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Deciphering the mechanisms that link biodiversity with ecosystem functions is critical to understanding the consequences of changes in biodiversity. The hypothesis that complementarity and selection effects drive relationships between biodiversity and ecosystem functions is well accepted, and an approach to statistically untangle the relative importance of these effects has been widely applied. In contrast, empirical demonstrations of the biological mechanisms that underlie these relationships remain rare. Here, on the basis of a field experiment with young trees, we provide evidence that one form of complementarity in plant communities-complementarity among crowns in canopy space-is a mechanism, related to light interception and use, that links biodiversity with ecosystem productivity. Stem biomass overyielding increased sharply in mixtures with greater crown complementarity. Inherent differences among species in crown architecture led to greater crown complementarity in functionally diverse species mixtures. Intraspecific variation, specifically neighbourhood-driven plasticity in crowns, further modified spatial complementarity and strengthened the positive relationship with overyielding-crown plasticity and inherent interspecific differences contributed near equally in explaining patterns of overyielding. We posit that crown complementarity is an important mechanism that may contribute to diversity-enhanced productivity in forests.

BackgroundThe diversity and composition of the microbial community of tree leaves (the phyllosphere) varies among trees and host species and along spatial, temporal, and environmental gradients. Phyllosphere community variation within the canopy of an individual tree exists but the importance of this variation relative to among-tree and among-species variation is poorly understood. Sampling techniques employed for phyllosphere studies include picking leaves from one canopy location to mixing randomly selected leaves from throughout the canopy. In this context, our goal was to characterize the relative importance of intra-individual variation in phyllosphere communities across multiple species, and compare this variation to inter-individual and interspecific variation of phyllosphere epiphytic bacterial communities in a natural temperate forest in Quebec, Canada.MethodsWe targeted five dominant temperate forest tree species including angiosperms and gymnosperms:Acer saccharum,Acer rubrum,Betula papyrifera,Abies balsameaandPicea glauca. For one randomly selected tree of each species, we sampled microbial communities at six distinct canopy locations: bottom-canopy (1–2 m height), the four cardinal points of mid-canopy (2–4 m height), and the top-canopy (4–6 m height). We also collected bottom-canopy leaves from five additional trees from each species.ResultsBased on an analysis of bacterial community structure measured via Illumina sequencing of the bacterial 16S gene, we demonstrate that 65% of the intra-individual variation in leaf bacterial community structure could be attributed to the effect of inter-individual and inter-specific differences while the effect of canopy location was not significant. In comparison, host species identity explains 47% of inter-individual and inter-specific variation in leaf bacterial community structure followed by individual identity (32%) and canopy location (6%).DiscussionOur results suggest that individual samples from consistent positions within the tree canopy from multiple individuals per species can be used to accurately quantify variation in phyllosphere bacterial community structure. However, the considerable amount of intra-individual variation within a tree canopy ask for a better understanding of how changes in leaf characteristics and local abiotic conditions drive spatial variation in the phyllosphere microbiome.

International audience; The impact of mechanical site preparation (MSP) on soil biogeochemical structure in young larch plantations was investigated. Soil samples were collected in replicated plots comprising simple trenching, double trenching, mounding and inverting site preparation. Unlogged natural mixed forest areas were used as a reference. Analysis of soil nutrients, abundance of bacteria and gas exchanges unveiled no significant difference among the plots. However, inverting site preparation resulted in higher variations of gas exchanges when compared with trenching, mounding and unlogged natural forest. A combination of the biological and physicochemical variables was used to define a multifunctional classification of the soil samples into four distinct groups categorized as a function of their deviation from baseline ecological conditions. According to this classification model, simple trenching was the approach that represented the lowest ecological risk potential at the microsite level. No relationship was observed between MSP method and soil bacterial community structure as assessed by high-throughput sequencing of bacterial 16S rRNA gene; however, indicator genotypes were identified for each multifunctional soil class. This is the first identification of multifunctional molecular indicators for baseline and disturbed ecological conditions in soil, demonstrating the potential of applied microbial ecology to guide silvicultural practices and ecological risk assessment.

The area of forest plantations is increasing worldwide helping to meet timber demand and protect natural forests. However, with global change, monospecific plantations are increasingly vulnerable to abiotic and biotic disturbances. As an adaption measure we need to move to plantations that are more diverse in genotypes, species, and structure, with a design underpinned by science. TreeDivNet, a global network of tree diversity experiments, responds to this need by assessing the advantages and disadvantages of mixed species plantations. The network currently consists of 18 experiments, distributed over 36 sites and five ecoregions. With plantations 1-15 years old, TreeDivNet can already provide relevant data for forest policy and management. In this paper, we highlight some early results on the carbon sequestration and pest resistance potential of more diverse plantations. Finally, suggestions are made for new, innovative experiments in understudied regions to complement the existing network. ispartof: Ambio vol:45 issue:1 pages:29-41 ispartof: location:Sweden status: published

Small-seeded tree species such as yellow birch (YB, Betula alleghaniensis Britt.) require deadwood or mineral soil for their establishment. Although much research has been done comparing YB germination on leaf litter vs. exposed mineral soil, less is known about deadwood as a seedbed and how different seedbeds affect YB early growth along light availability and size gradients. We examine how three common seedbeds (deadwood, moss cover on deadwood, and mineral soil) affected establishment and growth, biomass partitioning, and morphological traits of YB juveniles growing in the understory of temperate mixed deciduous and coniferous forests in southern Quebec. A total of 274 YB were sampled in four sugar maple (Acer saccharum Marsh.) dominated northern hardwood stands where selective cuts had been applied 6 and 15 years prior to sampling. Over 75% of the YB found on deadwood were on material of birch and conifer origin, although these species made less than 40% of the basal area. YB juveniles growing on deadwood showed traits that improve survival in shade such as reduced height growth for tall plants, higher efficiency in resource capture, and multilayered crowns. Our results demonstrate the importance of deadwood of birch and conifer origin in maintaining an abundant, natural, spatially well-distributed, and multistoried regeneration of YB. The authors are particularly grateful to I. Pratte, D. Schönig, A. Paquin, N. Bergeron, M. Messier, E. Messier, S. Carpentier, Y. Gauthier, and E. Mauri for their valuable help with the field and lab work. They also thank M. Mazerolle and S. Daigle for their assistance with statistical models. This work was funded by a MVRMF grant from the Ministry of Natural Resources of Quebec (MRNQ). J.-B. Lambert was also awarded with a Fonds de recherche du Québec — Nature et technologies (FRQNT) scholarship.

Isabelle Aubin [et al.].- Received October 15, 2015, Published on the web 23 February 2016. ., The integration of functional traits into vulnerability assessments is a promising approach to quantitatively capture differences in species sensitivity and adaptive capacity to climate change, allowing the refinement of tree species distribution models. In response to a clear need to identify traits that are responsive to climate change and applicable in a management context, we review the state of knowledge of the main mechanisms, and their associated traits, that underpin the ability of boreal and temperate tree species to persist and/or shift their distribution in a changing climate. We aimed to determine whether current knowledge is sufficiently mature and available to be used effectively in vulnerability assessments. Marshalling recent conceptual advances and assessing data availability, our ultimate objective is to guide modellers and practitioners in finding and selecting sets of traits that can be used to capture differences in species’ ability to persist and migrate. While physiological mechanisms that determine sensitivity to climate change are relatively well understood (e.g. drought-induced cavitation), many associated traits have not been systematically documented for North American trees and differences in methodology preclude their widespread integration into vulnerability assessments (e.g. xylem recovery capacity). In contrast, traits traditionally associated with the ability to migrate and withstand fire are generally well documented, but new key traits are emerging in the context of climate change that have not been as well characterized (e.g. age of optimum seed production). More generally, the extent of, and patterns in, intraspecific trait variation, as well as co-variation and interaction among traits, limit our ability to use this approach to assess tree adaptive capacity. We conclude by outlining research needs and potential strategies for the development of trait-based knowledge applicable in large-scale modelling efforts, sketching out important aspects of trait data organization that should be part of a coordinated effort by the forest science community.

While trees store substantial amounts of nonstructural carbon (NSC) for later use, storage regulation and mobilization of stored NSC in long-lived organisms like trees are still not well understood. At two different sites with sugar maple (Acer saccharum), we investigated ascending sap (sugar concentration, δ(13) C, Δ(14) C) as the mobilized component of stored stem NSC during early springtime. Using the bomb-spike radiocarbon approach we were able to estimate the average time elapsed since the mobilized carbon (C) was originally fixed from the atmosphere and to infer the turnover time of stem storage. Sites differed in concentration dynamics and overall δ(13) C, indicating different growing conditions. The absence of temporal trends for δ(13) C and Δ(14) C indicated sugar mobilization from a well-mixed pool with average Δ(14) C consistent with a mean turnover time (TT) of three to five years for this pool, with only minor differences between the sites. Sugar maple trees hence appear well buffered against single or even several years of negative plant C balance from environmental stress such as drought or repeated defoliation by insects. Manipulative investigations (e.g. starvation via girdling) combined with Δ(14) C measurements of this mobilized storage pool will provide further new insights into tree storage regulation and functioning.

Norway maple (Acer platanoides L), which is among the most invasive tree species in forests of eastern North America, is associated with reduced regeneration of the related native species, sugar maple (Acer saccharum Marsh) and other native flora. To identify traits conferring an advantage to Norway maple, we grew both species through an entire growing season under simulated light regimes mimicking a closed forest understorey vs. a canopy disturbance (gap). Dynamic shade-houses providing a succession of high-intensity direct-light events between longer periods of low, diffuse light were used to simulate the light regimes. We assessed seedling height growth three times in the season, as well as stem diameter, maximum photosynthetic capacity, biomass allocation above- and below-ground, seasonal phenology and phenotypic plasticity. Given the north European provenance of Norway maple, we also investigated the possibility that its growth in North America might be increased by delayed fall senescence. We found that Norway maple had significantly greater photosynthetic capacity in both light regimes and grew larger in stem diameter than sugar maple. The differences in below- and above-ground biomass, stem diameter, height and maximum photosynthesis were especially important in the simulated gap where Norway maple continued extension growth during the late fall. In the gap regime sugar maple had a significantly higher root shoot ratio that could confer an advantage in the deepest shade of closed understorey and under water stress or browsing pressure. Norway maple is especially invasive following canopy disturbance where the opposite (low root shoot ratio) could confer a competitive advantage. Considering the effects of global change in extending the potential growing season, we anticipate that the invasiveness of Norway maple will increase in the future. © 2012 The Author. Published by Oxford University Press. All rights reserved.

The majority of the world's forests are used for multiple purposes, which often include the potentially conflicting goals of timber production and biodiversity conservation. A scientifically validated management approach that can reduce such conflicts is retention forestry, an approach modeled on natural processes, which emerged in the last 25 years as an alternative to clearcutting. A portion of the original stand is left unlogged to maintain the continuity of structural and compositional diversity. We detail retention forestry's ecological role, review its current practices, and summarize the large research base on the subject. Retention forestry is applicable to all forest biomes, complements conservation in reserves, and represents bottom-up conservation through forest manager involvement. A research challenge is to identify thresholds for retention amounts to achieve desired outcomes. We define key issues for future development and link retention forestry with land-zoning allocation at various scales, expanding its uses to forest restoration and the management of unevenage forests. Fil: Gustafsson, Lena. Sveriges Lantbruksuniversitet; Suecia Fil: Baker, Susan C.. University of Tasmania; Australia Fil: Bauhus, Jrgen. Universität Freiburg Im Breisgau; Alemania Fil: Beese, William J.. Vancouver Island University; Canadá Fil: Brodie, Angus. No especifica; Fil: Kouki, Jari. Itä-suomen Yliopisto; Finlandia Fil: Lindenmayer, David B.. Australian National University; Australia Fil: Lhmus, Asko. University Of Tartu; Estonia Fil: Martínez Pastur, Guillermo José. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina Fil: Messier, Christian. Le Réseau de L'université Du Québec; Canadá Fil: Neyland, Mark. Forestry Tasmania; Australia Fil: Palik, Brian. United States Department of Agriculture; Argentina Fil: Sverdrup Thygeson, Anne. Universitetet For Miljø- Og Biovitenskap; Noruega Fil: Volney, W. Jan A.. Northern Forestry Centre; Canadá Fil: Wayne, Adrian. Western Australia Department Of Environment And Conservation; Australia Fil: Franklin, Jerry F.. University Of Washington, Seattle

Approximately 85% of the global forest estate is neither formally protected nor in areas dedicated to intensive wood production (e.g., plantations). Given the spatial extent of unprotected forests, finding management approaches that will sustain their multiple environmental, economic, and cultural values and prevent their conversion to other uses is imperative. The major global challenge of native forest management is further demonstrated by ongoing steep declines in forest biodiversity and carbon stocks. Here, we suggest that an essential part of such management—supplementing the protection of large reserves and sensitive areas within forest landscapes (e.g., aquatic features)—is the adoption of the retention approach in forests where logging occurs. This ecological approach to harvesting provides for permanent retention of important selected structures (e.g., trees and decayed logs) to provide for continuity of ecosystem structure, function, and species composition in the postharvest forest. The retention approach supports the integration of environmental, economic, and cultural values and is broadly applicable to tropical, temperate, and boreal forests, adaptable to different management objectives, and appropriate in different societal settings. The widespread adoption of the retention approach would be one of the most significant changes in management practice since the onset of modern high-yield forestry.

Numerous crown parameters (e.g., leaf area index, diameter, height, volume) can be obtained via the analysis of tree crown photographs. In all cases, parameter values are functions of the position of the crown outline. However, no standardized method to delineate crowns exists. To explore the effect of different outlines on tree crown descriptors, in this case crown openness (CO), and facilitate the adoption of a standard method free of user bias, we developed the program Crown Delineator that automatically delineates any outline around tree crowns following predetermined sensibility settings. We used different outlines to analyze tree CO in contrasting settings: using saplings from four species in young boreal mixedwood forests and medium-sized hybrid poplar trees from a low-density plantation. In both cases, the estimated CO increases when calculated from a looser outline, which had a strong influence on understory available light simulations using a forest simulator. These results demonstrate that the method used to trace crown outlines is an important step in the determination of CO values. We provide a much-needed computer-assisted solution to help standardize this procedure, which can also be used in many other situations in which the delineation of tree crowns is needed (e.g., competition and crown shyness).

Competition is a major determinant of plant growth and is often used in studies of tree growth and species coexistence. However, these approaches are usually temporally static, i.e., assessed at a single point or period in time. While constantly changing forest conditions due to natural and human-induced disturbances potentially alter competition among individuals, static approaches cannot qualify the temporal variability of competitive interactions. Here we present a longitudinal analysis of competitive interactions among trees and discuss the implication of our results for ecological interpretation. Spatially-explicit tree growth data were obtained from 18 study plots (0.4 ha each) in sugar maple (Acer saccharum Marsh.) stands in Quebec, Canada. During the studied period (1980–2003), these stands had been disturbed by insect outbreaks (forest tent caterpillar, Malacosoma disstria Hubner) and by commercial partial harvest. We analyzed radial growth rates (outcome of competition) on an annual basis and as a function of tree biology (bole diameter, crown position), competition (above- and belowground competition from neighbours) and environmental conditions (light availability, harvest disturbance). Competitive interactions changed throughout the studied period. Canopy disturbance from partial harvest interacted with defoliators and influenced competition symmetry by favoring smaller trees. Competitive interactions seemed to have switched from below- to above-ground following canopy recovery after harvest. Release from competition due to partial harvest increase neighbourhood size (radius of effective competition) and enhanced the competitive pressure from larger individuals. The temporal variability in parameter estimates may be used for setting confidence intervals on competitive success (growth rates), thereby yielding a more robust basis for ecological interpretation. Our results also show that temporal variability in competitive interactions could contribute to the maintenance of high tree species diversity and structural complexity in some ecosystems by temporally altering species-specific responses to environmental change and disturbance.

Protected areas cover over 12% of the terrestrial surface of Earth, and yet many fail to protect species and ecological processes as originally envisioned. Results of recent studies suggest that a critical reason for this failure is an increasing contrast between the protected lands and the surrounding matrix of often highly altered land cover. We measured the isolation of 114 protected areas distributed worldwide by comparing vegetation-cover heterogeneity inside protected areas with heterogeneity outside the protected areas. We quantified heterogeneity as the contagion of greenness on the basis of NDVI (normalized difference vegetation index) values, for which a higher value of contagion indicates less heterogeneous land cover. We then measured isolation as the difference between mean contagion inside the protected area and mean contagion in 3 buffer areas of increasing distance from the protected-area border. The isolation of protected areas was significantly positive in 110 of the 114 areas, indicating that vegetation cover was consistently more heterogeneous 10-20 km outside protected areas than inside their borders. Unlike previous researchers, we found that protected areas in which low levels of human activity are allowed were more isolated than areas in which high levels are allowed. Our method is a novel way to assess the isolation of protected areas in different environmental contexts and regions.

Partial harvesting, where different numbers and arrangements of live trees are retained in forest stands, has been proposed for maintaining late-successional structure and associated vertebrate species within managed boreal forests. Using the stand dynamics model SORTIE-ND, we examined 80-year patterns of structural change in response to different intensities (30–70% basal area removal) and spatial patterns (22–273 m 2 mean patch size) of harvesting. We also applied habitat models for seven late-successional vertebrates to the structural conditions present after harvesting to assess potential species responses. Partial harvesting increased understory and downed woody debris (DWD) cover and decreased overstory structure for the first 25 years after harvest, in comparison to unharvested stands, with this effect subsequently reversing as harvest-induced regeneration reached the canopy. Although harvesting enhanced long-term structural development in this regard, large trees, large snags, and large DWD all remained below unharvested levels throughout the simulation period. Harvesting also produced transient increases in early-decay DWD and ground exposure. Most changes in structural attributes increased in proportion to harvest intensity, but structural differences among harvest patterns were generally small. Dispersed harvesting induced somewhat less pronounced decreases in vertical structure, and produced more post-harvest slash, than aggregated harvesting. All seven vertebrate species decreased in abundance as harvest intensity increased from 30 to 70%. In comparison to their pre-harvest abundances in old stands, vertebrates associated with DWD (redback salamander, marten, red-backed vole) showed neutral or positive responses at one or more harvest intensities, whereas those associated with large trees and snags (brown creeper, flying squirrel) consistently exhibited substantial adverse impacts.

Selection cutting, where approximately 30% of the trees are removed every 30 years, is the main silvicultural treatment used in temperate deciduous forests of Quebec (Canada). Concerns have been raised that the use of heavy machinery is creating soil disturbances that are negatively affecting the growth and survival of remaining trees. The aim of the study was to determine if heavy machinery is affecting the growth, morphology, and architecture of sugar maple (Acer saccharum Marsh.) fine roots in and around machinery tracks left after selection logging. The study site, a sugar maple dominated stand, was located in southern Quebec. Root ingrowth bags and standard root cores were used to compare fine root growth, morphology, and architecture in and around machinery tracks one year after logging. Fine root growth of maple was reduced fivefold in both primary (multiple trip) and secondary (only one trip) machinery tracks compared with the control. There was a nonstatistical reduction in fine root growth within 1 m of the tracks. Because machinery tracks cover between 15% and 25% of a stand after selection logging, such reduction in fine root growth could be significant for the growth and survival of the remaining mature maple trees.

Northern hardwoods have undergone a marked change in their dynamics, with American beech (Fagus grandifo- lia Ehrh.) increasing in abundance relative to sugar maple (Acer saccharum Marsh.). This study aims to better understand this sudden shift in recruitment dynamics. We performed an extensive analysis of the age structure, radial growth pattern, and release history on >700 saplings from 34 mature maple-beech stands of southern Quebec. We found (i) that the sapling age structures showed a progressive decrease in the establishment of maple relative to beech starting about 40 years ago, (ii) a change in the species growth hierarchy that started in the 1980s due to increasing radial growth of beech, (iii) that this growth trend is negligible for both maple and beech when we account for size and suppression status, and finally (iv) that the growth trend appears to be independent of present soil conditions. These results contrast with previous studies conducted at the adult stage that reported a growth decline for maple. We conclude that this change in recruitment dynamics is not re- lated to growth, and consequently, further studies investigating this phenomenon should concentrate on establishment and survival. Resume : La dynamique des forets de feuillus temperees a change de facon marquee avec l'augmentation de l'abondance du hetre a grandes feuilles (Fagus grandifolia Ehrh.) par rapport a celle de l'erable a sucre (Acer saccharum Marsh.). Cette etude vise a mieux comprendre ce changement soudain de dynamique du recrutement. Nous avons realise une analyse de la structure d'âge, du patron de croissance radiale et de l'historique de degagement de plus de 700 gaules provenant de 34 peu- plements matures d'erable et de hetre du sud du Quebec. Nous avons trouve (i) que la structure d'âge des gaules indiquait une diminution progressive de l'etablissement de l'erable par rapport au hetre, processus qui a commence il y a environ 40 ans, (ii) un changement de hierarchie dans la croissance des especes qui a commence dans les annees 1980 a la suite d'une hausse de croissance radiale du hetre, (iii) que cette tendance de croissance est negligeable pour l'erable et le hetre lorsqu'on tient compte de la taille et du statut d'oppression des individus, et enfin (iv) que la tendance de croissance semble indepen- dante des presentes conditions de sol. Ces resultats sont differents de ceux d'etudes anterieures realisees au stade adulte et qui rapportaient un declin de la croissance de l'erable. Nous concluons que ce changement de dynamique du recrutement n'est pas relie a la croissance et, par consequent, que les futures etudes portant sur ce phenomene devraient se concentrer sur l'etablissement et la survie. (Traduit par la Redaction)

The silviculture of hybrid poplars and other fast-growing tree species is a promising solution to reduce the pressure on natural forests while maintaining wood supplies to industries. However, hybrid poplars are very sensitive to competing vegetation and to inadequate soil conditions and fertility. Possible management tools include mechanical site preparation (MSP), vegetation control (VC), and fertilization. Experimental plantations of hybrid poplars (one clone, Populus balsamea×Populus maximowiczii) were established at eight formerly forested sites on acidic soil in the southern boreal forest of Quebec, Canada. The objective was to test the response of hybrid poplars to the interaction of several silvicultural tools, which has been rarely done. FourMSPtreatments (in decreasing order of intensity: mounding, harrowing, heavy disk trenching, light disk trenching) and a control (unprepared) were all combined with four different frequencies of plant competition control by brushing (from never up to once a year). Fertilization with N or N+ P was also tested in three selected MSP treatments. After five years, hybrid poplar tree growth among MSP treatments increased in the following order: unprepared < light disk trenching < heavy disk trenching < harrowing < mounding. MSP was also essential in favouring early tree survival, as illustrated by mortality rates of over 20% in unprepared plots and below 5% in all other MSP treatments. The effect of competition control on hybrid poplar growth was greatest in the less intensive MSP treatments, where competing vegetation was the most abundant. On the contrary, fertilization effect was significant only in the most intensive MSP (mounding). Moreover, neither fertilization nor VC could compensate for inadequate soil preparation. Of all the silvicultural treatments tested, mounding provided the best tree growth despite a nitrogen and carbon impoverished surface soil.

Partitioning of elements in tree xylem is being increasingly studied, as it suggests that elements are potentially mobile within the xylem long after their uptake. A recent study revealed that only the most mobile xylem fraction (water-soluble) of base cations (calcium [Ca], magnesium [Mg], and potassium [K]) increased at higher soil acidity, while the two mobile fractions (water- and acid-soluble) of acidic metals—potentially phytotoxic aluminium (Al), cadmium (Cd) and manganese (Mn)—were significantly enhanced on very acid soils. The current paper presents an investigation of soil–wood chemistry relationships with basal area tree growth. It was hypothesized that the growth of sugar maple would be reduced by low base cation and high acidic metal concentrations in the xylem mobile fractions. Sugar maple trees (n = 55) from six watersheds in southern Quebec, Canada were analysed by sequential chemical extractions for the water-soluble, acid-soluble and residual fractions of base cations (Ca, K, Mg) and acidic metals (Al, Cd, Mn) in xylem. Generally, tree growth was positively correlated to concentrations of base cations in wood (ρ = 0.27–0.50) and soil (ρ = 0.41–0.67), and negatively correlated to concentrations of acidic metals in wood (ρ = −0.33 to −0.52) and soil (ρ = −0.67). However, these relations differed depending on the element fraction considered. Water- and acid-soluble xylem concentrations of base cations and Al were among the best predictors of growth trends (R2 = 0.46–0.51). The relationship between acidic metals and tree growth is further discussed.

In the context of partial harvesting, adequately managing post-harvest light conditions are essential to obtain a desired composition of tree species regeneration. The objective of this study was to determine how varying the intensity and spatial pattern of harvest would affect understory light conditions in boreal mixedwood stands of northwestern Quebec using the spatially explicit SORTIE-ND light model. The model was evaluated based on comparisons of observed and predicted light levels in both mapped and un-mapped plots. In mapped plots, reasonably accurate predictions of the overall variation in light levels were obtained, but predictions tended to lack spatial precision. In un-mapped plots, SORTIE-ND accurately predicted stand-level mean GLI (Gap Light Index) under a range of harvest intensities. The model was then used to simulate nine silvicultural treatments based on combinations of three intensities of overstory removal (30%, 45% and 60% of basal area) and three harvest patterns (uniform, narrow strips, large gaps). Simulations showed that increasing overstory removal had less impact on light conditions with uniform harvests, and a more marked effect with more aggregated harvest patterns. Whatever the harvest intensity, uniform cuts almost never created high light conditions (GLI > 50%). Gap cuts, on the other hand, resulted in up to 40% of microsites receiving GLI > 50%. Our results suggest that either a 30% strip or gap cut or a 45–60% uniform partial harvest could be used to accelerate the transition from an aspen dominated composition to a mixedwood stand because both types of cut generate the greatest proportion of moderately low light levels (e.g., 15–40% GLI). These light levels tend to favour an accelerated growth response among shade-tolerant conifers, while preventing excessive recruitment of shade-intolerant species. A better understanding of how spatial patterns of harvest interact with tree removal intensity to affect understory light conditions can provide opportunities for designing silvicultural prescriptions that are tailored to species’ traits and better suited to meet a variety of management objectives.

L’analisi delle problematiche inerenti le risorse naturali in un periodo di profondi cambiamenti globali presenta nuove sfide per il mondo forestale, sfide che hanno dato vita a nuovi tipi di approcci alla gestione delle foreste in considerazione della molteplicita di beni e servizi che queste forniscono. Uno di questi approcci e basato sulla scienza dei sistemi complessi. Considerare le foreste come Sistemi Complessi Adattativi (o CAS) rappresenta un paradigma emergente basato sullo sviluppo della teoria dei sistemi in altri settori come la fisica, la medicina, l’economia. Questo nuovo quadro concettuale riconosce la complessita dei sistemi (ecologici, economici e sociali), le loro strutture gerarchiche, le interazioni e i flussi di energia tra queste gerarchie, e la loro capacita di auto-organizzazione e di adattamento. Una delle maggiori sfide in campo forestale e come imparare a utilizzare questi concetti per favorire la capacita dei sistemi forestali di auto-organizzarsi e adattarsi di fronte al cambiamento globale, in modo da permettere alla foreste di continuare a soddisfare i bisogni umani quali beni e servizi ecosistemici. In questo lavoro si analizza brevemente la scienza della complessita applicata alla gestione forestale e si discute come questa scienza possa essere rapportata agli strumenti di modellistica utilizzati nella gestione forestale. In particolare, ci si sofferma sul concetto di prevedibilita e sulle sfide legate all’uso di modelli, per mettere in evidenza le differenze tra la visione convenzionale della foresta e la visione della foresta come un sistema complesso adattativo. Si conclude proponendo un nuovo paradigma di gestione forestale, piu adatto alle sfide del futuro, basato su eterogeneita, imprevedibilita e adattabilita, piuttosto che su uniformita e prevedibilita e sull’approccio “comando e controllo”.

Forest management practices conducted primarily at the stand scale result in simplified forests with regeneration problems and low structural and biological diversity. Landscape models have been used to help design management strat- egies to address these problems. However, there remains a great deal of uncertainty that the actual management practices result in the desired sustainable landscape structure. To investigate our ability to meet sustainable forest management goals across scales, we assessed how two models of forest dynamics, a scaled-up individual-tree model and a landscape model, simulate forest dynamics under three types of harvesting regimes: clearcut, gap, and uniform thinning. Althougth 50- 100 year forecasts predicted average successional patterns that differed by less than 20% between models, understory dy- namics of the landscape model were simplified relative to the scaled-up tree model, whereas successional patterns of the scaled-up tree model deviated from empirical studies on the driest and wettest landtypes. The scale dependencies of both models revealed important weaknesses when the models were used alone; however, when used together, they could pro- vide a heuristic method that could improve our ability to design sustainable forest management practices. Resume´ : Les pratiques d'amenagement forestier qui sont surtout appliquees a l'echelle du peuplement entraoˆnent une sim- plification de la foret et engendrent des problemes de regeneration ainsi qu'une faible diversitestructurale et biologique. Des modeles de paysage ont eteutilises pour aider a elaborer des strategies qui s'attaquent aces problemes. Cependant, beaucoup d'incertitude persiste quant asavoir si les pratiques actuelles d'amenagement produisent une structure vraiment durable du paysage. Dans le but d'examiner notre capaciteaatteindre les objectifs de l'amenagement forestier durable peu importe l'echelle, nous avons evaluede quelle facon deux modeles de dynamique forestiere, un modele elargi d'arbre indi- viduel et un modele de paysage, simulent la dynamique de la foret soumise a trois regimes de coupe : rase, par trouees et eclaircie uniforme. Meme si les previsions sur 50 a 100 ans produisaient des patrons de succession moyens qui differaient par moins de 20 % selon le modele, la dynamique de sous-bois du modele de paysage etait simplifiee relativement au mo- dele elargi d'arbre individuel, tandis que les patrons de succession du modele elargi d'arbre individuel deviaient des etudes empiriques sur les stations les plus seches et les plus humides. Le fait que les deux modeles soient dependants de l'echelle fait ressortir d'importantes faiblesses lorsqu'ils sont utilisess eparement. Par contre, utilises ensemble ils pourraient fournir une methode heuristique capable d'ameliorer notre capaciteaconcevoir des pratiques d'amenagement forestier durable. (Traduit par la Redaction)

Aim An important issue regarding biodiversity concerns its influence on ecosystem functioning.Experimental work has led to the proposal of mechanisms such as niche complementarity.However,few attempts have been made to confirm these in natural systems, especially in forests. Furthermore, one of the most interesting unresolved questions is whether the effects of complementarity on ecosystem functioning (EF) decrease in favour of competitive exclusions over an increasing productivity gradient. Using records from permanent forest plots, we asked the following questions.(1) Is tree productivity positively related to diversity? (2) Does the effect of diversity increase in less productive forests? (3) What metric of diversity (e.g. functional or phylogenetic diversity) better relates to tree productivity? Location Temperate, mixed and boreal forests of eastern Canada. Methods Over 12,000 permanent forest plots, from temperate to boreal forests, were used to test our hypotheses in two steps.(1) Stepwise regressions were used to identify the best explanatory variables for tree productivity. (2) The selected climatic and environmental variables,as well as density and biodiversity indices,were included in a structural equation model where links (paths) between covarying variables are made explicit, making structural equation modelling the best tool to explore such complicated causal networks. Results This is the first large-scale demonstration of a strong, positive and significant effect of biodiversity on tree productivity with control for climatic and environmental conditions. Important differences were noted between the two forest biomes investigated. Main conclusions We show for the first time that complementarity may be less important in temperate forests growing in a more stable and productive environment where competitive exclusion is the most probable outcome of species interactions, whereas in the more stressful environment of boreal forests, beneficial interactions between species may be more important. The present work is also a framework for the analysis of large datasets in biodiversity‐ecosystem functioning (B-EF) research.

Neighbourhood competition indices (NCI), where position and species identity of neighbours are known, have been used to investigate growth and competitive interactions among adult trees. In this study, we used NCI in 8–15-year-old stands following clear-cutting in a boreal mixedwood forest of eastern Canada to improve our understanding of early successional forest dynamics. Trees of increasing diameter from the center (≥1 cm) to the edge (≥5 cm) were mapped in twenty-five circular 450m2 plots. Target trees (DBH≥1 cm) were sampled in plot center to determine their annual radial stem growth. For each species, we compared a set of growth models using either a spatially explicit NCI or a non-spatial competition index. Both types of indices estimated a species-specific competition coefficient for each pair of competitor–target species. NCI were selected as the best competition model for all target species although differences in variance explained relative to the non-spatial index were small. This likely indicates that competition occurs at the local level but that the high density and the relative uniformity of these young stands creates similar neighbourhoods for most trees in a given stand. The effective neighbourhood radius for competitors varied among species and was smaller for shade tolerant species. Intraspecific neighbours were the strongest competitors for most species. Aspen (Populus tremuloides) was a weak competitor for all species as opposed to balsam fir (Abies balsamea) which was a strong competitor in all cases. These results are in contradiction with some widely used forest policies in North America (e.g. free-to-grow standards) that consider broadleaf species, such as aspen, as the strongest competitors. For these early successional forests, the decision regarding the use of spatial or non-spatial competition indices should rest on the intended use. For even-age management, spatial indices might not justify their use in highdensity stands but they are needed for the simulation of novel harvest techniques creating complex stand structure.

The challenge of sustainable forest management is to integrate diverse and sometimes conflicting management objectives. In order to achieve this goal, we need a better understanding of the aspects influencing the preferences of diverse groups and how these groups make trade-offs between different attributes of SFM. We compare the SFM preferences of interest groups in regions with different forest use histories based on the reasoning that the condition of the forest reflects the forest use history of the area. The condition of the forest also shapes an individual's forest values and attitudes. These held values and attitudes are thought to influence SFM preferences. We tested whether the SFM preferences vary amongst the different interest groups within and across regions. We collected data from 252 persons using a choice experiment approach, where participants chose multiple times among different options described by a combination of attributes that are assigned different levels. The novelty of our approach was the use of choice experiments in the assessment of regional preference differences. Given the complexity of inter-regional comparison and the small sample size, this was an exploratory study based on a purposive rather than random sample. Nevertheless, our results suggest that the aggregation of preferences of all individuals within a region does not reveal all information necessary for forest management planning since opposing viewpoints could cancel each other out and lead to an interpretation that does not reflect possibly polarised views. Although based on a small sample size, the preferences of interest groups within a region are generally statistically significantly different from each other; however preferences of interest groups across regions are also significantly different. This illustrates the potential importance of assessing heterogeneity by region and by group.

In the Neotropics, there is a growing interest in establishing plantations of native tree species for commerce, local consumption, and to replant on abandoned agricultural lands. Although numerous trial plantations have been established, comparative information on the performance of native trees under different regional environments is generally lacking. In this study, we evaluated the accumulation and partitioning of above-ground biomass in 16 native and two exotic tree species growing in replicated species selection trials in Panama under humid and dry regional environments. Seven of the 18 species accumulated greater total biomass at the humid site than at the dry site over a two-year period. Speciesspecific biomass partitioning among leaves, branches and trunks was observed. However, a wide range of total biomass found among species (from 1.06 kg for Dipteryx panamensis to 29.84 kg for Acacia mangium at Soberania) justified the used of an Aitchison log ratio transformation to adjust for size. When biomass partitioning was adjusted for size, a majority of these differences proved to be a result of the ability of the tree to support biomass components rather than the result of differences in the regional environments at the two sites. These findings were confirmed by comparative ANCOVAs on Aitchison-transformed and non-Aitchison-transformed variables. In these comparisons, basal diameter, height and diameter at breast height were robust predictors of biomass for the pooled data from both sites, but Aitchison-transformed variables had little predictive power. © 2010 Elsevier B.V. All rights reserved.

We examined how the density, growth and survival of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton) regeneration are influenced by gap size, soil nutrient availability and understory vegetation. We used a factorial combination of (1) three gap sizes (small

Th e belief that canopy gaps are important for the maintenance of tree species diversity appears to be widespread, but there have been no formal theoretical models to assess under what conditions gap phase processes allow coexistence. Much of the empirical research on niche diff erentiation in response to gaps has focused on evidence for an interspecifi c tradeoff between low light survival and high light growth. Th e objectives of this study are fi rst to distinguish the possible mechanisms allowing coexistence based on this tradeoff , and second, to explore their limitations. We present a theory of forest dynamics driven by small-scale disturbances as a special case of the theory of coexistence in variable environments. We demonstrate that temporal and spatial heterogeneity in light conditions that results from canopy gaps can allow stable coexistence as a result of three previously documented general mechanisms: ‘relative non-linearity’, ‘the successional niche’ and the ‘storage eff ect’. We fi nd that temporal fl uctuations in light availability alone allow the stable coexistence of only two species. Spatial variation in disturbance synchronicity and intensity allows three species to coexist in a narrow parameter space. Th e rate of extinction is, however, extremely slow and there is transient coexistence of a larger number of species for a long period of time. We conclude that while the low light survival/high light growth tradeoff may be ubiquitous in forest tree species, it is unlikely to function as an important mechanism for the stable coexistence of several tree species.

The public view of tree plantations is somewhat ambiguous. While planting a single tree is generally considered good for the environment, planting a million trees raises concerns in some circles. Although plantations are often used to compensate for bad forestry practices, to willingly simplify otherwise complex forest ecosystems, or as a strategy for allowing the current petroleum-based economy to continue on its course, we believe plantations have a legitimate place in the sustainable management of forests. Multi-purpose plantations, designed to meet a wide variety of social, economic, and environmental objectives, can provide key ecosystem services, help preserve the world's remaining primary forests, and sequester an important proportion of the atmospheric carbon released by humans over the past 300 years.

Forest management has been criticised in the last 20 years for its negative impact on the native species, structures and functions of the forest. Of many possible alternatives proposed to minimize these effects, the functional zoning (or TRIAD) approach is gaining popularity in North America. The goal of this approach is to minimize the negative environmental impacts of forestry while maintain timber supply by dividing the forest into three broad land-use zones: (1) conservation, (2) ecosystem management, and (3) wood production. In this study, we used a spatially explicit landscape model to simulate the effects of fire and six different forest management scenarios on a boreal mixedwood forest management unit in central Quebec. The management scenarios examined included the current practices scenario, a scenario proposed by the provincial government, and four TRIAD scenarios varying in the amount of forest allocated to each of the three zones. For each scenario, we examined the harvest volume, percentage old-growth forest or old forest managed to favour old-growth attributes, and effective mesh size of forest patches by 20-year age classes. With more area set aside for conservation and high-retention partial cut harvesting techniques designed to maintain the attributes of old-growth stands, all TRIAD scenarios resulted in higher percentages of stands with old-growth attributes than the current practices scenario and the government proposed scenario, and two of the four TRIAD scenarios also resulted in higher harvest volume over the long term. All forest management scenarios resulted in significantly lower effective mesh size than the fire-only scenario, but this difference was not as pronounced for the four TRIAD scenarios as for the current practice and government proposed scenarios. We conclude that the TRIAD approach has the potential to minimize some of the negative impacts of forestry on the landscape, while maintaining timber supply over the long term.

The TRIAD approach to forest management involves dividing the forest into 3 zones, each with its own management objectives, but with the overall goal of increasing the ecological and economic sustainability of the forest. For the past 5 years, we have been experimenting with TRIAD zoning in central Quebec, incorporating social interests into the original concept of TRIAD management. Results generally indicate that this approach is economically viable, socially acceptable, and preferable ecologically in this area. Although much remains to be done, thus far the consensus among the various project participants is that this approach may be a good fit for the public forest of Canada. Key words: TRIAD, functional zoning, sustainable forest management, ecosystem management, natural disturbance based management, intensive forestry, mixed plantations, partial cuts, eco-efficiency, conservation, new forestry, green accounting

Large-scale synchronous variations in community dynamics are well documented for a vast array of organisms, but are considerably less understood for forest trees. Because of temporal variations in canopy gap dynamics, forest communities-even old-growth ones-are never at equilibrium at the stand scale. This paucity of equilibrium may also be true at the regional scale. Our objectives were to determine (1) if nonequilibrium dynamics caused by temporal variations in the formation of canopy gaps are regionally synchronized, and (2) if spatiotemporal variations in canopy gap formation affect the relative abundance of tree species in the understory. We examined these questions by analyzing variations in the suppression and release history of Acer saccharum Marsh. and Fagus grandifolia Ehrh. from 481 growth series of understory saplings taken from 34 mature stands. We observed that (1) the proportion of stems in release as a function of time exhibited a U-shaped pattern over the last 35 years, with the lowest levels occurring during 1975-1985, and that (2) the response to this in terms of species composition was that A. saccharum became more abundant at sites that had the highest proportion of stems in release during 1975-1985. We concluded that the understory dynamics, typically thought of as a stand-scale process, may be regionally synchronized.

In this study, we quantified the effects of local neighbourhood competition, light availability, and proximity to skid trails on the growth of sugar maple (Acer saccharum Marsh.) trees following selection harvest. We hypothesized that growth would increase with decreasing competition and increasing light availability, but that proximity to skid trails would negatively affect growth. A total of 300 sugar maples were sampled ∼10 years after selection harvesting in 18 stands in Temiscamingue (Quebec, Canada). Detailed tree and skid trail maps were obtained in one 0.4 ha plot per stand. Square-root transformed radial growth data were fitted to a linear mixed model that included tree diameter, crown position, a neighbourhood competition index, light availability (estimated using the SORTIE light model), and distance to the nearest skid trail as explanatory variables. We considered various distance-dependent or -independent indices based on neighbourhood radii ranging from 6 to 12 m. The competition index that provided the best fit to the data was a distance-dependent index computed in a 6 m search radius, but a distance-independent version of the competition index provided an almost equivalent fit to data. Models corresponding to all combinations of main effects were fit to data using maximum likelihood, and weighted averages of parameter estimates were obtained using multimodel inference. All predictors had an influence on growth, with the exception of light. Radial growth decreased with increasing tree diameter, level of competition and proximity to skid trails, and varied among crown positions with trees in suppressed and intermediate positions having lower growth rates than codominants and dominants. Our results indicate that in selection managed stands, the radial growth of sugar maple trees depends on competition from close (≤6 m) conspecific neighbours, and is still affected by proximity to skid trails ∼10 years after harvesting. Such results underscore the importance of minimizing the extent of skid trail networks by careful pre-harvest planning of trail layout. We also conclude that the impact of heterogeneity among individual-tree neighbourhoods, such as those resulting from alternative spatial patterns of harvest, can usefully be integrated into models of post-harvest tree growth.

Questions: What climate variables best explain fire occurrence and area burned in the Great Lakes-St Lawrence forest of Canada? How will climate change influence these climate variables and thereby affect the occurrence of fire and area burned in a deciduous forest landscape in Temiscamingue, Quebec, Canada? Location: West central Quebec and the Great Lakes-St Lawrence forest of Canada. Methods: We first used an information-theoretic framework to evaluate the relative role of different weather variables in explaining occurrence and area burned of large fires (4200 ha, 1959-1999) across the Great Lakes- St Lawrence forest region. Second, we examined how these weather variables varied historically in Temiscamingue and, third, how they may change between the present and 2100 according to different scenarios of climate change based on two Global Circulation Models. Results: Mean monthly temperature maxima during the fire season (Apr-Oct) and weighted sequences of dry spells best explained fire occurrence and area burned. Between 1910 and 2004, mean monthly temperature maxima in Temiscamingue showed no apparent temporal trend, while dry spell sequences decreased in frequency and length. All future scenarios show an increase in mean monthly temperature maxima, and one model scenario forecasts an increase in dry spell sequences, resulting in a slight increase in forecasted annual area burned. Conclusion: Despite the forecasted increase in fire activity, effects of climate change on fire will not likely affect forest structure and composition as much as natural succession or harvesting and other disturbances, principally because of the large relative difference in area affected by these processes.

Silviculture to maintain old-growth forest attributes appears to be an oxymoron since the late developmental phases of forest dynamics, described by the term old-growth, represent forests that have not experienced human intervention or timber removal for a long time. In the past, silvicultural systems applied to old-growth aimed to convert it into simplified, more productive regrowth forests substantially different in structure and composition. Now it is recognised that the maintenance of biodiversity associated with structural and functional complexity of late forest development successional stages cannot rely solely on old-growth forests in reserves. Therefore, in managed forests, silvicultural systems able to develop or maintain old-growth forest attributes are being sought. The degree to which old-growth attributes are maintained or developed is called “old-growthness”. In this paper, we discuss silvicultural approaches that promote or maintain structural attributes of old-growth forests at the forest stand level in (a) current old-growth forests managed for timber production to retain structural elements, (b) current old-growth forests requiring regular, minor disturbances to maintain their structure, and (c) regrowth and secondary forests to restore old-growth structural attributes. While the functions of different elements of forest structure, such as coarse woody debris, large veteran trees, etc., have been described in principle, our knowledge about the quantity and distribution, in time and space, of these elements required to meet certain management objectives is rather limited for most ecosystems. The risks and operational constraints associated with managing for structural attributes create further complexity, which cannot be addressed adequately through the use of traditional silvicultural approaches. Silvicultural systems used in the retention and restoration of old-growthness can, and need, to employ a variety of approaches for managing spatial and temporal structural complexity. We present examples of silvicultural options that have been applied in creative experiments and forestry practice over the last two decades. However, these largely comprise only short-term responses, which are often accompanied by increased risks and disturbance. Much research and monitoring is required still to develop and optimise new silvicultural systems for old-growthness for a wide variety of forest ecosystem types.

This research tested whether demonstration of the long term effect of different forest management scenarios in a large forested area changes people's forest values and attitudes. Forestry professionals and other forest users in Central Labrador were shown simulation results of three alternative forest management scenarios illustrating possible long term effects on various indicators. Forest values and attitudes towards forestry were measured before and after the presentation. Our conception of values and attitudes is based on the cognitive hierarchy model of human behaviour which states that values are more enduring and more difficult to change than attitudes. It was thus hypothesized that attitudes would change but not values and that change in forestry professionals would be less than in other forest users since foresters are trained to think about long-term effects and large-scale processes of forest management scenarios. We also hypothesized that a greater number of people would have an opinion on forest management after the presentation. All three hypotheses were partially supported by the results. The results indicated that some attitude change occurred, but that values also changed somewhat. Most of the significant changes occurred when persons with no clear opinion on several forest-related questions formed an opinion. Long-term, landscape simulation results provide valuable information and enhance understanding of both forestry professionals and other forest users. However, being provided the same information, the two groups learned different things. While forest users gained more confidence in the current forest management plan and were motivated to further participate, professionals learned more specific things. This reflects differences between technical and local knowledge.

This research aims at evaluating the effectiveness of communicating simulation results to enhance learning. This was done by testing whether showing simulation results which demonstrate the long-term effects of different man- agement strategies in a large area changes people's SFM preferences. Forest users in Central Labrador were shown simu- lation results of three alternative forest management scenarios illustrating possible long term effects on various indicators. SFM preferences were measured using an attribute-based choice experiment before and after the presentation. Based on earlier research and issues closely related to the attributes used in the choice experiment, we expected significant prefer- ence change and more consistent choices after presentation. No significant change was found, however. This may be due to the relative stability of SFM preferences in the region. It is also possible that more participants and a longer time frame would be needed to reveal change. However, detected changes in the parameter estimates for cut block size and alterna- tive-specific constant (ASC), measuring the tendency of selecting the status quo alternative, indicate that some learning occurred during the process.

Differences in the way local and regional interest groups perceive Sustainable Forest Management in regions with different forest use histories were studied using Southeastern Finland, the Mauricie in Quebec and Central Labrador in Canada as examples of regions with high, medium and low importance of commercial forestry. We present a conceptual model illustrating the cyclic interaction between the forest, cultural models about forests and forest management. We hypothesized that peoples' perceptions would be influenced by their cultural models about forests and would thus vary amongst regions with different forest use histories and among different interest groups. The weightings of the environmental, economic and social components of sustainability as well as themes important for each of the interest groups were elicited using individual listing of SFM indicators and group work aimed at developing a consensus opinion on a common indicator list. In Southeastern Finland the views of the different groups were polarized along the environment-economy axis, whereas in Central Labrador all groups were environmentally oriented. The social dimension was low overall except among the Metis and the Innu in Labrador. Only environmental groups were similar in all three research regions, the largest differences between regions were found among the forestry professionals in their weightings concerning economy and nature. As the importance of commercial forestry increased, a greater importance of economic issues was expressed whereas the opposite trend was observed for issues regarding nature. Also inter-group differences grew as the importance of commercial forestry increased in the region. Forest management and forest use can be seen as factors strongly influencing peoples' cultural models on forests.

Tree mortality is a major force driving forest dynamics. To foresters, however, tree mortality is often considered a loss in productivity. To reduce tree mortality, silvicultural systems, such as selection cuts, aim at removing trees that are more likely to die. In order to identify trees with higher risks of mortality, field classifications are employed that assess vigour based on external characteristics of trees. We used a novel longitudinal approach for estimating survival probabilities based on ring-width measurements, initially developed by Bigler and Bugmann [Bigler, C., Bugmann, H., 2004. Predicting the time of tree death using dendrochronological data. Ecol. Appl. 14 (3), 902–914], to parameterize a survival probability model for sugar maple (Acer saccharum Marsh.) and to test whether field-assessed tree vigour classes are corroborated by survival probabilities determined from radial growth history. Data from 56 dead and 321 live sugar maples were collected in stands in western Quebec (Canada) that had undergone a selection cut � 10 years prior to sampling. Our results showed that tree vigour established from external defects and

With an increasing proportion of natural forests being replaced by plantations, there is a need to determine their potential to fulfill ecological purposes other than wood production. This study evaluated the extent to which deciduous and coniferous plantations develop understory attributes comparable to those of naturally regenerated stands. A functional group approach was used to synthesise species responses in terms of their ecological traits. Multivariate analyses of ecological traits revealed 16 emergent groups that shared common traits associated with a similar life history strategy. Responses of these groups, understory structure, and understory environmental conditions to plantation types and stand stages were analyzed and compared to naturally regenerated stands. Clear associations of trait responses to stand developmental stages and plantation types emerged. Light-demanding and wind-dispersed species groups were associated with early-successional stages, while woody groups, ferns and ant-dispersed spring-flowering herbs were associated with late-successional stages. Analyses also revealed an indicator group associated with old naturally regenerated forest. The understory functional groups and environmental conditions of deciduous plantations converged toward those of old naturally regenerated forests. However, understory structure in deciduous plantations remained poorly developed and richness of the indicator group was low compared to unplanted stands. Conifer plantations, currently the most common plantation type in the northern hardwood biome, showed a completely different pathway of understory development. Modifications to current plantation management practices are proposed to help recreate or maintain natural understory biological and physical attributes.

Understanding coexistence of highly shade-tolerant tree species is a longstanding challenge for forest ecologists. A conceptual model for the coexistence of sugar maple (Acer saccharum) and American beech (Fagus grandibfolia) has been proposed, based on a low-light survival/high-light growth trade-off, which interacts with soil fertility and small-scale spatiotemporal variation in the environment. In this study, we first tested whether the spatial distribution of seedlings and saplings can be predicted by the spatiotemporal variability of light availability and soil fertility, and second, the manner in which the process of environmental filtering changes with regeneration size. We evaluate the support for this hypothesis relative to the one for a neutral model, i.e., for seed rain density predicted from the distribution of adult trees. To do so, we performed intensive sampling over 86 quadrats (5 x 5 m) in a 0.24-ha plot in a mature maple-beech community in Quebec, Canada. Maple and beech abundance, soil characteristics, light availability, and growth history (used as a proxy for spatiotemporal variation in light availability) were finely measured to model variation in sapling composition across different size classes. Results indicate that the variables selected to model species distribution do effectively change with size, but not as predicted by the conceptual model. Our results show that variability in the environment is not sufficient to differentiate these species' distributions in space. Although species differ in their spatial distribution in the small size classes, they tend to correlate at the larger size class in which recruitment occurs. Overall, the results are not supportive of a model of coexistence based on small-scale variations in the environment. We propose that, at the scale of a local stand, the lack of fit of the model could result from the high similarity of species in the range of environmental conditions encountered, and we suggest that coexistence would be stable only at larger spatial scales at which variability in the environment is greater.

• Background and Aims: Natural and anthropogenic disturbances can act as stresses on tree vigour. According to Manion's conceptual model of tree disease, the initial vigour of trees decreases as a result of predisposing factors that render these trees more vulnerable to severe inciting stresses, stresses that can then cause final vigour decline and subsequent tree death. This tree disease model was tested in sugar maple (Acer saccharum) by assessing the roles of natural and anthropogenic disturbances in tree decline and death. • Methods: Radial growth data from 377 sugar maple trees that had undergone both defoliations by insects and partial harvest were used to estimate longitudinal survival probabilities as a proxy for tree vigour. Radial growth rates and survival probabilities were compared among trees subjected to different levels of above- and below-ground disturbances, between periods of defoliation and harvest, and between live and dead trees. • Key Results: Manion's tree disease model correctly accounts for vigour decline and tree death in sugar maple; tree growth and vigour were negatively affected by a first defoliation, predisposing these trees to death later during the study period due to a second insect outbreak that initiated a final vigour decline. This decline was accelerated by the partial harvest disturbance in 1993. Even the most severe anthropogenic disturbances from partial harvest did not cause, unlike insect defoliation, any growth or vigour declines in live sugar maple. • Conclusions: Natural disturbances acted as predisposing and inciting stresses in tree sugar maple decline and death. Anthropogenic disturbances from a partial harvest at worst accelerated a decline in trees that were already weakened by predisposing and inciting stresses (i.e. repeated insect defoliations). Favourable climatic conditions just before and after the partial harvest may have alleviated possible negative effects on growth resulting from harvesting.

American beech (Fagus grandifolia Ehrh.) reproduces sexually, and vegetatively by root suckers. Although many studies have investigated its regeneration response, most did not account for differences that may exist between its two modes of reproduction. This study was performed in an old-growth Acer - Fagus forest in southern Quebec, where beech bark disease had only a minor effect at the time of the study. We compared the density and frequency of occurrence of beech seedlings and root suckers (height < 30 cm), as well as their morphology, growth, survival, and response to experimental defoliation. Root suckers accounted for ∼13% of beech regeneration at our site. Density and frequency of occurrence were greater for seedlings than suckers, but did not vary with light availability, which was low at our study site (mean: 2.9%). Seedlings and suckers did not differ in leaf characteristics, but several differences were observed in terms of plant morphology, growth, and survival. Root suckers showed more lateral growth than height growth, and had a lower leaf area index than seedlings. Root suckers had both a greater growth in height and diameter, and a higher survivorship than seedlings (height and diameter growth were, respectively, five and two times greater for suckers than seedlings, and 74% of suckers survived more than 1 year, compared to 52% for seedlings). Defoliation treatments, which included levels of defoliation of 50% and 100% (1) did not affect current-year extension growth of seedlings and suckers; (2) did not affect seedling diameter growth, but had a negative impact on sucker diameter growth; and (3) affected survivorship for both origins, but had a much greater negative impact on seedling survivorship (none of the completely defoliated seedlings survived over one year, while 55% of the suckers did). This study showed that several differences exist between small beech seedlings and root suckers in traits that are important determinants of a species' competitive ability. We therefore expect that variation in the relative importance of root suckering among sites might have several community-level implications.

Dendrochemistry has been used for the historical dating of pollution. Its reliability is questionable due primarily to the radial mobility of elements in sapwood. In the present study, the extractability of seven elements was characterized to assess their suitability for the monitoring of environmental conditions. Nine mature sugar maple trees (Acer saccharum Marsh.), a wide-ranging species in eastern North America that has suffered decline in past decades, were sampled in three Quebec watersheds along a soil acidity gradient. Five-year groups of annual tree rings were treated by sequential chemical extractions using extractants of varying strength (deionized H2O, 0.05 M HCl, and concentrated HNO(3)) to selectively solubilize the elements into three fractions (water-soluble, acid-soluble, and residual). Monovalent K; divalent Ba, Ca, Cd, Mg, Mn; and trivalent Al cations were found mostly in the water-soluble, acid-soluble, and residual fractions, respectively. Forms more likely to be mobile within the tree (water-soluble and acid-soluble) do not seem to be suitable for temporal monitoring because of potential lateral redistribution in sapwood rings. However, certain elements (Cd, Mn) were responsive to current soil acidity and could be used in spatial variation monitoring. Extractability of Al varied according to soil acidity; at less acidic sites, up to 90% of Al was contained in the residual form, whereas on very acidic soils, as much as 45% was found in the water-soluble and acid-soluble fractions. Sequential extractions can be useful for determining specific forms of metals as key indicators of soil acidification.

Crown openness (CO) of mature trees influences light transmission within the forest canopy. However, in modeling, this variable is often considered constant within species, and its potential regional variability is ignored. The objective of this study was to evaluate if CO values of yellow birch ( Betula alleghaniensis Britt.), sugar maple ( Acer saccharum Marsh.), and eastern hemlock ( Tsuga canadensis (L.) Carrière) vary according to the following factors: (i) species, (ii) regional actual evapotranspiration (AET), (iii) tree size (i.e., diameter at breast height, DBH), and (iv) angle of transmission from zenith. To achieve this, CO was evaluated for 136 yellow birches, 109 sugar maples, and 68 hemlocks from different regions of western Quebec, southern Ontario, and northern Michigan. Results showed that all of the studied factors affected CO. While dominant trees can intercept light laterally as well as vertically, smaller trees are more efficient at intercepting light vertically. Increasing AET is associated with more open crowns. Given its importance in light transmission in the understory, a better understanding of how CO varies between individuals, species, and regions is needed.

Question: In deciduous-dominated forest landscapes, what are the relative roles of fire weather, climate, human and biophysi cal landscape characteristics for explaining variation in large fire occurrence and area burned? Location: The Great Lakes-St. Lawrence forest of Canada. Methods: We characterized the recent (1959-1999) regime of large (≥ 200 ha) fires in 26 deciduous-dominated landscapes and analysed these data in an information-theoretic framework to compare six hypotheses that related fire occurrence and area burned to fire weather severity, climate normals, population and road densities, and enduring landscape characteristics such as surficial deposits and large lakes. Results: 392 large fires burned 833 698 ha during the study period, annually burning on average 0.07% ± 0.42% of forested area in each landscape. Fire activity was strongly seasonal, with most fires and area burned occurring in May and June. A combination of antecedent-winter precipitation, fire season precipitation deficit/surplus and percent of landscape covered by well-drained surficial deposits best explained fire occurrence and area burned. Fire occurrence varied only as a function of fire weather and climate variables, whereas area burned was also explained by percent cover of aspen and pine stands, human population density and two enduring characteristics: percent cover of large water bodies and glaciofluvial deposits. Conclusion: Understanding the relative role of these variables may help design adaptation strategies for forecasted increases in fire weather severity by allowing (1) prioritization of land scapes according to enduring characteristics and (2) management of their composition so that substantially increased fire activity would be necessary to transform landscape structure and composition.

Tree-ring chronologies have been widely used in studies of tree mortality where variables of recent growth act as an indicator of tree physiological vigour. Comparing recent radial growth of live and dead trees thus allows estimating probabilities of tree mortality. Sampling of mature dead trees usually provides death-year distributions that may span over years or decades. Recent growth of dead trees (prior to death) is then computed during a number of periods, whereas re- cent growth (prior to sampling) for live trees is computed for identical periods. Because recent growth of live and dead trees is then computed for different periods, external factors such as disturbance or climate may influence growth rates and, thus, mortality probability estimations. To counteract this problem, we propose the truncating of live-growth series to obtain similar frequency distributions of the ''last year of growth'' for the populations of live and dead trees. In this paper, we use different growth scenarios from several tree species, from several geographic sources, and from trees with different growth patterns to evaluate the impact of truncating on predictor variables and their selection in logistic regression analy- sis. Also, we assess the ability of the resulting models to accurately predict the status of trees through internal and external validation. Our results suggest that the truncating of live-growth series helps decrease the influence of external factors on growth comparisons. By doing so, it reinforces the growth-vigour link of the mortality model and enhances the model's accuracy as well as its general applicability. Hence, if model parameters are to be integrated in simulation models of greater geographical extent, truncating may be used to increase model robustness. Resume´ : La dendrochronologie etelargement utilisee dans les etudes portant sur la mortalitedes arbres oudes variables de croissance recente sont utilisees comme indicateur de la vigueur physiologique des arbres. La comparaison de la crois- sance radiale recente d'arbres vivants et morts permet donc d'estimer la probabilitede mortalitedes arbres. L'echantil- lonnage d'arbres matures morts fournit generalement la distribution des annees de mortalitequi peuvent s'etendre sur plusieurs annees ou decennies. La croissance recente des arbres morts (avant leur mort) est ensuite calculee pour un certain nombre de periodes alors que celle des arbres vivants (avant leur echantillonnage) est calculee pour des periodes identi- ques. Puisque la croissance recente des arbres vivants et morts est ensuite calculee pour des periodes differentes, des fac- teurs externes tels les perturbations ou le climat peuvent influencer le taux de croissance et, par consequent, l'estimation de la probabilitede mortalite´. Pour resoudre ce probleme, nous proposons de tronquer les series de croissance des arbres vivants de facon a obtenir des distributions de frequence similaires de «la derniere annee de croissance» pour les popula- tions d'arbres vivants et morts. Dans cette etude, nous utilisons differents scenarios de croissance apartir de plusieurs es- peces d'arbre et de plusieurs origines geographiques ainsi que differents patrons de croissance pour evaluer l'impact des series tronquees sur les variables de prediction et sur leur selection dans les analyses de regression logistique. De plus, nous evaluons la capacitedes modeles qui en resultent apredire avec exactitude le statut des arbres al'aide d'une valida- tion interne et externe. Nos resultats indiquent que les series de croissance tronquees des arbres vivants contribuent adi- minuer l'influence des facteurs externes sur les comparaisons de croissance. De ce fait, elles renforcent le lien entre la croissance et la vigueur dans le modele de mortaliteet ameliorent l'exactitude et l'applicabilitegenerale du modele. Par consequent, si les parametres du modele doivent etre integres dans des modeles de simulation aplus grande porteeg eogra- phique, les series tronquees peuvent etre utilisees pour augmenter la robustesse du modele. (Traduit par la Redaction)

Forest age structure and its spatial arrangement are important elements of sustainable forestry because of their effects on biodiversity and timber availability. Forest management objectives that include specific forest age structure may not be easily attained due to constraints imposed by the legacies of historical management and natural disturbance. We used a spatially explicit stochastic model to explore the synergetic effects of forest management and fire on boreal forest age structure. Specifically, we examined (1) the duration of spatial legacies of different management practices in the boreal forest, (2) how multiple shifts in management practices affect legacy duration and the spatial trajectories of forest age structure, and (3) how fire influences legacy duration and pattern development in combination with harvesting. Results based on 30 replicates of 500 years for each scenario indicate that (1) spatial legacies persist over 200 years and the rate at which legacies are overcome depends on whether new management targets are in synchrony with existing spatial pattern; (2) age specific goals were met faster after multiple management shifts due to the similar spatial scale of the preceding management types; (3) because large fires can erase the spatial pattern created by smaller disturbances, scenarios with fire had shorter lags than scenarios without fire. These results suggest that forest management goals can be accelerated by applying management at a similar spatial scale as existing spatial patterns. Also, management planning should include careful consideration of historical management as well as current and likely future disturbances.

Evaluating forest ecological integrity remains a major challenge for ecologists. We analyzed understory vegetation using an approach that combined plant functional types and vertical stratification to evaluate the effects of human disturbances on the ecological integrity of sugar maple-dominated stands in southern Quebec. Ecological integrity was evaluated by analyzing the divergence of understory species assemblages from those observed in comparable unmanaged forest. Multivariate analyses of biological traits revealed 13 emergent groups that share common traits associated with a similar life history strategy. Responses of these groups, of specific traits, and of understory structure to different human disturbances were tested. Nine of the 13 emergent groups varied in occurrence or diversity among disturbance types. Analyses also revealed a set of traits specifically associated with unmanaged old growth forest, indicating that species possessing these traits may be sensitive to human disturbance....

In forest communities, species coexistence can be favoured by disturbance-related variations in light regime coupled with rank reversal in species performance. The objective of this study was to determine if a major canopy disturbance, resulting from an ice storm, would favour the coexistence of sugar maple (Acer saccharum) and beech (Fagus grandifolia) in an old-growth forest located near the northern limit of the species' range. The growth, density and frequency of occurrence of understorey stems were evaluated 7 years after the ice storm and compared with pre-disturbance values to determine if sugar maple, a slightly less shade-tolerant species, would be favoured over beech. Although height and radial growth increased three- to five-fold, and > 70% of the 5-10 cm d.b.h. stems showed a release, sugar maple did not benefit more from the opening of the canopy than beech. The inability of sugar maple to outgrow beech might be related to the high proportion of beech root sprouts at our site, suggesting that the potential for the disturbancemediated mechanism to slow the competitive exclusion of maple might decrease as the importance of vegetative reproduction increases in beech. Prior to the disturbance, sugar maple was more abundant and occurred in a larger proportion of the plots among small size classes, while beech dominated in the larger size classes. Although some increases in the density of both species were detected after the disturbance, the general trends of relative abundance and distribution of the species were not modified significantly. Our results do not indicate that a single canopy disturbance such as the major 1998 ice storm significantly favoured sugar maple over beech. When a disturbance opens the canopy, competition for light in gaps may tend to maintain or reinforce a pre-existing hierarchy instead of reversing it, especially when interspecific differences in postdisturbance growth are small or absent, as in this study. This study shows that the dynamics of sugar maple - beech communities are more complex than previously envisaged and that revised models should incorporate additional factors (e.g. vegetative reproduction) that might also play an important role in regulating community dynamics.