Forests are expected to be strongly affected by modifications in climate and disturbance regimes, threatening their ability to sustain the provision of essential services. Promoting drought-tolerant species or functionally diverse stands have recently emerged as management options to cope with global change. Our study aimed at evaluating the impact of contrasting stand-level management scenarios on the resilience of temperate forests in eastern North America and central-western Europe using the individual process-based model HETEROFOR. We simulated the evolution of eight stands over 100 years under a future extreme climate according to four management scenarios (business as usual- BAU; climate change adaptation- CC; functional diversity approach- FD; no management- NM) while facing multiple disturbances, resulting in a total of 160 simulations. We found that FD demonstrated the greatest resilience regarding transpiration and tree biomass, followed by CC and then BAU, while these three scenarios were equivalent concerning the net primary production. These results were however dependent on forest type: increasing functional diversity was a powerful option to increase the resilience of coniferous plantations whereas no clear differences between BAU and adaptive management scenarios were detected in broadleaved and mixed stands. The FD promoted a higher level of tree species diversity than any other scenario, and all scenarios of management were similar regarding the amount of harvested wood. The NM always showed the lowest resilience, demonstrating that forest management could be an important tool to mitigate adverse effects of global change. Our study highlighted that tree-level process-based models are a relevant tool to identify suitable management options for adapting forests to global change provided that model limitations are considered, and that alternative management options, particularly those based on functional diversity, are promising and should be promoted from now on.

The fundamental trade-off between current and future reproduction has long been considered to result in a tendency for species that can grow large to begin reproduction at a larger size. Due to the prolonged time required to reach maturity, estimates of tree maturation size remain very rare and we lack a global view on the generality and the shape of this trade-off. Using seed production from five continents, we estimate tree maturation sizes for 486 tree species spanning tropical to boreal climates. Results show that a species' maturation size increases with maximum size, but in a non-proportional way: the largest species begin reproduction at smaller sizes than would be expected if maturation were simply proportional to maximum size. Furthermore, the decrease in relative maturation size is steepest in cold climates. These findings on maturation size drivers are key to accurately represent forests' responses to disturbance and climate change.

Process-based forest growth models with spatially explicit representation are relevant tools to investigate innovative silviculture practices and/or climate change effects because they are based on key ecophysiological processes and account for the effects of local competition for resources on tree growth. Such models are rare and are often calibrated for a very limited number of species and rarely for mixed and/or uneven-aged stands, and none are suitable for the temperate forests of Québec. The aim of this study was to calibrate and evaluate HETEROFOR (HETEROgeneous FORest), a process-based and spatially explicit model based on resource sharing, for 23 functionally diverse tree species in forest stands with contrasting species compositions and environmental conditions in southern Quebec. Using data from the forest inventory of Quebec, we evaluated the ability of HETEROFOR to predict the short-term growth (5–16 years) of these species at the tree and stand levels and the long-term dynamics (120 years) of red and sugar maple stands. The comparison between the prediction quality of the calibration and evaluation datasets showed the robustness of the model performance in predicting individual-tree growth. The model reproduced correctly the individual basal area increment (BAI) of the validation dataset, with a mean Pearson's correlation coefficient of 0.56 and a mean bias of 18 %. Our results also highlighted that considering tree position is of importance for predicting individual-tree growth most accurately in complex stands with both vertically and horizontally heterogeneous structures. The model also showed a good ability to reproduce BAI at the stand level, both for monospecific (bias of −3.7 %; Pearson's r=0.55) and multi-species stands (bias of −9.1 %; Pearson's r=0.62). Long-term simulations of red maple and sugar maple showed that HETEROFOR was able to accurately predict the growth (basal area and height) and mortality processes from the seedling stage to the mature stand. Our results suggest that HETEROFOR is a reliable option to simulate forest growth in southern Quebec and to test new forestry practices under future climate scenarios.

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands. A new method to quantify three masting components from individual tree-years has revealed that globally, masting is uncommon in tree species that depend on mutualist dispersers, with its distribution further mediated by climate and nutrient availability.

Aim Our understanding of the mechanisms that maintain forest diversity under changing climate can benefit from knowledge about traits that are closely linked to fitness. We tested whether the link between traits and seed number and seed size is consistent with two hypotheses, termed the leaf economics spectrum and the plant size syndrome, or whether reproduction represents an independent dimension related to a seed size-seed number trade-off.Location Most of the data come from Europe, North and Central America and East Asia. A minority of the data come from South America, Africa and Australia.Time period 1960-2022.Major taxa studied Trees.Methods We gathered 12 million observations of the number of seeds produced in 784 tree species. We estimated the number of seeds produced by individual trees and scaled it up to the species level. Next, we used principal components analysis and generalized joint attribute modelling (GJAM) to map seed number and size on the tree traits spectrum.Results Incorporating seed size and number into trait analysis while controlling for environment and phylogeny with GJAM exposes relationships in trees that might otherwise remain hidden. Production of the large total biomass of seeds [product of seed number and seed size; hereafter, species seed productivity (SSP)] is associated with high leaf area, low foliar nitrogen, low specific leaf area (SLA) and dense wood. Production of high seed numbers is associated with small seeds produced by nutrient-demanding species with softwood, small leaves and high SLA. Trait covariation is consistent with opposing strategies: one fast-growing, early successional, with high dispersal, and the other slow-growing, stress-tolerant, that recruit in shaded conditions.Main conclusions Earth system models currently assume that reproductive allocation is indifferent among plant functional types. Easily measurable seed size is a strong predictor of the seed number and species seed productivity. The connection of SSP with the functional traits can form the first basis of improved fecundity prediction across global forests.

Using nurse shrubs to improve tree seedling establishment in stressed environments is a common practice in forestry. Recent refinements of the stress-gradient hypothesis suggest that positive nurse effects occur under intermediate stress and decline in the harshest conditions. Additionally, indirect facilitation (e.g. protection from herbivory) is expected in low-stressed/productive systems according to the initial stress-gradient hypothesis. However, there have been fewer investigations into the use of nurse shrubs to decrease herbivore impacts in stressed systems compared with the role of nurses in stress alleviation. This raises the need to clarify conditions in which the use of nurse shrubs would be most appropriate. We conducted a field experiment in the sand dune forest of southwest France characterized by severe summer water shortages and the presence of different types of wild herbivores. In 2015 and 2016, Pinus pinaster seedlings were planted under shrubs Arbutus unedo or in shrub-removed locations, in fenced/unfenced areas. Survival, cause of mortality and water stress were monitored for 1 year after plantation. Summer drought was the main cause of mortality. Water stress increased in both years as summer progressed. We found direct facilitation only in July 2016 at intermediate water stress, due to a decrease in vapour pressure deficit under shrubs. These positive effects declined in late summer when stress was at a maximum. Rodent damage occurred under shrubs in spring whereas ungulate browsing was observed in open areas in autumn/winter. Overall seedling survival was equivalent under or outside shrubs during the first year, but was better under shrubs in the second one. Synthesis and applications. Our results partially validate the refinement of the stress-gradient hypothesis, with a decline of positive interactions in the most water-stressed conditions. This draws attention to the difficulty of using shrubs as nurses to improve tree regeneration in the most xeric systems. In addition, we show that indirect associational effects, both positive and negative, could be as important as abiotic stress alleviation in a stressful ecosystem, and using shrubs to facilitate seedling survival could be efficient depending on the type of herbivore involved. This highlights the fact that considering plant-plant interactions can help in the design of management options, providing that up-to-date ecological theories are considered and that the different biotic and abiotic constraints are accurately evaluated.

Stand regeneration is a crucial step in the management of many forests and its failure can jeopardize future forest growth and production. Thus, adapting forestry practices to improve seedling establishment is of prime importance to maintain sustainable forest management. In the coastal dune forests of maritime pine in SW France, regeneration failure after clear-cutting has greatly increased in the last decades. The aim of this study was to quantify the different stages involved in the regeneration process (seed rain, germination, survival), and to assess the impact of harvesting methods (partial cutting vs. clear-cutting) and of the use of direct seeding (seeding vs. no seeding) on these stages. We established five trials located in areas with contrasting regeneration statuses (of which two sites were in an area characterized by chronic regeneration failure), and we investigated the effect of the harvesting method and the use of direct seeding in a factorial design. We monitored the seed rain, germination and first-year survival for three years. Due to the transient nature of the seed bank, we found that the seed rain of the previous year was the only possible seed source for tree regeneration, and it increased with the proximity and size of surrounding mature trees. Nonetheless, seed rain did not limit regeneration in these stands. In fact, germination was the bottleneck stage of the regeneration process in all sites because it was short-lived and consequently exposed to failure in case of unfavourable conditions. Once established, seedling death was mainly due to summer drought. Clear-cutting without seeding resulted in severe regeneration failures in 4 out of 5 sites, with the most severe in the failure area. Direct seeding increased seedling emergence only in the first year, while partial cutting had a longer-lasting effect by providing a regular seed supply over several years. Partial cutting also decreased biotic and abiotic stresses through microhabitat modification under the tree canopy, resulting in a higher number of germinations and greater seedling survival. This positive effect was more pronounced in the two sites within the failure area, suggesting that conditions were more stressful in this part of the forests. Consequently, we recommend avoiding clear-cutting in favour of partial cutting in all parts of these forests, and ensuring the maintenance of sufficient forest cover to promote regeneration. Direct seeding could be used in addition to partial cutting to maximize the chances of success, but only in areas where natural regeneration is low.

Symbiosis with ectomycorrhizal (ECM) fungi can be important for regeneration success. In a context of increasing regeneration failures in the coastal forest of maritime pine in Southwest France, we tried to identity whether differences in ECM communities could partly explain the variation of regeneration success and how they are influenced by forest practices and stand characteristics. In particular, we focused on the effects of harvesting methods (comparing mature forest with seed-tree regeneration and clear-cuts) and topography (bottom-, mid-, and top positions). Five field trials (two in regeneration failure areas and three in successful areas) were used to sample 450 one-year-old seedlings. Assessments of ECM of seedling nutrient concentrations and of seedling growth based on exploration types were made. ECM root colonisation was similar in all harvesting treatments, suggesting that enough inoculum remained alive after logging. Harvesting-induced effects modifying soil properties and light availability respectively impacted ECM composition and seedling growth. Topography-induced variations in water and nutrient availability led to changes in ECM composition, but had little impact on seedling growth. Contact, short-distance, and long-distance exploration types improved the nutritional status of seedlings (Ca, K, and N), showing that mycorrhization could play an important role in seedling vitality. However, neither ECM root colonisation nor exploration types could be related to regeneration failures.

Les forêts dunaires d’Aquitaine de pin maritime (Pinus pinaster) représentent un intérêt économique et écologique important de par leur rôle multifonctionnel (production de bois,protection contre l’érosion, préservation de la biodiversité, tourisme). La régénération naturelle est pratiquée après coupe rase pour les renouveler. Mais, de nombreux échecs de régénération sont observés depuis 20 ans, entrainant des pertes économiques non négligeables.L’objectif de ce travail a été d’identifier les mécanismes écologiques impliqués dans ces échecs récurrents, afin de proposer des recommandations de gestion pour favoriser le renouvellement de ces pinèdes. Pour cela, nous avons étudié quatre étapes-clés du processus de régénération : dispersion des graines, germination, survie, et croissance. Un réseau de sites d’observation le long du cordon dunaire a été utilisé pour quantifier la régénération, en interactions avec des pratiques sylvicoles (coupe rase vs coupe progressive, semis de graines de pin maritime vs absence de semis). Deux sites semi-expérimentaux ont été installé afin d’étudier en détail certains facteurs explicatifs potentiellement importants dans notre contexte(climat, interactions plante-plante, herbivorie). Nos résultats confirment que la régénération naturelle est un processus complexe, influencé par de nombreux facteurs environnementaux et d’autres liés à la gestion, pouvant être eux mêmes en interaction. La quantité de graines atteignant le sol ne semble pas être limitante. La germination des graines est fortement liée à l’humidité du sol, et est influencée par l’hétérogénéité de la microtopographie. La sécheresse estivale, cause majeure de mortalité des plantules, peut-être modulée par le maintien d’arbres semenciers. Les interactions biotiques directes et indirectes avec la végétation de sous-bois impactent fortement la survie des plantules, mais varient grandement en direction et en intensité selon les saisons et les années.Bien que nos résultats permettent de mieux comprendre les mécanismes impliqués dans le cycle de régénération du pin maritime, il apparaît difficile de mettre en avant un seul facteur expliquant les échecs de régénération sur certains secteurs. Cependant, dans l’ensemble, le maintien d’arbres semenciers pendant quelques années permet d’obtenir une régénération réussie sur la globalité des forêts dunaires, grâce à l’apport pluriannuel de graines mais aussi à travers les modifications du microclimat sous leur canopée. Contrôler la végétation spontanée afin de réduire la compétition avec les plantules de pin, et augmenter les plans de chasse afin de réduire la pression des herbivores sur les plantules, doit aussi être envisagé dans les secteurs où les échecs de régénération sont fréquents.