Evans for kindly providing the SORTIE parameters for species from Japan and United Kingdom, respectively. We are also grateful to Timothy Paine and two anonymous referees for their constructive comments on an earlier version of this manuscript. This research was supported by a Juan de la Cierva Fellowship to A.A. (FJCI‐2014‐20739), a NSERC Strategic Grant to D.G. and the NSERC industrial Chair programme to C.M. Additional support came from the CERCA Programme of the Generalitat de Catalunya. The authors declare no conflict of interests.

Use of fast-growing tree plantations on dedicated areas is proposed as a means of reconciling fibre production with conservation objectives. Success of this approach however requires fine-tuning silvicultural scenarios so that survival and growth are optimized while management and environmental costs are minimized. This is particularly challenging for hybrid larch ( Ã Coaz), a shade-intolerant species planted on fertile sites in Quebec (Canada) where legislation prevents the use of chemical herbicides. In this context, multiple motor-manual release treatments are often required, with high impacts on costs and social issues related to the scarcity of a qualified workforce. We established a split-split-plot design on a recently harvested site to assess the main and interaction effects of mechanical site preparation (MSP) intensity (five modalities of trenching or mounding), motor-manual release scenario (one or two treatments) and planting depth (0â3 cm or 3â10 cm) on hybrid larch seedling growth and survival six years after planting. Mechanical site preparation intensity and planting depth did not influence seedling growth after 6 years. The lack of significant interaction between MSP and release scenarios indicates that these operations should be planned independently. A more intensive MSP treatment cannot replace a second motor-manual release on fertile sites, as proposed to reduce costs. Our results also show the significant advantage of performing two motor-manual release treatments two years apart (the first one early in the scenario), over performing a single treatment. Our study provides silvicultural guidelines for the establishment of high-yield exotic larch plantations.Larixmarschlinsii

Some site preparation is generally recommended to enhance the growth and survival of planted and naturally regenerated seedlings, but it must be justified both economically and environmentally. More severe preparation is thought to be necessary for intensive plantation silviculture, e.g., using fast-growing, ameliorated stocks, especially in boreal ecosystems. Although not justified scientifically, deep-planting of seedlings is often discouraged and may even be financially penalized in eastern Canada. We thus evaluated early seedling growth and survival of hybrid larch (Larix × marschlinsii Coaz) in an experiment including mechanical site preparation and planting depth treatments. Our results suggest that satisfactory early hybrid larch establishment and growth could be met using low environmental impact or low cost treatments (such as soil inversion using an excavator or single-pass disk trenching), and that deeper planting has no negative effect. Structural equation modelling (SEM) was used to explore causal relationships between factors influencing seedling performance at the local scale (planting microsites), including soil moisture, soil temperature, surrounding vegetation, and seedling nutrition. SEM analysis supported the absence of overall differences among treatments, while also highlighting the negative impact of increased soil water content where drainage was suboptimal, as well as the unexpected positive impact of increased competition on growth mostly through seedling nutrition, among others. These early observations will need to be confirmed over a longer period, as well as with a more comprehensive assessment of site environmental conditions and competition intensity.

We analysed the spatial distribution of fine roots and light availability in a tree-based intercrop system (TBI) composed of Quercus rubra L. (QUR), hybrid poplar (Populus deltoides × Populus nigra—HYP) and hay (CROP) in southern Quebec (Canada) to evaluate interactions between trees and crop. Trees in the 8-year-old TBI system had superficial fine root profiles, which is common in tree species grown in conventional plantations and natural forests. More than 95 % of fine roots were found within the first 25 and 45 cm for QUR and HYP, respectively, and 35 cm for CROP. However, vertical separation between the fine root systems of QUR and CROP was evident, as QUR allocated less fine roots to the top 10 cm of soil, and more to depths between 10 and 30 cm, as opposed to CROP which had a greater proportion of fine roots in the top 10 cm. HYP fine roots showed no adaption when intercropped with hay. High tree fine root length density (FRLD) in the top soil layer was observed near the tree stems while hay FRLD was reduced by 45 %, suggesting strong competition for resources. Hay yield analysis revealed significant reduction near trees, particularly HYP. However, light did seem to be the main driver of intercrop yield, as it not only accounted for the effect of competition by roots (being correlated), but also had a singular effect.

Partial harvesting during the snow-free season disturbs the forest floor and modifies seedbed characteristics. Quantitative information is lacking about the distribution of changes in seedbed proportions between areas located in and outside skid trails, and about how long these changes persist after harvest. These effects could interact with species’ seedbed requirements and seed input and influence spatio-temporal patterns of seedling establishment in forests, which would have important implications for regeneration dynamics. The objectives of this study were to determine how selection cutting affects seedbed proportions in and outside skid trails in northern hardwood stands, how these seedbed proportions vary over time following harvest, and how seedbed proportions in selection cuts compare with unharvested stand conditions. We sampled 12 sugar maple-dominated stands in southeastern Quebec, Canada. Two had not been harvested in the recent decades, while 10 had been harvested through selection cutting, 1–3 years earlier. A total of 3600 quadrats were sampled to determine the proportions of 8 seedbed types and whether or not a quadrat was located in a skid trail. This was the case for 24.1% of the quadrats in selection cuts. Outside skid trails, seedbed proportions in selection cuts did not vary from those in unharvested stands (P ⩾ 0.097). In these stands, leaf litter was the most abundant substrate, covering 87.3% of the forest floor, followed by rotten wood (4.9%) and fresh wood (3.0%). Humus, rocks and live tree bases occupied 1–2% the forest floor, while mineral soil and moss covered less than 1%. In selection cuts, proportions of rotten wood and live tree bases were lower in skid trails than outside, and this difference persisted 13 years after harvest. In 1- and 2-year-old cuts, the proportion of litter was lower in skid trails than outside, but not later on. The proportions of mineral soil and disturbed humus increased sharply in the skid trails after harvest (17.1-fold and 2.7-fold increases, respectively), but these effects lasted only 3 years for mineral soil and 1 year for humus. Power functions were used to model variation in litter, mineral soil and disturbed humus proportions as a function of time since harvest. We discuss the implications for regeneration dynamics of the marked, but short term increase in mineral soil and disturbed humus availability. Spatially explicit models that can simulate the mid- to long-term availability of substrates in both time and space would be useful for assessing the long-term implications of harvesting on seedbed proportions and regeneration patterns.

While post-harvest residue (“slash”) is increasingly viewed as a source of biofuel, few studies have considered the potential ecological impact of the spatial distribution of forest harvest residue. We hypothesized that slash piles create islands of high soil fertility and light, leading to greater abundance, growth, and reproduction of plants. In 6-yr-old intensively managed hybrid poplar (Populus balsamifera L. × P. maximowiczii A. Henry) plantations, we showed that soluble organic N, NO₃–N, NH₄–N, and P decreased as a negative exponential function with distance from the pile. Pin cherry (Prunus pensylvanica L. f.) relative growth rate was fastest near piles in the first few years after harvest. We found significantly greater stem size, flowers per plant, and foliar P near piles for cherry, and this inverse gradient sharpened in a 16-yr-old white spruce [Picea glauca (Moench) Voss] plantation. For the two other flowering species, strawberry (Fragaria virginiana Mill.) and raspberry (Rubus idaeus L.), flower and fruit abundance were also significantly and strongly negatively correlated with distance to the pile. Further, directly correlating soil nutrient availability with plant traits, we found significant positive relationships between plant growth, reproductive output, and N availability. Partial correlation analysis indicated that more of the variance in plant traits was explained by distance than by soil nutrition. We conclude that in industrial forests, piles replace canopy gaps as sites where understory plant species can episodically reproduce and are therefore important for many plant species as well as the pollinators and frugivores dependent on them.

We report a new silvicultural approach that is well suited for the management of uneven-aged forests in which timber production is an important objective. The approach recognizes two main components in the stand, i.e. a fiber production component, which provides veneer/sawlog quality products from the high-quality trees (HQT), and an ecological component, which contributes to the overall ecosystem functioning through the lower value stems. The objective of the study was to verify if it is possible to sustainably harvest only HQT in northern hardwood (NH) and therebyproduce aviable alternative to high-grading the stands. To do so, a simple stand growthsimulator, based on empirical growth rates of HQT in Sugar Maple/Yellow Birch stands in southwestern Quebec, was combined with an optimization tool. The optimization parameters aimed to identify possible tree marking regimes (TMRs) under 10-year rotation partial cutting, which would ensure that the basal area of HQT was maintained for 40 years. Results suggest that sustainability is achievable starting from very different initial stand structures and the application of a wide range of alternative TMRs. We argue that this new approach is one way to apply emerging concepts in forest management, such as ecological integrity, attempts to emulate natural disturbance regimes and provides new possibilities managing for resilience and for adaptation to climate change.

Partial cutting has been proposed as a means to better conserve biodiversity in managed forest landscapes. However, partial cutting encompasses many forms of silviculture; some with implicit goals of maintaining biodiversity such as multicohort harvesting or others which may specifically focus on regeneration of stands but may still provide some additional benefits for biodiversity such as shelterwood harvesting. Here we compared ground beetle assemblages of clear cuts, shelterwoods, multicohort harvested stands and uncut stands collected using pitfall traps both 2 and 3-years post-harvest. We hypothesized that partial cutting treatments would maintain assemblages that were more similar to uncut stands than to clear cuts. We further hypothesized that among partial cuts the multicohort harvested stands, with relatively high levels of retention (66%), would maintain beetle assemblages that were more similar to uncut stands than would shelterwoods, which had lower levels of retention (50%). We collected 6692 individuals, representing 42 species. Catch rates of beetles were similar among all harvested treatments (shelterwood, multicohort and clear cuts) and lower than uncut stands. Species richness and composition was similar between shelterwood and multicohort stands. Both partial cut treatments fell between clear cuts and uncut stands in terms of species richness and compositional similarity. Compositional differences between uncut stands and partial cut stands were defined primarily by reduced abundances of forest associated species such as Agonum retractum (LeConte), Synuchus impunctatus (Say) and four Pterostichus species within partial cuts. Within partial cuts, beetle assemblages differed between machine corridors with 0% retention and adjacent partial cut strips (50% retention) and uncut vegetation corridors (100%). We conclude that both shelterwoods and multicohort harvesting stands provide at least initially similar benefits for biodiversity compared to clear cutting although neither maintains assemblages consistent with those found in uncut stands. We expect that these similarities will end once trees are removed from shelterwoods. The reductions in abundances within partial cuts may extend the time necessary for individual populations to increase to pre-harvest levels in partial cuts. For land-managers, similar initial responses of beetle assemblages in multicohort and shelterwood harvests may permit some flexibility for conservation planning whereby final removal of seed trees within shelterwoods could be delayed depending on the status of recovering beetle populations.

Direct tree seeding is potentially an economical technique for restoring forests on abandoned fields. However, the success of tree establishment depends on many factors related to species and seed characteristics, environmental conditions, competition and predation. We compared seedling emergence, survival and growth of six tree species of different seed sizes in a forest restoration project of abandoned fields. Species were seeded in plots with and without herbaceous vegetation and with and without protection from bird and mammal predation. Yellow birch (Betula alleghaniensis) did not emerge in all treatments, paper birch (Betula papyrifera) and tamarack (Larix laricina) had a seedling emergence rate lower than 1%, and sugar maple (Acer saccharum) had a low overall emergence rate of 6%. Seedling emergence reached 57% for northern red oak (Quercus rubra) and 34% for red pine (Pinus resinosa), but survival of oak after one year was much higher (92%) than pine seedlings (16%). Overall, protection from birds and mammals and elimination of the herbaceous vegetation cover had no detectable effects on seedling emergence, survival and height. Nonetheless, red oak seedlings growing in the presence of vegetation had a smaller diameter and shoot biomass and a larger specific leaf area. We conclude that only large seeded species, such as oak, should be used for forest restoration of abandoned fields by direct seeding in our region.