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In a context where the disturbances could be more severe and/or more frequent, the choice of silvicultural management to be implemented following disturbances comes up against the lack of knowledge of the spontaneous dynamics of regeneration obtained in the context of climate change. From permanent plots installed in France following the storms of 1999, the objective of this study is to analyse the medium-term vegetation dynamics observed within forest gaps, to identify its main determinants and assess the interaction between disturbances and adaptation of plant communities to climate change. The natural tree regeneration, obtained in the post-storm gaps, seems to be sufficient in terms of density and species diversity. Most sites have in average more than 2000 stems ha-1 and 4.4 species present in the stratum greater than 2 m. The few concerning situations have been identified from the young stages of regeneration and have common characteristics: they are old coniferous stands on acidic soil where competing vegetation has been able to develop, inducing regeneration blocking. In these contexts, a silvicultural intervention would have been desirable to promote regeneration. À specific study of the natural regeneration of three major European hardwood species was carried out: oak, hornbeam and beech. We have highlighted a decline in the abundance of oak over time, in favour of the two most competitive species of beech and hornbeam. Oak is still present on 22% of the sites 19 years after the canopy was opened despite its low density. It also shows growth equivalent or even superior to that of hornbeam and beech, suggesting different development strategies between the three species. Nevertheless, in view of the observed dynamics of the oak, without intervention to promote its regeneration, its future seems uncertain in the coming decades. Finally, our study revealed a greater thermophilisation of the plant communities present in forests that have had an open canopy compared to those in undisturbed forests. By stimulating thermophilisation, disturbance thus plays a major ecological role in the adaptation of plant communities to climate change. The importance of large-scale disturbances should therefore not be underestimated, especially in a future climate context where they are likely to intensify.; Dans un contexte où les perturbations sont prédites plus sévères et/ou plus fréquentes, le choix des itinéraires sylvicoles à mettre en œuvre à la suite de perturbations se heurte à la méconnaissance de la dynamique spontanée de la régénération obtenue en contexte de changement climatique. À partir d’un réseau de placettes permanentes installées en France après les tempêtes de 1999, l’objectif de cette étude est d'analyser la dynamique de la végétation de moyen terme observée au sein de trouées forestières, d’identifier ses principaux déterminants et d’évaluer l’interaction entre perturbations et adaptation des communautés végétales au changement climatique. La régénération naturelle ligneuse, obtenue dans les trouées post-tempête, semble être satisfaisante en termes de densité et de diversité en essences, la majorité des sites ayant une densité de plus de 2000 tiges ha-1 avec en moyenne 4,4 espèces présentes dans la strate supérieure à 2 m. Les quelques situations préoccupantes ont pu être identifiées dès les jeunes stades de la régénération et regroupent des caractéristiques communes : ce sont des anciens peuplements résineux, sur sol acide où une végétation compétitrice s’est développée, induisant un blocage de la régénération. Dans ces contextes, une intervention sylvicole aurait été souhaitable pour favoriser la régénération. Une étude plus spécifique de la régénération naturelle de trois grandes essences feuillues européennes a été menée : le chêne, le charme et le hêtre. Nous avons ainsi pu mettre en évidence un déclin de l’abondance du chêne au cours du temps, au profit des deux espèces plus compétitrices qui sont le hêtre et le charme. Bien qu’ayant une densité faible, le chêne reste tout de même présent sur 22 % des sites 19 ans après l’ouverture du couvert. Il présente également une croissance équivalente, voire supérieure, à celle du charme et du hêtre, ce qui suggère des stratégies de développements différentes entre les trois essences. Néanmoins, au vu de la dynamique observée du chêne, sans intervention pour favoriser sa régénération, son avenir semble incertain dans les décennies à venir. Enfin, notre étude a permis de mettre en évidence une thermophilisation plus importante des communautés végétales présentes au sein des forêts ayant connues une période d’ouverture du couvert par rapport à celles des peuplements non perturbés. En stimulant la thermophilisation, la perturbation joue donc un rôle écologique majeur dans l’adaptation des communautés végétales au changement climatique. L’importance des perturbations de grande ampleur ne doit donc pas être sous-estimée, en particulier dans un contexte climatique futur où celles-ci risquent de s’intensifier.

Le paysage est devenu le fil rouge de la charte du Parc naturel régional du Morvan 2020-2035. Les parties prenantes ont établi le double constat que considérer l’avenir d’un paysage forestier oblige à prendre en compte de façon systémique les activités qui se déploient sur ce territoire et surtout que le paysage peut être un excellent levier de motivation et de médiation entre acteurs. Par un traitement des attentes culturelles ou sociétales, l’enrichissement et la validation du projet par la population sont appelés à faire vivre le paysage. Un paysage doit fonctionner dans ses composantes écologiques, socioéconomiques et culturelles. Si l’évolution de ses composantes s’opère de manière logique et compréhensible dans un certain « état du monde », alors l’acceptation de cette évolution est facile. Le paysage ne peut plus être « la partie d’un pays que la nature présente à un observateur ». Dans ce découpage objet-sujet, les paysages forestiers du Morvan nous offrent parfois un spectacle de désolation, de vide. Ce couplage sujet-objet propre à la définition occidentale du paysage pourrait laisser place à une vision issue de la pensée chinoise. Nous verrons comment la production de catégories émergentes à partir d’enceintes de concertation territorialisée permet de faire travailler ensemble les acteurs autour de ce qui peut être préservé en commun, le vivre de paysage. L’issue à ces antagonismes se trouve dans la construction d’une vision partagée de l’avenir du paysage du massif. Nous étayerons nos propos grâce aux nombreux bilans et enquêtes réalisées lors de la révision de la charte. The landscape has become the red thread of the 2020-2035 Charter for the Morvan Regional Nature Park. Stakeholders realise that in considering the future of a forest landscape they need to systemically take into account the activities deployed within it and, above all, to recognise the landscape as an excellent lever for motivation and mediation between actors. By addressing cultural or societal expectations, enhancing the project, and gaining the population’s approval can ensure the landscape is experienced as a living space. A landscape must function in relation to its ecological, socio-economic, and cultural components. If these components evolve in a logical and comprehensible way in tune with a certain "state of the world”, their change is easily accepted. The landscape can therefore no longer be perceived as "the part of a region nature presents to an observer". From this object-subject perspective, the forest landscapes of the Morvan sometimes offer us a spectacle of desolation and emptiness. This coupling of subject and object which is specific to the Western definition of the landscape could be replaced by a vision derived from Chinese thought. We will see how the production of emerging categories based on regional consultation procedures can enable actors to work together on preserving the shared experience of the landscape. The solution to such oppositions lies in the construction of a shared vision of the future of the landscape of the Morvan Massif. We will back up the arguments put forward in this article with the many evaluations and surveys carried out during the revision of the charter.

Although fires are common disturbances in North American forests, the extent to which soil invertebrate assemblages recover from burning remains unclear. Here, we examine long-term (14- to 101-yr) recoveries of soil invertebrate communities from common cut and burn treatments conducted at 6 to 26-yr intervals since 1911 in a deciduous forest in the upper Great Lakes region (USA). We characterize soil surface macro-invertebrate communities during both fall and spring across a long-term, experimental fire chronosequence to characterize invertebrate community recovery at decadal time-scales and community changes between seasons. We posited that changes in invertebrate community structure might, in turn, impact decomposition process. We sampled active organisms at the soil surface using pitfall traps. We described understory vegetation, measured soil properties, and conducted a 4-year litter bag study with big-toothed aspen leaves (Populus grandidentata). Invertebrate community responses followed a habitat accommodation model of succession showing that invertebrate succession is dependent on the soil surface properties. The fall and spring measures revealed that the densities of active invertebrates were highest 101 years after fire. For a given pair of stands, a pattern of sharing higher percentage of taxa was denoted when stands were of similar age. Some species such as the beetle Stelidota octomaculata appeared to be indicator of the chronosequence succession stage because it tracks the successional increase of Quercus and acorn production at the study site. We also found a significant positive correlation between leaf decomposition of soil macrofaunal accessible leaves and millipedes density across the chronosequence. We show that vegetation cover changes and related shifts in habitat structure occurring during post-fire succession are important in shaping communities assemblages. This finding highlights the importance of simultaneously considering abiotic-biotic factors together with above- and belowground measurements to better characterize controls on successional community dynamics after disturbance.

Forest ecosystems are one of the main providers of terrestrial ecosystem services, whose functioning has already been altered by recent climate change. Increases in tree mortality rates have been highlighted in different biomes worldwide, as well as increases in the frequency of massive mortality events following droughts. However, tree mortality is a multi-causal process. It is difficult to quantify the importance of the different factors that can possibly lead to tree death, and particularly the importance of climate change in comparison with forest dynamics and competition-related effects, environmental or biotic factors. This thesis aims at assessing the drivers of background tree mortality, which is the mortality observed in a stand in the absence of extreme perturbation, for the main European tree species. We used data from the French forest inventory of the National Geographic Institute (IGN) and historic climate data from Météo-France spanning the years 1961 to 2015. First, we modelled background tree mortality for 43 tree species in order to identify the drivers of background tree mortality. We used 372.974 trees, including 7.312 dead trees surveyed between the years 2009 and 2015. We found that factors related with competition, tree development stage, stand structure and species composition and logging intensity explained 85% of the recent tree mortality. Environmental factors (soil and climate conditions) accounted for 9% of the total modelled mortality. Temperature increases and rainfall decreases since the period 1961 – 1987 had a significant effect on the mortality of 45% of the 43 species and explained in average 6% of the total modelled mortality.Secondly, we focused on the link between trees locations along temperature and rainfall gradients and their sensitivity to changes of temperature and rainfall. We found that, for 9 species out of 12, temperature increases and rainfall decreases effects were more important in areas with high mean temperature and low mean rainfall. These results show that climate change-related tree mortality could be exacerbated towards the species’ warm and dry edges.Finally, we sought to evaluate how climate change-related tree mortality varied along trees social statuses and sizes gradients. We found that suppressed trees were more sensitive to temperature increases than dominant trees. On the contrary, dominant trees, and particularly large dominant trees, appear to be more sensitive to rainfall decrease than suppressed trees. Overall, our results show that climate change-related tree mortality is globally more important for suppressed than dominant trees.We highlighted the existence of a link between recent temperature increases and rainfall decreased and observed tree mortality rates on around half of the species of the French forest. We also showed that these effects were exacerbated towards the warm and dry edges of the species ranges. Finally, we showed that these effects differed according to trees social statuses and development stages. These results allow us to better understand the impacts of climate change on French and European forest and to better anticipate their effects through the adaptation of silvicultural practices.; Les écosystèmes forestiers sont l’un des principaux fournisseurs de services écosystémiques terrestres, dont le changement climatique récent a déjà altéré le fonctionnement. Des augmentations des taux de mortalité ont été mis en évidence à différents endroits du globe, de même que des évènements de mortalité massifs attribués à des sécheresses de plus en plus fréquentes et intenses. Toutefois, la mortalité d’un arbre est un phénomène multifactoriel. Il est difficile d’évaluer l’importance relative des différents facteurs pouvant conduire à la mort d’un arbre, et notamment l’importance du changement climatique par rapport aux facteurs liés à la dynamique forestière, aux facteurs environnementaux ou encore biotiques. Cette thèse porte sur l’étude des déterminants de la mortalité de fond, qui est la mortalité observée dans un peuplement en l’absence de perturbation extrême, pour les principales espèces forestières européennes. L’ensemble de l’étude a été réalisée à partir des données de l’inventaire forestier de l’Institut national de l'information géographique et forestière (IGN) et de données climatiques de Météo France couvrant la période 1961 - 2015. Dans un premier temps, nous avons modélisé les déterminants de la mortalité de fond pour 43 espèces d’arbres représentatives des forêts tempérées européennes. Nous avons utilisé les relevés de 372.974 arbres dont 7.312 arbres morts. Les facteurs liés à la compétition, au stade de développement des arbres, à la structure et composition du peuplement et à l’intensité des travaux sylvicoles expliquent 85% de la mortalité récente modélisée, et les facteurs environnementaux (sol et conditions climatiques moyennes) 9%. Les augmentations de températures et baisses de précipitations entre la période contemporaine du relevé et la période 1961 – 1987 jouent un rôle pour 45% des espèces, expliquant 6% de la mortalité modélisée. Ensuite, nous nous sommes intéressés au lien entre la position des arbres le long des gradients environnementaux (température et précipitations) et la sensibilité aux changements de températures et de régimes de précipitations. Nous avons mis en évidence, pour 9 espèces sur 12, une exacerbation de la mortalité due aux augmentations de température et baisses de précipitations récentes lorsque celles-ci sont combinées à des conditions climatiques moyennes chaudes et sèches. Ces résultats mettent en évidence une vulnérabilité exacerbée pour les arbres évoluant vers les limites chaudes et sèches de leur aire de distribution. Enfin, nous avons cherché à évaluer le lien entre la sensibilité au changement climatique et le statut social et la taille moyenne des arbres. Nos résultats indiquent que les arbres dominés sont plus sensibles aux augmentations de températures que les arbres dominants. A l’inverse, les arbres dominants apparaissent plus sensibles aux baisses de précipitations, avec un effet maximal atteint pour les arbres dominants de gros diamètre. Globalement, nos résultats indiquent que la mortalité due aux augmentations de températures ou aux diminutions de précipitations est nettement plus importante pour les arbres dominés que pour les dominants, et que le statut social de l’arbre joue un rôle plus important que la taille de l’arbre.Ces travaux ont permis de mettre en évidence l’existence d’un lien entre les augmentations de températures, les baisses de précipitations récentes et les taux de mortalité observés en forêt sur près de la moitié des espèces arborées de la forêt française. Ils ont également permis de démontrer que ces effets étaient exacerbés sur les arbres évoluant à proximité des limites chaudes de leur aire de distribution. Enfin, ils ont permis d’évaluer les différences de sensibilité au changement climatique en fonction du statut social et de la taille des arbres. Ces résultats permettent de mieux comprendre l’impact des changements climatiques sur les forêts françaises et européennes, contribuant à mieux anticiper leurs effets.

Engineering school; In a context of the increasing pressure of wild ungulates on the timber industry, silviculture must adapt to ensure a good renewal of forest stands. We have studied the responses of sessile oak (Quercus petraea) seedlings and adventitious vegetation to herbivory pressure of the main wild ungulates of the plains (Cervus elaphus, Capreolus capreolus and Sus scrofa). We followed a cohort of sessile oak seedlings from 2018 and studied its behaviour according to two main parameters. The presence or absence of predation by ungulates through a device of enclosures and exclosures. The edaphic modalities and the presence or absence of competing for vegetation thanks to two working methods: a control modality without intervention and a so-called VEGHYDR modality on which the grasses have been removed, the soil decompaction and furrows dug. We have shown that during the first two growing seasons, the main damage is due to wild boar populations. They represent 12% of seedling damage (2nd degradation factor after slugs and phytophagous insects) and the leading cause of death (40%). Consequences of their high consumption of acorns and cotyledons and the selection of a population unfavourable to forestry. Wild boars are opportunistic omnivores and attack the most vigorous acorns by neglecting the less palatable. As there is a relationship between acorn size, seedling growth and survival, this harms these last two parameters. When the cotyledons disappear, the impact of wild boars decreases. They are no longer consumed but can still betrampled or snatched by wild boars in search of food. The impacts of cervids are still absent. We also found that the survival rate of sessile oak seedlings increases with the density of competing for herbaceous vegetation. This goes against our original hypothesis. Finally, it emerged from this study that the morphology of the seedlings during their first growing seasons is mainly dependent on the quality of the acorns rather than the biotic and abiotic variables studied. The results of our study are consistent with the literature except for the positive correlation observed between seedling survival rate and competing for vegetation density. In the next years of monitoring the device, we will see if this trend will be confirmed or reversed. Finally, it has become apparent that soil decompaction and drainage are detrimental to the establishment of the natural regeneration of sessile oak. This work is, therefore, to be avoided for this type of forest stand management. Besides, the economic value of placing fences around regeneration plots is not confirmed, since when excluding wild boar, the other causes of mortality increase considerably.; Dans un contexte de pression croissante des ongulés sauvages sur la filière bois, la sylviculture doit s’adapter afin de garantir un bon renouvellement des peuplements forestiers. Nous avons étudié les réponses des semis de chêne sessile (Quercus petraea) et de la végétation adventice face à la pression d’herbivorie des principaux ongulés sauvages de plaine (Cervus elaphus, Capreolus capreolus et Sus scrofa). Nous avons effectué le suivi d’une cohorte de semis de chênes sessiles de 2018 et étudié son comportement en fonction de deux grands paramètres. La présence ou l’absence de prédation par les ongulés grâce à un dispositif d’enclos et d’exclos. Les modalités édaphiques et la présence ou l’absence de végétation concurrente grâce à deux modalités de travail : une modalité témoin sans intervention et une modalité dite VEGHYDR sur laquelle les graminées ont été supprimé, le sol décompacté et des sillons creusés. Nous avons montré qu’au cours des deux premières saisons de végétation, les principaux dégâts sont dus aux populations de sangliers. Ils représentent 12% des dégradations des semis (2em facteur de dégradation après les limaces et les insectes phytophages) et la première cause de mortalité (40%). Conséquences de leur consommation importante de glands et de cotylédons et de la sélection d’une population peu favorable à la sylviculture. De fait les sangliers sont des omnivores opportunistes et s’attaquent aux glands les plus vigoureux en délaissant les moins appétents. Comme il existe une relation entre taille des glands, croissance et survie des semis, cela a un effet négatif sur ces deux derniers paramètres. Lorsque les cotylédons disparaissent, l’impact des sangliers diminue. Ils ne sont alors plus consommés mais peuvent encore être piétinés ou arrachés par des sangliers à la recherche de nourriture. Les impacts des cervidés sont eux encore absents. Nous avons également constaté que le taux de survie des semis de chêne sessile augmente avec la densité de la végétation concurrente herbacée. Ce qui va à l’encontre de notre hypothèse de départ. Enfin, il est ressorti de cette étude que la morphologie des semis au cours de leurs premières saisons de végétation est majoritairement dépendante de la qualité des glands plus que des variables biotiques et abiotiques étudiées. Bien que l’assainissement du sol soit également néfaste à la survie des semis du fait d’une diminution critique de la réserve en eau durant les périodes de sécheresse. Le taux de survie est en effet plus important sur nos modalités témoins sans intervention, donc sans travail ni assainissement du sol. Les résultats de notre étude sont conformes à la littérature à l’exception de la corrélation positive observée entre le taux de survie des semis et la densité de végétation concurrente. Dans les prochaines années de suivi du dispositif, nous verrons si cette tendance sera confirmée ou bien inversée. Enfin il est apparu que la décompaction et le drainage du sol sont néfastes à l’installation de la régénération naturelle du chêne sessile. Ces travaux sont donc à éviter pour ce type de gestion de peuplement forestier. De plus, l’intérêt économique de la mise en place de clôtures autour des placettes de régénération n’est pas confirmé, puisque lorsque l’on exclut le sanglier, les autres causes de mortalité augmentent considérablement.

International audience; Temperate forests are under climatic and economic pressures. Public bodies, NGOs and the wood industry are looking for accurate, current and affordable data driven solutions to intensify wood production while maintaining or improving long term sustainability of the production, biodiversity, and carbon sequestration. Free tools and open access data have already been exploited to produce accurate quantitative forest parameters maps suitable for policy and operational purposes. These efforts have relied on different data sources, tools, and methods that are tailored for specific forest types and climatic conditions. We hypothesized we could build on these efforts in order to produce a generic method suitable to perform as well or better in a larger range of settings. In this study we focus on building a generic approach to create forest parameters maps and confirm its performance on a test site: a maritime pine (Pinus pinaster) forest located in south west of France. We investigated and assessed options related with the integration of multiple data sources (SAR L- and C-band, optical indexes and spatial texture indexes from Sentinel-1, Sentinel-2 and ALOS-PALSAR-2), feature extraction, feature selection and machine learning techniques. On our test case, we found that the combination of multiple open access data sources has synergistic benefits on the forest parameters estimates. The sensibility analysis shows that all the data participate to the improvements, that reach up to 13.7% when compared to single source estimates. Accuracy of the estimates is as follows: aboveground biomass (AGB) 28% relative RMSE, basal area (BA) 27%, diameter at breast height (DBH) 20%, age 17%, tree density 24%, and height 13%. Forward feature selection and SVR provided the best estimates. Future work will focus on validating this generic approach in different settings. It may prove beneficial to package the method, the tools, and the integration of open access data in order to make spatially accurate and regularly updated forest structure parameters maps effortlessly available to national bodies and forest organizations.

Partie 3 : Douglas-fir ecology; International audience

Partie 3 : Douglas-fir ecology; International audience

Forests are complex three-dimensional ecosystems, but little is known about the influence of vertical stratification of forest structure on biodiversity and species turnover. Saproxylic beetles make a substantial contribution to forest biodiversity and ecosystem functions such as nutrient cycling. Management measures aimed at supporting saproxylic biodiversity are becoming an integral part of sustainable forest management practices. Yet, monitoring is carried out under the assumption that saproxylic activity at ground level will be a realistic reflection of saproxylic biodiversity in the forest as a whole. To investigate the validity of this assumption we compare vertical stratification and composition of saproxylic beetle assemblages between three forest types of varying altitude and latitude, including a tropical lowland forest in Panama, a temperate lowland forest and a temperate montane forest, both in the eastern Czech Republic. Beetles were sampled following a standardized sampling protocol using flight intercept traps arranged in vertical transects. Overall, the tropical forest was estimated to harbour two to three times more saproxylic beetle species than the temperate lowland and the montane forest, respectively. However, point richness estimates within vertical strata were remarkably similar between biomes. Species richness was similar in the understorey of all three forests. It peaked in the canopy of the tropical forest but in the understorey of temperate forests. So, while the beetle assemblages were clearly vertically stratified in all three forest types, stratification patterns varied markedly between tropical and temperate forests. This trend is driven primarily by the high richness of saproxylic beetles in the tropical forest canopy. However these richness differences belie the strong similarities in stratification of feeding guild composition observed all three forest types. This would tend to suggest that similar trophic structuring forces might operate across very different forest biomes of the world. Similarities in feeding guild composition suggest that management measures aiming at conserving biodiversity of saproxylic beetles are likely to be effective across different forest types. The differences in vertical stratification, however, suggest that understorey monitoring alone will be insufficient to understand management effects on saproxylic biodiversity. In the tropics, in particular, more emphasis will need to be placed on managing the diverse, three-dimensional structure of forest canopies as habitat for saproxylic beetles.

One of the main usages of wood in Europe is renewable energy supply that implies intensification of forest management to respond to this increasing demand. However, the impact of intense forestry practices on soil microbial communities remains poorly investigated. In the frame of my PhD thesis, I evaluated effects of artificial organic matter removal on functional and taxonomical diversity of soil bacterial and fungal communities in temperate forest, using six experimental sites across France (INRA MOS experimental network). In parallel, I also characterised impact of intensified forest management practices on functional microbial communities in tropical plantation of Eucalyptus trees. This work permitted to identify several sensitive functional indicators of organic matter degradation. Notably, the degradation of chitin – a nitrogen polymer main component of arthropods and fungal cell walls – was revealed to be particularly sensitive to organic matter removal. Genomics and enzymatic approaches were then used to estimate chitinolytic potentials of the different genera of soil fungi. In controlled conditions, we were able to quantify ectomycorrhizal fungus carbon and nitrogen mobilisation and transfer capacities from chitin enriched organic matter to its host during symbiotic interaction. Finally, we evaluated chitinolytic functions of ectomycorrhizal fungi at large scale by combining enzymatic and isotopic approaches. Taken together, the results acquired in the frame of my PhD thesis, illustrate the significant role of ectomycorrhizal fungi in carbon and nitrogen mobilisation from organic matter. We particularly highlight that microbial compartment in soil must be considered in studies of forest management practices; En Europe, le bois est la première source d’énergie renouvelable. La transition énergétique se traduit par une intensification de l’exploitation des forêts. L’effet de ces pratiques sylvicoles sur les communautés microbiennes du sol est encore peu étudié. Au cours de ma thèse, j’ai évalué les conséquences d’une manipulation artificielle de matière organique en forêt tempérée sur la diversité fonctionnelle et taxonomique des communautés bactériennes et fongiques telluriques dans six sites expérimentaux (réseau expérimental MOS). Parallèlement, une caractérisation fonctionnelle des communautés microbiennes a également été réalisée dans un contexte proche des réalités de l’intensification des pratiques sylvicoles sous climat tropical en plantation d’Eucalyptus. Si certains descripteurs fonctionnels de la dégradation de la matière organique sont particulièrement informatifs, les activités microbiennes de dégradation de la chitine, polymère azoté des arthropodes et champignons, sont apparues très sensibles au retrait de matière organique. C’est pourquoi, par des approches de génomiques comparatives, nous avons cherché à estimer le potentiel chitinolytique des différentes guildes fongiques des sols. En conditions contrôlées, nous avons ensuite quantifié les capacités potentielles de mobilisation et de transfert du carbone et de l’azote, à partir d’une matière organique microbienne riche en chitine, par un champignon ectomycorhizien en symbiose avec son hôte. Enfin, la généricité des fonctions chitinolytiques d’un plus large spectre d’espèces fongiques ectomycorhiziennes a été évaluée par le couplage d’approches enzymatiques et isotopiques. L’ensemble de nos résultats met en lumière le rôle significatif des champignons ectomycorhiziens dans la mobilisation du carbone et de l’azote à partir de certaines formes de matière organique, et la nécessité de prendre en compte le compartiment microbien dans les études d’impact des pratiques sylvicoles

Eddy covariance (EC) measurements are widely used to estimate the amount of carbon sequestrated by terrestrial biomes. The decision to exclude an EC flux from a database (bad quality records, turbulence regime not adequate, footprint problem,...) becomes an important step in the CO2 flux determination procedure. In this paper an innovative combination of existing assessment tests is used to give a relatively complete evaluation of the net ecosystem exchange measurements. For the 2005 full-leaf season at the Hesse site, the percentage of rejected half-hours is relatively high (59.7%) especially during night-time (68.9%). This result strengthens the importance of the data gap filling method. The data rejection does not lead to a real improvement of the accuracy of the relationship between the CO2 fluxes and the climatic factors especially during the nights. The spatial heterogeneity of the soil respiration (on a site with relatively homogenous vegetation pattern) seems large enough to mask an increase of the goodness of the fit of the ecosystem respiration measurements with a dependence on soil temperature and water content when the tests are used to reject EC data. However, the data rejected present some common characteristics. Their removal lead to an increase in the total amount of CO2 respired (24%) and photosynthesised (16%) during the 2005 full-leaf season. Consequently the application of our combination of multiple quality tests is able improve the inter-annual analysis. The systematic application on the large database like the CarboEurope and FLUXNET appears to be necessary.

For at least the past two centuries, human activities have caused strong environmental changes in the biosphere. Many studies have shown responses of vegetation to global changes. However, many unknowns remain. First, most explicitly temporal studies have been conducted at a single site with a common intensity of environmental changes and historical land-use legacies. Results are highly variable among studies, and we have a very limited understanding of mechanisms underlying this variation. Second, despite the major contribution of bryophytes to ecosystem functioning, very few temporal studies have focused on bryophytes.This Ph.D. contributes to filling these two knowledge gaps. The overarching question for the three research projects presented here is: what is the impact of environmental change on biodiversity? We built a set of hypotheses around two main questions: (i) What is the effect of environmental changes on forest vegetation? (ii) Which taxon, bryophytes or vascular plants, is most sensitive to global changes? Chapter 2 tests the hypothesis that bryophytes are more sensitive than vascular plants to the combination of atmospheric deposition and warming in an industrial region in north-eastern France. Chapter 3 tests the hypothesis that forest vegetation changes have been greatest in regions with the strongest warming trends along a continental gradient in eastern Canada. The last chapter combines the two first approaches, quantifying temporal changes in bryophyte and vascular plant communities in sites with different warming intensities along elevational gradients in eastern Canada. To answer to these questions, I used an historical ecological approach by resurveying botanical plots initially surveyed in the 1970s. Plot selection followed a reproducible and detailed procedure to minimize confounding factors. Our results show a direct effect of global changes on forest vegetation. First, bryophytes appear more sensitive to atmospheric deposition than vascular plants (Chapter 2). Second, temporal changes in vascular plant communities were stronger in areas where warming has been greatest (Chapter 3). Third, in response to warming, changes in bryophyte and vascular plant communities show idiosyncratic differences, depending on the community property under study (Chapter 4). Results of the three chapters clearly show systematic changes in community composition, that are not necessarily accompanied by changes in local diversity. In sum, we provide empirical evidence that historical ecology is a powerful method to disentangling mechanisms of vegetation response to global changes. Only a holistic approach based on different biodiversity components, different spatial scales and wide variety of community properties permit an understanding of the complexity of temporal dynamics of vegetation., Depuis environ deux siècles les activités humaines modernes ont profondément modifié les conditions environnementales sur la surface de la Terre. De nombreuses études ont mis en évidence une réponse de la végétation face à ces changements. Cependant, il persiste plusieurs incompréhensions. Premièrement, les réponses des communautés varient fortement entre les études, et les mécanismes responsables de cette variation sont encore mal connus. Deuxièmement, malgré leur importance écologique, les bryophytes restent largement sous étudiées dans les études temporelles, limitant notre compréhension de leur dynamique temporelle. Ce doctorat a pour objectif d’apporter des éléments de réponses à ces deux points. Les trois projets de recherche gravitent autour de la grande question : quels sont les effets des changements environnementaux sur la biodiversité? Les hypothèses que j’ai développées se construisent autour de ces deux grandes questions (i) Quels sont les effets des dépositions et du réchauffement de la température sur la végétation forestière? (ii) Il y a-t-il une différence de sensibilité entre bryophytes et trachéophytes face aux changements environnementaux? Le chapitre 2 teste l’hypothèse que les bryophytes sont plus sensibles que les plantes vasculaires face aux dépositions atmosphériques et au réchauffement de la température dans une région industrielle du nord-ouest de la France. Le chapitre 3 teste les mécanismes de réponse de la végétation forestière le long d’un gradient de réchauffement climatique dans l’est du Canada. Enfin, le chapitre 4 est une approche mixe entre les deux premiers chapitres, il teste la réponse des plantes vasculaires et des bryophytes face à différentes intensités de réchauffement de la température sur des gradients altitudinaux dans l’est de la province de Québec. Dans les trois chapitres, j’ai utilisé les méthodes de l’écologie historique. Après un long travail d’archive, j’ai rééchantillonné des relevés botaniques faits dans les années 1970. La sélection des sites à rééchantillonner suit un protocole finement détaillé afin de minimiser tous effets confondants. Les résultats des trois chapitres mettent en évidence le lien direct entre réponse de végétation et changements environnementaux. Premièrement, les bryophytes sont plus sensibles aux dépositions atmosphériques que les plantes vasculaires (chapitre 2). Secondement, les changements temporels de la végétation vasculaire sont plus grands dans les zones ou le réchauffement climatique fut le plus fort (chapitre 3). Dernièrement, face au réchauffement climatique, les réponses des bryophytes et plantes vasculaires diffèrent selon la propriété de la communauté qui est étudiée (chapitre 4). Les trois chapitres, montrent un changement systématique de la composition des communautés, sans pour autant de changement de la richesse spécifique. Ce doctorat fournit trois exemples de la force des méthodes historiques dans la compréhension des mécanismes de réponse de la végétation face aux changements globaux. Mes travaux supportent l’importance d’analyser la dynamique de la végétation avec une vision holistique. La compréhension des mécanismes liés à la dynamique temporelle de la végétation doit passer par l’étude de plusieurs groupes taxonomiques, avec différentes propriétés des communautés sur plusieurs échelles spatiales.

(IF 3.13; Q1); Numerous studies predict a short-term important decrease in fossil resources and stress the need to develop alternative renewable energies, thus European countries aim at increasing biomass production for energetic purposes.One such source of bioenergy could be obtained from forest biomass pools by exporting logging residues, yet this practice would have strong impacts on forest ecosystems (e.g. disturbance of soil biodiversity and chemical properties). Most studies on biomass removal effects focused on boreal forests but responses in temperate forests are still scarcely studied. Soil macrofauna is involved in forest ecosystem functioning through numerous chemical, physical and biological processes and multiple interactions with other organisms. A disturbance of the soil macrofauna community can thus lead to a response of the forest ecosystem as a whole.Experimental plots were set up in 6 northern France forests (either dominated by oak or beech), with a treatment corresponding to Whole Tree Harvesting (WTH) practices compared to control, in order to characterize the response of soil macrofaunal communities to organic matter (OM) export. Our study showed that OM export leads to a short-term loss of abundance in macrofaunal communities in temperate deciduous forest soils that can also lead to an alteration of soil OM cycle, and revealed that the breadth of this negative impact is modulated by tree species and faunal trophic groups. Furthermore, some results of community responses were directly applicable to forest management as marked responses of several taxa revealed four bioindicators of OM disturbance.Therefore, the level of disturbance related to new forest management practices, such as WTH to increase forest biofuel yields, (i) could be characterized by surveying either total soil macrofauna or using tools such as bioindicator taxa and (ii) should be assessed regarding tree stand identity.

Mediterranean forests are found in the Mediterranean basin, California, the South African Cape Province, South and southwestern Australia and parts of Central Chile. They represent 1.8 % of the world forest areas of which the vast majority is found in the Mediterranean basin, where historical and paleogeographic episodes, long-term human influence and geographical and climatic contrasts have created ecosystemic diversity and heterogeneity. Even if evergreen is dominant, deciduous trees are also represented, with different forest types including dense stands with a closed canopy (forests sensu stricto) and pre-forestal or pre-steppic structures with lower trees density and height. The Mediterranean basin is also a hot spot of forest species and genetic diversity, with 290 woody species versus only 135 for non-Mediterranean Europe. However, the characteristics of the Mediterranean area (long-standing anthropogenic pressure, significant current human activity and broad biodiversity) make it one of the world’s regions most threatened by current changes. Four examples of Mediterranean forest types, present in south and north of the Mediterranean basin and more or less threatened, are developed in order to show that linking “hard sciences” and humanities and social sciences is necessary to understand these complex ecosystems. We show also that these forests, in spite of specific climatic constraints, can also be healthy and productive and play a major ecological and social role. Furthermore, even if the current human activity and global change constitute a risk for these exceptional ecosystems, Mediterranean forests represent a great asset and opportunities for the future of the Mediterranean basin.

Generally, the effects of climate change on tree growth focus on changes in one dimension of a tree. However, diameter increment along the main stem reacts differently to climatic variables, which in turn influences tree form. These differences can thus have important implications on stem volume, which could induce biases in future forest biomass estimation. A stem taper model including climatic variables was fitted to stem analysis data of five different species (Abies balsamea, Betula papyrifera, Picea glauca, Picea mariana, Populus tremuloides) distributed along a gradient from the temperate to the boreal forest of Eastern Canada. The effects of shifts in stem form on tree volume between different climatic scenarios were then estimated and related to different functional traits. Changes in stem form with climatic variables were observed for four of the five species, with up to 5% differences in stem volume between different climatic situations. Changes in stem volume were found to decrease with increasing waterlogging and shade tolerance. The most important differences in stem volume can induce changes of 3–4% in the biomass of a single tree. Not taking into account shifts in stem form could have implications in forest biomass estimations.

UMR SYSTEM : équipe SYME; Background and aims :Plant phenology is a sensitive indicator of plant response to climate change. Observations of phenological events belowground for most ecosystems are difficult to obtain and very little is known about the relationship between tree shoot and root phenology. We examined the influence of environmental factors on fine root production and mortality in relation with shoot phenology in hybrid walnut trees (Juglans sp.) growing in three different climates (oceanic, continental and Mediterranean) along a latitudinal gradient in France.Methods : Eight rhizotrons were installed at each site for 21 months to monitor tree root dynamics. Root elongation rate (RER), root initiation quantity (RIQ) and root mortality quantity (RMQ) were recorded frequently using a scanner and time-lapse camera. Leaf phenology and stem radial growth were also measured. Fine roots were classified by topological order and 0–1 mm, 1–2 mm and 2–5 mm diameter classes and fine root longevity and risk of mortality were calculated during different periods over the year.Results :Root growth was not synchronous with leaf phenology in any climate or either year, but was synchronous with stem growth during the late growing season. A distinct bimodal pattern of root growth was observed during the aerial growing season. Mean RER was driven by soil temperature measured in the month preceding root growth in the oceanic climate site only. However, mean RER was significantly correlated with mean soil water potential measured in the month preceding root growth at both Mediterranean (positive relationship) and oceanic (negative relationship) sites. Mean RIQ was significantly higher at both continental and Mediterranean sites compared to the oceanic site. Soil temperature was a driver of mean RIQ during the late growing season at continental and Mediterranean sites only. Mean RMQ increased significantly with decreasing soil water potential during the late aerial growing season at the continental site only. Mean root longevity at the continental site was significantly greater than for roots at the oceanic and Mediterranean sites. Roots in the 0–1 mm and 1–2 mm diameter classes lived for significantly shorter periods compared to those in the 2–5 mm diameter class. First order roots (i.e. the primary or parents roots) lived longer than lateral branch roots at the Mediterranean site only and first order roots in the 0–1 mm diameter class had 44.5% less risk of mortality than that of lateral roots for the same class of diameter.Conclusions :We conclude that factors driving root RER were not the same between climates. Soil temperature was the best predictor of root initiation at continental and Mediterranean sites only, but drivers of root mortality remained largely undetermined.

Climate change affects ecosystem functioning directly through impacts on plant physiology, resulting in changes of global productivity. However, climate change has also an indirect impact on ecosystems, through changes in the composition and diversity of plant communities. The relative importance of these direct and indirect effects has not been evaluated within a same generic approach yet. Here we took advantage of a novel approach for disentangling these two effects in European temperate forests across a large climatic gradient, through a large simulation-based study using a forest succession model. We first showed that if productivity positively correlates with realized tree species richness under a changed climate, indirect effects appear pivotal to understand the magnitude of climate change impacts on forest productivity. We further detailed how warmer and drier conditions may affect the diversity-productivity relationships (DPRs) of temperate forests in the long term, mostly through effects on species recruitment, ultimately enhancing or preventing complementarity in resource use. Furthermore, losing key species reduced the strength of DPRs more severely in environments that are becoming climatically harsher. By disentangling direct and indirect effects of climate change on ecosystem functioning, these findings explain why high-diversity forests are expected to be more resilient to climate change.

Background and aims: In temperate conifer forests, soil temperature is an important driver of fine root growth and winter root growth can occur during aerial dormancy. We hypothesize that in conifers, stocks of non-structural carbohydrates (NSC) in fine roots are high enough to provide energy for root growth and production throughout the year, even when photosynthesis is reduced. Methods: We measured monthly root production (i.e. the number of roots undergoing elongation) and their elongation rate (RER) in mature Picea abies for one year, along a soil temperature gradient (three altitudes of 1400, 1700 and 2000 m). Every two months we harvested needles, branches, stem, large, medium and fine roots, and quantified starch and soluble sugars in each organ using analytical methods and near infra-red spectroscopy (NIRS). Soil water potential was monitored continuously. We analysed RER data with regard to climate variables and NSC levels of the current and preceding month. Results: NIRS was a reliable method for measuring starch and soluble sugars. NSC was high in the crown and roots but very low in the trunk all year round. Soil temperature was positively correlated to RER (of the current month) between 0 and 8 °C, above which RER stabilised and was not explained by NSC levels or soil water potential. However, mean RER of fine roots in the month following the measurement of NSC was significantly and negatively correlated to soluble sugar and positively correlated with starch content. Very fine root starch content was also positively correlated with root production in the month following the starch measurement. Conclusion: Soil temperature was a major driver of fine root elongation, but at low temperatures only. At soil temperatures >8 °C, no particular driver was dominant. NSC levels were negligible in the stem and root-bases, suggesting that wood production is a major sink that depletes carbohydrates. A large pool of NSC, principally in the form of starch, existed in fine roots of P. abies, and acted as an energy supply for root production throughout the year, even when photosynthesis was limited. Soluble sugars were depleted in fine roots during the growing season, but no relationship was found between fine root production and soluble sugars. The physiological mechanism by which NSC accumulation actively or passively occurs in fine roots is not known but could be due to the symplastic pathway of phloem unloading in conifer root tips, which suggests a passive mechanism.

The following work is focused in the study of the effects of forest management on a mixed forest of Pinus sylvestris and Fagus sylvatica on the pines growth at Aspurz, in the Pyrenees of Navarra. This study started in 1999 when the forest area that was going to be managed was chosen. The zone was divided in 3 blocs of 3 treatment repetitions: 0%, 20% and 30% (in 1999) – 40% (in 2009) of the basal area extracted from the forest in 9 different plots. The final objective of this study is to determine whether the thinning intensity has had any significant effects on the scots pine growth, how they keep after 9 years from the second and last thinning. During the field work, the trees were marked, and the diameters of all the trees and the height of five of each social class and plot were measured. After an analysis of the different parameters (number of trees, mean diameter, mean height and mean basal area of the different treatments) and a variance analysis to test the influence of the forest management treatment and the tree social class were performed through the programs “Excel” and “JMP”. Significant differences were obtained for some variables due to the treatment and the social class, but the main tendency of the results and the analysis proved that the thinning treatments were not anymore effective 9 years after the last and second intervention. Le travail présent est centré dans l’étude des effets des éclaircies forestières dans une forêt mixte de pin sylvestre et hêtre sur les pins sylvestres à Aspurz, aux Pyrénées de Navarre. L’étude a commencé en 1999 quand la surface du bois qui allait être éclairée a été choisie. Elle a été divisée par 3 blocs avec 3 répétitions de traitements d’éclairage : 0%, 20% et 30 % (en 1999) - 40% (en 2009) de la surface basale extraites dans un total de 9 parcelles. L’objectif général de ce projet est de déterminer si les intensités d’éclaircies ont eu un effet significatif diffèrent aux parcelles témoin sur la croissance des pins sylvestres ou si elles sont disparues. Pendant le travail sur champs, les arbres ont été marqués, et les diamètres de tous les arbres et les hauteurs de cinq arbres de chaque classe sociologique et chaque parcelle ont été mesurés. Après une analyse des différents paramètres (nombre d’arbres, diamètre moyen des arbres, hauteur moyenne des arbres et surface basale moyenne par traitement) et une analyse statistique de variance pour tester l’influence du traitement forestier ont été faits avec les programmes « Excel » et « JMP ». Des différences significatives ont été obtenues pour certaines variables en fonction du traitement et de la classe sociologique mais en gros il a été prouvé que les éclaircies n’ont plus d’effet sur cette forêt 9 ans après la dernière et deuxième intervention. Graduado o Graduada en Ingeniería Agroalimentaria y del Medio Rural por la Universidad Pública de Navarra Nekazaritzako Elikagaien eta Landa Ingurunearen Ingeniaritzan graduatua Nafarroako Unibertsitate Publikoan

Quantifying the accuracy of remote sensing products is a timely endeavor given the rapid increase in Earth observation missions. A validation site for Sentinel-2 products was hence established in central Germany. Automatic multispectral and hyperspectral sensor systems were installed in parallel with an existing eddy covariance flux tower, providing spectral information of the vegetation present at high temporal resolution. Normalized Difference Vegetation Index (NDVI) values from ground-based hyperspectral and multispectral sensors were compared with NDVI products derived from Sentinel-2A and Moderate-resolution Imaging Spectroradiometer (MODIS). The influence of different spatial and temporal resolutions was assessed. High correlations and similar phenological patterns between in situ and satellite-based NDVI time series demonstrated the reliability of satellite-based phenological metrics. Sentinel-2-derived metrics showed better agreement with in situ measurements than MODIS-derived metrics. Dynamic filtering with the best index slope extraction algorithm was nevertheless beneficial for Sentinel-2 NDVI time series despite the availability of quality information from the atmospheric correction procedure.

International audience; There are strong uncertainties regarding LAI dynamics in forest ecosystems in response to climate change. While empirical growth & yield models (G&YMs) provide good estimations of tree growth at the stand level on a yearly to decennial scale, process-based models (PBMs) use LAI dynamics as a key variable for enabling the accurate prediction of tree growth over short time scales. Bridging the gap between PBMs and G&YMs could improve the prediction of forest growth and, therefore, carbon, water and nutrient fluxes by combining modeling approaches at the stand level. Our study aimed to estimate monthly changes of leaf area in response to climate variations from sparse measurements of foliage area and biomass. A leaf population probabilistic model (SLCD) was designed to simulate foliage renewal. The leaf population was distributed in monthly cohorts, and the total population size was limited depending on forest age and productivity. Foliage dynamics were driven by a foliation function and the probabilities ruling leaf aging or fall. Their formulation depends on the forest environment. The model was applied to three tree species growing under contrasting climates and soil types. In tropical Brazilian evergreen broadleaf eucalypt plantations, the phenology was described using 8 parameters. A multi-objective evolutionary algorithm method (MOEA) was used to fit the model parameters on litterfall and LAI data over an entire stand rotation. Field measurements from a second eucalypt stand were used to validate the model. Seasonal LAI changes were accurately rendered for both sites (R-2 = 0.898 adjustment, R-2 = 0.698 validation). Litterfall production was correctly simulated (R-2 = 0.562, R-2 = 0.4018 validation) and may be improved by using additional validation data in future work. In two French temperate deciduous forests (beech and oak), we adapted phenological sub-modules of the CASTANEA model to simulate canopy dynamics, and SLCD was validated using LAI measurements. The phenological patterns were simulated with good accuracy in the two cases studied. However, IA/max was not accurately simulated in the beech forest, and further improvement is required. Our probabilistic approach is expected to contribute to improving predictions of LAI dynamics. The model formalism is general and suitable to broadleaf forests for a large range of ecological conditions. (C) 2014 Elsevier B.V. All rights reserved.

This paper documents the accuracy of a post-correction method applied to precipitation regionalized by the Weather Research and Forecasting (WRF) Regional Climate Model (RCM) for improving simulated rainfall and feeding impact studies. The WRF simulation covers Burgundy (northeastern France) at a 8-km resolution and over a 20-year long period (1989–2008). Previous results show a strong deficiency of the WRF model for simulating precipitation, especially when convective processes are involved. In order to reduce such biases, a Quantile Mapping (QM) method is applied to WRF-simulated precipitation using the mesoscale atmospheric analyses system SAFRAN («Système d'Analyse Fournissant des Renseignements Adaptés à la Nivologie») that provides precipitation data at an 8 km resolution. Raw and post-corrected model outputs are next used to compute the soil water balance of 30 Douglas-fir and 57 common Beech stands across Burgundy, for which radial growth data are available. Results show that the QM method succeeds at reducing the model's wet biases in spring and summer. Significant improvements are also noted for rainfall seasonality and interannual variability, as well as its spatial distribution. Based on both raw and post-corrected rainfall time series, a Soil Water Deficit Index (SWDI) is next computed as the sum of the daily deviations between the relative extractible water and a critical value of 40% below which the low soil water content induce stomatal regulation. Post-correcting WRF precipitation does not significantly improve the simulation of the SWDI upon the raw (uncorrected) model outputs. Two characteristic years were diagnosed to explain this unexpected lack of improvement. Although the QM method allows producing realistic precipitation amounts, it does not correct the timing errors produced by the climate model, which is yet a major issue to obtain reliable estimators of local-scale bioclimatic conditions for impact studies. A realistic temporality of simulated precipitation is thus required before using any systematic post-correction method for appropriate climate impact assessment over temperate forests. Fil: Boulard, Damien. Universite de Bourgogne; Francia Fil: Castel, Thierry. Universite de Bourgogne; Francia Fil: Camberlin, Pierre. Universite de Bourgogne; Francia Fil: Sergent, Anne Sophie. Institut National de la Recherche Agronomique; Francia. Instituto Nacional de Tecnología Agropecuaria. Centro Regional Patagonia Norte. Estación Experimental Agropecuaria San Carlos de Bariloche; Argentina Fil: Asse, Daphné. Crea Centre de Recherches Sur Les Ecosystèmes Daltitud; Francia Fil: Bréda, Nathalie. Institut National de la Recherche Agronomique; Francia Fil: Badeau, Vincent. Institut National de la Recherche Agronomique; Francia Fil: Rossi, Aurélien. Universite de Bourgogne; Francia Fil: Pohl, Benjamin. Universite de Bourgogne; Francia

Quantifying the accuracy of remote sensing products is a timely endeavor given the rapid increase in Earth observation missions.[br/] A validation site for Sentinel-2 products was hence established in central Germany. Automatic multispectral and hyperspectral sensor systems were installed in parallel with an existing eddy covariance flux tower, providing spectral information of the vegetation present at high temporal resolution. Normalized Difference Vegetation Index (NDVI) values from ground-based hyperspectral and multispectral sensors were compared with NDVI products derived from Sentinel-2A and Moderate-resolution Imaging Spectroradiometer (MODIS). The influence of different spatial and temporal resolutions was assessed.[br/] High correlations and similar phenological patterns between in situ and satellite-based NDVI time series demonstrated the reliability of satellite-based phenological metrics. Sentinel-2-derived metrics showed better agreement with in situ measurements than MODIS-derived metrics.[br/] Dynamic filtering with the best index slope extraction algorithm was nevertheless beneficial for Sentinel-2 NDVI time series despite the availability of quality information from the atmospheric correction procedure.

How temperate forests will respond to climate change is uncertain; projections range from severe decline to increased growth. We conducted field tests of sessile oak (Quercus petraea), a widespread keystone European forest tree species, including more than 150,000 trees sourced from 116 geographically diverse populations. The tests were planted on 23 field sites in six European countries, in order to expose them to a wide range of climates, including sites reflecting future warmer and drier climates. By assessing tree height and survival, our objectives were twofold: (1) to identify the source of differential population responses to climate (genetic differentiation due to past divergent climatic selection versus plastic responses to ongoing climate change), (2) to explore which climatic variables (temperature or precipitation) trigger the population responses. Tree growth and survival were modeled for contemporary climate and then projected using data from four regional climate models for years 2071-2100, using two greenhouse gas concentration trajectory scenarios each. Overall results indicated a moderate response of tree height and survival to climate variation, with changes in dryness (either annual or during the growing season) explaining the major part of the response. Whilst, on average, populations exhibited local adaptation, there was significant clinal population differentiation for height growth with winter temperature at the site of origin. The most moderate climate model (HIRHAM5-EC; rcp4.5) predicted minor decreases in height and survival, whilst the most extreme model (CCLM4-GEM2-ES; rcp8.5) predicted large decreases in survival and growth for southern and southeastern edge populations (Hungary and Turkey). Other non-marginal populations with continental climates were predicted to be severely and negatively affected (Bercé, France), while populations at the contemporary northern limit (colder and humid maritime regions; Denmark and Norway) will probably not show large changes in growth and survival in response to climate change.

The timing of leaf phenophases greatly influences the functioning of trees. Phenological traits are thus considered major components of tree fitness, and are expected to be strongly selected under environmental or biotic pressures. To date, most phenological studies have been conducted at the population scale, with comparatively very few works at the scale of individuals. We take advantage of a unique phenological database, consisting of leaf unfolding (LU) and leaf senescence (LS) observations done at the individual scale for 5 years over 35 populations, representing >1200 dominant and co-dominant trees of three species (Quercus robur, Quercus petraea and Fagus sylvatica), to document the within-population variability of phenological traits in temperate deciduous forest trees. We show that individual phenological ranks mostly repeat from year to year among trees of a given population, though LU ranks tend to repeat more than LS ranks. Using simulated dynamics of soil water content, we suggest that inter-individual differences in the access to soil water may play a role in determining tree phenological ranks. The timing of leaf phenology is further correlated with individual growth in a given tree population. In Beech populations, early-leafers tend to grow more. On the other hand, Oak trees entering senescence later tend to grow more. The growth of Oak trees, being more prone to spring pathogenic attacks than Beech, may benefit less from a spring extension of leaf display.

Nous avons distingué trois types d'interactions entre la biodiversité et l'adaptation des forêts au changement climatique : (i) l'adaptation de la sylviculture en vue du maintien des services rendus par la forêt, en particulier la production de bois, aura des conséquences indirectes sur la biodiversité; (ii) la biodiversité pourra en partie aider à l'adaptation des forêts; (iii) les modèles pour le siècle à venir et, de façon moins tranchée, les observations de la dynamique récente des écosystèmes forestiers tempérés montrent que le changement climatique, s'il n'est pas contenu, aura des impacts directs importants sur la biodiversité forestière. Quelle mesure d'adaptation faut-il prendre pour la protéger ?

European forests are an important carbon sink; however, the relative contributions to this sink of climate, atmospheric CO 2 concentration ([CO 2]), nitrogen deposition and forest management are under debate. We attributed the European carbon sink in forests using ORCHIDEE-FM, a process-based vegetation model that differs from earlier versions of ORCHIDEE by its explicit representation of stand growth and idealized forest management. The model was applied on a grid across Europe to simulate changes in the net ecosystem productivity (NEP) of forests with and without changes in climate, [CO 2] and age structure, the three drivers represented in ORCHIDEE-FM. The model simulates carbon stocks and volume increment that are comparable - root mean square error of 2 m 3 ha -1 yr -1 and 1.7 kg C m -2 respectively - with inventory-derived estimates at country level for 20 European countries. Our simulations estimate a mean European forest NEP of 175 ± 52 g C m -2 yr -1 in the 1990s. The model simulation that is most consistent with inventory records provides an upwards trend of forest NEP of 1 ± 0.5 g C m -2 yr -2 between 1950 and 2000 across the EU 25. Furthermore, the method used for reconstructing past age structure was found to dominate its contribution to temporal trends in NEP. The potentially large fertilizing effect of nitrogen deposition cannot be told apart, as the model does not explicitly simulate the nitrogen cycle. Among the three drivers that were considered in this study, the fertilizing effect of increasing [CO 2] explains about 61% of the simulated trend, against 26% to changes in climate and 13% only to changes in forest age structure. The major role of [CO 2] at the continental scale is due to its homogeneous impact on net primary productivity (NPP). At the local scale, however, changes in climate and forest age structure often dominate trends in NEP by affecting NPP and heterotrophic respiration.

European forests are intensively exploited for wood products, yet they also form a sink for carbon. European forest inventories, available for the past 50 years, can be combined with timber harvest statistics to assess changes in this carbon sink. Analysis of these data sets between 1950 and 2000 from the EU-15 countries excluding Luxembourg, plus Norway and Switzerland, reveals that there is a tight relationship between increases in forest biomass and forest ecosystem productivity but timber harvests grew more slowly. Encouragingly, the environmental conditions in combination with the type of silviculture that has been developed over the past 50 years can efficiently sequester carbon on timescales of decades, while maintaining forests that meet the demand for wood. However, a return to using wood as biofuel and hence shorter rotations in forestry could cancel out the benefits of carbon storage over the past five decades. European forests are intensively exploited for wood products, yet they are also a potential sink for carbon. European forest inventories combined with timber harvest statistics from sixteen European countries show that between 1950 and 2000 forest biomass increased faster than the amount of timber harvests. Silviculture, which has developed over the past 50 years, can efficiently sequester carbon on timescales of decades, while maintaining forests that meet the demand for wood.

Our aim was to investigate the possible enhancing role of long-term atmospheric CO2 increase on wood density as an essential component of biomass sequestration. We therefore assessed the long-term evolution of wood density over pre-industrial and contemporary periods, in a regional context free of management practices, atmospheric deposition and with restricted climatic change. Dominant trees of Araucaria araucana were sampled in 37 stands distributed throughout its natural distribution over temperate forests of the Chilean Andes Cordillera. Mean ring density (MRD) at 1.30 m was measured by X-ray micro-densitometry. A third-century MRD chronology was built after MRD standardisation by the effects of cambial age and radial growth, simultaneously estimated from a statistical model from rings of the pre-industrial period (1700-1850) to avoid any temporal bias. The age-alone standardised MRD chronology showed restricted fluctuations ranging between 2.0 and 3.2 % over the last three centuries. Multi-decennial fluctuations between ring width and MRD were found qualitatively synchronous and opposed. Accordingly, MRD fluctuations were removed with the age and growth standardisation, highlighting the absence of a historical wood density trend at constant ring size over the whole period. Over the most recent decades however, a slight increase in density was identified, with loss of synchronisation to radial growth. The absence of a long-term signal in wood density of A. araucana, in a context of restricted anthropogenic influence, differs markedly from reports of significant decreases in the Northern Hemisphere. We conclude as part of the analysis done that until now, increasing atmospheric CO2 unlikely would have an impact in wood density.

International audience; Mapping forest composition is a major concern for forest management, biodiversity assessment and for understanding the potential impacts of climate change on tree species distribution. In this study, the suitability of a dense high spatial resolution multispectral Formosat-2 satellite image time-series (SITS) to discriminate tree species in temperate forests is investigated. Based on a 17-date SITS acquired across one year, thirteen major tree species (8 broadleaves and 5 conifers) are classified in a study area of southwest France. The performance of parametric (GMM) and nonparametric (k-NN, RF, SVM) methods are compared at three class hierarchy levels for different versions of the SITS: (i) a smoothed noise-free version based on the Whittaker smoother; (ii) a non-smoothed cloudy version including all the dates; (iii) a non-smoothed noise-free version including only 14 dates. Noise refers to pixels contaminated by clouds and cloud shadows. The results of the 108 distinct classifications show a very high suitability of the SITS to identify the forest tree species based on phenological differences (average κ=0.93 estimated by cross-validation based on 1235 field-collected plots). SVM is found to be the best classifier with very close results from the other classifiers. No clear benefit of removing noise by smoothing can be observed. Classification accuracy is even improved using the non-smoothed cloudy version of the SITS compared to the 14 cloud-free image time series. However conclusions of the results need to be considered with caution because of possible overfitting. Disagreements also appear between the maps produced by the classifiers for complex mixed forests, suggesting a higher classification uncertainty in these contexts. Our findings suggest that time-series data can be a good alternative to hyperspectral data for mapping forest types. It also demonstrates the potential contribution of the recently launched Sentinel-2 satellite for studying forest ecosystems.

Key message We demonstrate that, beyond leaf phenology, the phenological cycles of wood and fine roots present clear responses to environmental drivers in temperate and boreal trees. These drivers should be included in terrestrial ecosystem models. Context In temperate and boreal trees, a dormancy period prevents organ development during adverse climatic conditions. Whereas the phenology of leaves and flowers has received considerable attention, to date, little is known regarding the phenology of other tree organs such as wood, fine roots, fruits, and reserve compounds. Aims Here, we review both the role of environmental drivers in determining the phenology of tree organs and the models used to predict the phenology of tree organs in temperate and boreal forest trees. Results Temperature is a key driver of the resumption of tree activity in spring, although its specific effects vary among organs. There is no such clear dominant environmental cue involved in the cessation of tree activity in autumn and in the onset of dormancy, but temperature, photoperiod, and water stress appear as prominent factors. The phenology of a given organ is, to a certain extent, influenced by processes in distant organs. Conclusion Inferring past trends and predicting future trends of tree phenology in a changing climate requires specific phenological models developed for each organ to consider the phenological cycle as an ensemble in which the environmental cues that trigger each phase are also indirectly involved in the subsequent phases. Incorporating such models into terrestrial ecosystem models (TEMs) would likely improve the accuracy of their predictions. The extent to which the coordination of the phenologies of tree organs will be affected in a changing climate deserves further research.

Présenté devant un groupe coopératif d'industriels forestiers et d'universitaires brésiliens, à l'invitation du CIRAD et de l'Ambassade de France au Brésil, l'exposé donne un point de vue européen sur les forêts et le changement climatique, le situant en particulier vis-à-vis de la diversité des forêts françaises à objectif de production et de leurs contrastes avec les sylviculture de plantation brésiliennes. La communication cherche, au-delà des différences de contexte biogéographique, social et industriel, à montrer l'importance de l'itinéraire technique plantation dans la mise en œuvre des différentes options de changement de gestion.

Global environmental changes may lead to a modification of abiotic factors and biological diversity. The production function in forest ecosystems has the particularity to be both subjected to, and a regulator of, these environmental changes. Understanding how forest productivity is driven by species diversity and environmental factors is therefore a critical issue. This PhD thesis studies how tree species mixture affects their productivity along edaphic and climatic gradients. Based on an approach using both an original dataset and the national forest inventory dataset, I focused on five major species of European forests in pure and two-species forests: Quercus petraea, Pinus sylvestris, Fagus sylvatica, Abies alba and Picea abies. In lowlands, abiotic factors had little impacts on the outcome of tree species mixture on productivity. In addition, the effect of tree species mixture was determined by the shade tolerance of companion species. In highlands, the positive effects of tree species mixture were strongest when the abiotic factors were the most limiting for growth. These results highlight the importance of considering abiotic factors and the functional characteristics of species as drivers of the effect of biological diversity on ecosystem functions., Les changements environnementaux à l’échelle globale peuvent se traduire par une modification des conditions abiotiques et de la diversité biologique. La fonction de production des écosystèmes forestiers a la particularité d'être à la fois soumise à, et régulatrice de ces changements environnementaux. Il apparaît alors primordial de comprendre la manière dont la diversité en essences et les facteurs abiotiques exercent le contrôle de cette fonction. Au cours de mes travaux j’ai étudié la variation de l'effet du mélange d’essences sur leur productivité dans un contexte climatique et édaphique changeant. Grâce à une double approche, basée sur des données locales et des données de l'Inventaire forestier national, je me suis concentrée sur cinq espèces forestières européennes majeures en peuplements purs et bispécifiques : Quercus petraea, Pinus sylvestris, Fagus sylvatica, Abies alba et Picea abies. En plaine je montre que l'effet du mélange d’essences varie peu avec les facteurs abiotiques. De plus cet effet est déterminé par la tolérance à l'ombrage de l'espèce accompagnatrice. En montagne l’effet du mélange d’essences sur leur productivité est plus fort et positif lorsque les facteurs abiotiques sont les plus limitants pour la croissance. Ces résultats illustrent l’intérêt d'intégrer les facteurs abiotiques et les caractéristiques fonctionnelles des espèces comme déterminants de l'effet de la diversité biologique sur les fonctions de l'écosystème.

Ecological models are increasingly used as decision-making tools and their reliability is becoming a key issue. At the same time, the sophistication of techniques for model development and analysis has given rise to a relative compartmentalization of model building and evaluation tasks. Several guidelines invite ecological modelers to follow an organized sequence of development and analysis steps and have coined the term "evaludation" for this process. The objective of this paper is to assess the feasibility and the value of a structured evaludation process, based on the working example of the Samsara2 model, a spatially explicit individual-based forest dynamics model. We implemented the six steps of model design, process level calibration, qualitative evaluation, quantitative evaluation, global sensitivity analysis, and partial recalibration using approximate Bayesian computing. We then evaluated how the evaludation process revealed model strengths and weaknesses, specified the model's conditions of use, clarified how the model works, and provided insights into forest ecosystem functioning. Finally, the efficiency/cost ratio of the process and future improvements are discussed.

International audience; With the evolution of ecosystem management and the actual climate change we are facing, there is a need to improve our knowledge of carbon (C) balance and more specifically of C allocation in the plants. In our study, we quantified the seasonal variation of gross primary production (GPP, obtained through eddy covariance measurements) and biomass production (BP, the C fixed into the biomass obtained thanks to inventory campaign) for a 60-year-old even-aged beech stand located in North East of France. We also assessed the seasonal evolution of the BP efficiency (BPE=BP/GPP; Vicca et al., 2012) and its potential determining factors for our site. For 2014, we found a net ecosystem exchange (NEE) of -549 gC m-2, corresponding to a C sequestration. This value breaks down between 1089 gC m-2 for the respiration of the ecosystem and -1639 gC m-2 for the GPP. On the same year, our stand built up 461.6 gC m-2 of tree biomass (leaves, trunk, branches, fine roots), leading to an annual BPE of 0.28, which is within the range of value found on other similar sites. There was a large temporal variation of C allocation to the different parts of the tree biomass during the growth season. Our results show that the growth first happened in the trunk and branches -with a peak value of 74.5 gC m-2 month-1 in May - whereas the fine roots biomass production started later (end of July) and reached a maximum at the end of the growth season (28.49 gC m-2 month-1 for September). The BPE varied also during the year from 0.13 in April to 0.31 in August, where the BP was the same than in July but the cumulated GPP was already decreasing. The seasonal variation may be mainly explained by climatic variations, whereas the shift between woody above-ground biomass and fine roots biomass could be explained by the phenology (linked to physiological mechanisms).

Characterizing the above- and belowground carbon stocks of ecosystems is vital for a better understanding of the role of vegetation in carbon cycling. Yet studies on forest ecosystems at high altitudes remain scarce. We examined above- and belowground carbon partitioning in trees growing in mixed montane/upper montane forest ecosystems in the French Alps. Field work was performed in three forests along a gradient of both altitude (1400 m, 1700 m, and 2000 m) and altitude-induced species composition (from lower altitude Abies alba and Fagus sylvatica to higher altitude Picea abies and Pinus uncinata). We performed forest inventories and root sampling along soil wall profiles, so that the stand basal area (SBA, in m2 ha-1) and root cross-sectional area (RCSA, in m2 ha-1) were estimated at each altitude. To characterize the carbon allocation trend between the above-and belowground compartments, the ratio of RCSA to SBA was then calculated. We found that both SBA and RCSA of coarse roots (diameter > 2 mm) were significantly different among the three altitudes. No significant difference in RCSA of fine roots (diameter ? 2 mm) was found among altitudes. The ratio of RCSA of fine roots to SBA augmented with increasing elevation, suggesting that forest ecosystems at higher altitudes allocate more carbon from above- to belowground organs. This increased allocation to fine roots would allow trees to scavenge nutrients more efficiently throughout the short growing season. Furthermore, this work highlighted the interest of using easy to measure area-based indicators as proxies of root and stem biomass when investigating carbon partitioning in highly heterogeneous montane/upper montane forests.

The aim of the present work was to provide a synopsis of the scientific literature concerning the effects of different tree spe- cies on soil and to quantify the effect of common European temperate forest species on soil fertility. The scientific literature dealing with the tree species effect on soil has been reviewed. The composition of forest overstory has an impact on the chemical, physical and biolo- gical characteristics of soil. This impact was highest in the topsoil. Different tree species had significantly different effects on water ba- lance and microclimate. The physical characteristics of soils also were modified depending on the overstory species, probably through modifications of the soil fauna. The rates of organic matter mineralization and nitrification seem to be dependent on tree species. A coni- ferous species, Picea abies, had negative input-output budgets for some nutrients, such as Ca and Mg. This species promoted a higher soil acidification and a decrease in pH. Thus, it should not be planted in very poor soils in areas affected by acidic atmospheric deposi- tions. Nevertheless, the effect of the canopy species on soil fertility was rarely significant enough to promote forest decline. The impact of a tree species on soil fertility varied depending on the type of bedrock, climate and forest management. forest soils / tree species / fertility / sustainability / resiliency

Insectivorous bats are increasingly recognized as potential regulators of pest insect populations.They also represent the group of European mammals with the most unfavorable conservation status. Forests are key habitats for many bat species but are currently under threat from climate change and fragmentation. It is therefore urgent to better understand the relationships between the bats, their prey and their habitats in forests. Our main objective was to quantify the effects, at multiple spatial scales, of the main attributes of forest habitats on the activity, species richness, functional diversity and composition of European bat communities. They were studied using manipulative experiments in Aquitaine plantation forests and automatic recordings in the network of exploratory plots set up in six European countries by the FunDivEurope project. From the plot to the continent scale, increasing tree diversity, amount of broad leaved trees and dead wood, had positive effects on bat communities through an increase in prey and roost resources. However these effects were not stationary, being stronger at higher latitudes, probably due to lower habitat carrying capacity in relation to harsher climatic conditions. In addition we experimentally demonstrated that the numerical and functional responses of bats to prey density could result in effective regulation of pine processionary moth populations. Forest management strategies aim at enhancing key habitat structures, are eventually proposed in order to improve the conservation of bats and to increase the service of pest regulation they can provide.; Les chiroptères sont reconnus comme de potentiels régulateurs des populations d’insectes. Ce sont aussi les mammifères européens pour lesquels les enjeux de conservation sont les plus importants. Ils trouvent dans les forêts des habitats favorables qui sont cependant menacés par les changements climatiques et la fragmentation. Il convient donc de mieux comprendre lesrelations entre les communautés de chiroptères, leurs habitats et leurs proies en forêt. L'objectif de cette thèse est de quantifier les effets, à différentes échelles spatiales, desprincipales composantes de l’habitat forestier sur l’activité, la richesse spécifique, la diversité fonctionnelle et la composition des communautés de chiroptères européens. Les résultats reposent sur des données collectées grâce à des protocoles expérimentaux en Aquitaine et dans les six pays du réseau de placettes forestières organisé par le projet FunDivEurope. De la parcelle au continent, l'accroissement de la diversité des essences forestières, de la proportion de feuillus et du bois mort, en augmentant les ressources en proies et en gîtes, ont des effets positifs sur les communautés de chiroptères. Ces effets, non stationnaires, se renforcent vers le nord avec la rigueur du climat. Nous confirmons également que les chiroptères forestiers, par leur réponse numérique et fonctionnelle aux densités de proie, peuvent limiter la démographie d’un insecte défoliateur. Des mesures de gestion, visant le renforcement des structures-clés des habitats forestiers, sont proposées pour favoriser la conservation des communautés de chiroptères et leur capacité de régulation des insectes ravageurs.

International audience

International audience; & Context European beech (Fagus sylvatica L.) is considered threatened by anticipated climate change, but the physiological causes of potential beech decline or mortality remains poorly understood. & Aims The purpose of the present study was to fuel debate about the assumption that carbohydrate depletion is involved in the decline of mature European beech. & Methods The health status of beech trees from a severely declining stand was visually assessed by examining their crown condition. Content and radial distribution of non-structural carbohydrates (starch and soluble carbohydrate) were analyzed in the trunks and compared to those reported earlier in trunks of healthy beech trees. & Results and discussion The distribution of carbohydrate in the beech trunks recorded here seemed affected by decline. We found a stronger radial decrease of starch content than those reported earlier for healthy beech trees. Carbohydrate reserves appear partially maintained in the outermost rings while starch depletion occurred in older wood rings in declining trees that may be able to mobilize carbohydrate reserves from older wood rings in response to successive climatic constraints.