Thibaut Fréjaville

Les forêts d'altitude connaissent une émergence croissante des feux. Via une analyse rétrospective des feux et du climat des dernières décennies dans le sud-est de la France, cette thèse montre d'abord que l'augmentation de l'activité des feux a été largement restreinte aux zones de montagne, les politiques de lutte contraignant les feux au sein des paysages méditerranéens. Les attributs d'inflammabilité et de résistance des forêts Alpines qui gouvernent ses effets et sa réponse au feu ont ensuite été analysés au sein des niches environnementales des espèces dominantes. Les résultats obtenus montrent que l'abondance et les propriétés d'inflammabilité des litières, des strates herbacées et arbustives varient en fonction de l'ouverture de canopée et la saisonnalité des précipitations. Par ailleurs, les traits de résistance au feu des arbres varient entre espèces en lien avec l'inflammabilité des communautés, selon un compromis entre tolérance (écorce épaisse en climat sec) et évitement (houppier haut en climat humide), suggérant une coévolution entre espèces et feux. Ces gradients biotiques et abiotiques gouvernent ainsi les niches d'inflammabilité des espèces, depuis les forêts denses et humides peu inflammables des Alpes externes du nord aux forêts ouvertes inflammables périméditerranéennes des Alpes du sud et subalpines des Alpes internes. Ainsi, du fait des propriétés de leur niche et de leurs traits d'histoire de vie, les arbres Alpins ne sont pas tous égaux face à l'émergence des feux au sein de la diversité des conditions de végétation et de climat des montagnes du sud de l'Europe.

Aim: Tree mortality is increasing world‐wide, leading to changes in forest composition and altering global biodiversity. Nonetheless, owing to the multifaceted stochastic nature of tree mortality, large‐scale spatial patterns of mortality across species ranges and their underlying drivers remain difficult to understand. Our main goal was to describe the geographical patterns and drivers of the occurrence of mortality (presence of a mortality event) and the intensity of tree mortality (amount of mortality related to that mortality event) in Europe. We hypothesized that the occurrence of mortality represents background mortality and is higher in the margin than in core populations, whereas the intensity of mortality could have a more even distribution according to the spatial and temporal stochasticity of die‐off events.Location: Europe (Spain, France, Germany, Belgium, Sweden and Finland).Major taxa studied: More than 1.5 million trees belonging to 20 major forest tree species.Methods: We developed binomial and truncated negative binomial models to tease apart the occurrence and intensity of tree mortality in National Forest Inventory plots at the range‐wide scale. The occurrence of mortality indicated that at least one tree had died in the plot, whereas the intensity of mortality referred to the number of dead trees per plot.Results: The highest occurrence of mortality was found in peripheral regions and the climatic trailing edge linked with drought, whereas the intensity of mortality was driven by competition, drought and high temperatures and was scattered uniformly across species ranges.Main conclusions: We show that tree background mortality, but not die‐off, is generally higher in the trailing‐edge populations. It remains to be explored whether other demographic traits, such as growth, reproduction and regeneration, also decrease at the trailing edge of European tree populations.

Aim Tree mortality is increasing worldwide, leading to changes in forest composition and altering global biodiversity. Yet, due to the multi-faceted stochastic nature of tree mortality, large-scale spatial patterns of mortality across species ranges and their underlying drivers remain difficult to understand. Our main goal is to describe the geographical patterns and drivers of the occurrence and intensity of tree mortality in Europe. We hypothesize that the occurrence of mortality represents background mortality and is higher in the margin than the core populations, whereas the intensity of mortality could have a more even distribution according to the spatial and temporal stochasticity of die-off events. Location Europe (Spain, France, Germany, Belgium, Sweden and Finland) Time period 1981 to 2014. Major taxa studied More than 1.5 million trees belonging to 20 major forest tree species Methods We develop hurdle models to tease apart the occurrence and intensity of tree mortality in National Forest Inventory plots at range-wide scale. The occurrence of mortality indicates that at least one tree has died in the plot and the intensity of mortality refers to the number of trees dead per plot. Results The highest mortality occurrence was found in peripheral regions and the climatic trailing edge linked with drought, whereas the intensity of mortality was driven by competition, drought and high temperatures and was uniformly scattered across species ranges. Main conclusions Our findings provide a new perspective in our understanding of tree mortality across species ranges. We show that tree background mortality but not die-off is generally higher in the trailing edge populations, but whether other demographic traits such as growth, reproduction and regeneration would also decrease at the trailing edge of European tree populations needs to be explored.

How populations of long‐living species respond to climate change depends on phenotypic plasticity and local adaptation processes. Marginal populations are expected to have lags in adaptation (i.e. differences between the climatic optimum that maximizes population fitness and the local climate) because they receive pre‐adapted alleles from core populations preventing them from reaching a local optimum in their climatically marginal habitat. Yet, whether adaptation lags in marginal populations are a common feature across phylogenetically and ecologically different species and how lags can change with climate change remain unexplored. To test for range‐wide patterns of phenotypic variation and adaptation lags of populations to climate, we (a) built model ensembles of tree height accounting for the climate of population origin and the climate of the site for 706 populations monitored in 97 common garden experiments covering the range of six European forest tree species; (b) estimated populations' adaptation lags as the differences between the climatic optimum that maximizes tree height and the climate of the origin of each population; (c) identified adaptation lag patterns for populations coming from the warm/dry and cold/wet margins and from the distribution core of each species range. We found that (a) phenotypic variation is driven by either temperature or precipitation; (b) adaptation lags are consistently higher in climatic margin populations (cold/warm, dry/wet) than in core populations; (c) predictions for future warmer climates suggest adaptation lags would decrease in cold margin populations, slightly increasing tree height, while adaptation lags would increase in core and warm margin populations, sharply decreasing tree height. Our results suggest that warm margin populations are the most vulnerable to climate change, but understanding how these populations can cope with future climates depend on whether other fitness‐related traits could show similar adaptation lag patterns.

Aim: To test whether intraspecific trait responses to climate among populations across species distribution ranges can be untangled using field observations, under the rationale that, in natural forest tree populations, long‐term climate shapes population responses while recent climate change drives phenotypic plasticity.Location: Europe.Time period: 1901–2014.Taxa: Silver fir (Abies alba Mill.) and sessile oak [Quercus petraea (Matt.) Liebl.].Methods: We estimated the variation of individual tree height as a function of long‐term and short‐term climates to tease apart provenance effects (variation among populations of different geographical origin), plasticity and their interaction, using mixed‐effect models calibrated with national forest inventory data (in‐situ models). To validate our approach, we tested the ability of in‐situ models to predict independently tree height observations in common gardens experiments where provenance and plastic effects can be measured and separated. In‐situ model predictions of tree height variation among provenances and among planting sites were compared to observations in common gardens and to predictions from a similar model calibrated using common garden data (ex‐situ model).Results: In Q. petraea, we found high correlations between in‐situ and ex‐situ model predictions of provenance and plasticity effects and their interaction for tree height (r > .80). We showed that the in‐situ models significantly predicted tree height variation among provenances and sites for A. alba and Q. petraea. Spatial predictions of phenotypic plasticity across species distribution ranges indicate decreasing tree height in populations of warmer climates in response to recent anthropogenic climate warming.Main conclusions: Our modelling approach using national forest inventory observations provides a new perspective for understanding patterns of intraspecific trait variation across species ranges. Its application is particularly interesting for species for which common garden experiments do not exist or do not cover the entire climatic range of the species.

Aim: Background tree mortality is a complex demographic process that affects forest structure and long-term dynamics. We aimed to test how drought intensity interacts with interspecific and intraspecific competition (or facilitation) in shaping individual mortality patterns across tree species ranges.Location: European latitudinal gradient (Spain to Finland).Time period: 1985 – 2014.Major taxa studied: Scots pine (Pinus sylvestrisL.) and European beech (Fagus sylvaticaL.).Methods: We performed logistic regression models based on individual tree mortality recorded in five European National Forest Inventories. We computed the relative importance of climatic drought intensity, basal area of conspecific and heterospecific trees (proxy of indirect intra- and interspecific competition or facilitation) and the effects of their interactions on mortality along the latitudinal gradient of both species range.Results: Increase in drought intensity over the study period was associated with higher mortality rates in both species. Drought was the most important driver of beech mortality at almost all latitudes while Scots pine mortality was mainly driven by basal area. High conspecific basal area was associated with high mortality rates in both species while high heterospecific basal area was correlated with mortality rates that were high in Scots pine but low in beech.Main conclusions: Beech mortality was directly affected by drought while Scots pine mortality was indirectly affected by drought through interactions with basal area. Despite their different sensitivity to drought and basal area, the highest predicted mortality rates for both species were at the ecotone between Mediterranean and cool temperate biomes, which can be explained by the combined effect of drought and competition. In the context of global warming, which is expected to be particularly strong in the Mediterranean biome, our results suggest that populations at the southern limit of species ranges may experience increased mortality rates in the near future.BIOSKETCH: The authors’ research is focused on functional trait ecology and global change, with special attention to mortality and demography processes. The authors use modelling multidisciplinary approaches to understand complex processes in ecology on a large geographical scale.

Aim: To test whether adaptive and plastic trait responses to climate across species distribution ranges can be untangled using field observations, under the rationale that, in natural forest tree populations, long-term climate shapes local adaptation while recent climate change drives phenotypic plasticity.Location: Europe.Time period: 1901-2014.Taxa: Silver fir (Abies alba Mill.) and sessile oak (Quercus petraea (Matt.) Liebl.).Methods: We estimated the variation of individual tree height as a function of long-term and short-term climates to tease apart local adaptation, plasticity and their interaction, using mixed-effect models calibrated with National Forest Inventory data (in-situ models). To validate our approach, we tested the ability of in-situ models to predict independently tree height observations in common gardens where local adaptation to climate of populations and their plasticity can be measured and separated. In-situ model predictions of tree height variation among provenances (populations of different geographical origin) and among planting sites were compared to observations in common gardens and to predictions from a similar model calibrated using common garden data (ex-situ model).Results: In Q. petraea, we found high correlations between in-situ and ex-situ model predictions of provenance and plasticity effects and their interaction on tree height (r > 0.80). We showed that the in-situ models significantly predicted tree height variation among provenances and sites for Abies alba and Quercus petraea. Spatial predictions of phenotypic plasticity across species distribution ranges indicate decreasing tree height in populations of warmer climates in response to recent anthropogenic climate warming.Main conclusions: Our modelling approach using National Forest Inventory observations provides a new perspective for understanding patterns of intraspecific trait variation across species ranges. Its application is particularly interesting for species for which common garden experiments do not exist or do not cover the entire climatic range of the species.

Fire resistance traits drive tree species composition in surface‐fire ecosystems, but how they covary at different scales of variation and with the environment is not well documented. We assessed the covariation of bark thickness (BT), tree height, and crown base‐to‐height ratio across Alpine forests, after accounting for the effects of tree diameter and competition for light on individual trait variation. Traits consistently correlated across individuals and communities, although the variance of BT mainly occurred among species, whereas crown elevation traits varied mainly within species. Aridity, temperature, and competition contributed to explain the variation of fire resistance traits among and within species, driving a trade‐off between fire resistance and the ability to compete for light. Thick‐barked species (fire‐tolerant) that self‐prune their lower branches (flame‐avoiders) dominated the most fire‐prone and flammable communities in sub‐Mediterranean southern Alps, whereas thin‐barked tree species that grow tall (competition for light) dominated the least fire‐prone communities in the northern Alps. Our findings suggest a long‐term interaction between mountain tree species and fire regime. Higher allocation to trunk elongation occurs in moist and shade environments, while higher allocation to thicken the bark and distancing the crown base from surface fuels occurs in open‐canopy, dry forests where fire spreads with higher intensity.