Microgeographic adaptation and the effect of pollen flow on the adaptive potential of a temperate tree species

Recent interest for microgeographic adaptation, i.e. adaptation at spatial scales compatible with substantial amount of gene dispersal, offers to reconsider the scale at which evolution occurs (Richardson et al. 2014). Whether gene flow is constraining or facilitating local adaptation at this fine spatial scale remains an unresolved question. Too important gene flow would overwhelm the effects of natural selection and decrease local adaptation along environmental gradients. Conversely, gene flow, and particularly long-distance dispersal events, could play a major role in resupplying the genetic variation of populations and favouring the spread of beneficial alleles (Kremer et al. 2012). Hence, the high dispersal capacities of trees are often assumed to be the main process maintaining the large levels of genetic variation measured in their natural populations. However, evidences for microgeographic adaptation and the quantitative assessment of the impact of gene flow on adaptive genetic variation are still limited in most temperate trees. Here, we sampled 60 open-pollinated families of European beech (Fagus sylvatica L.) from three natural plots, spreading along a short elevation gradient (∼1.5 km) at the warm margin of this species distribution. We analysed the phenotypic and genotypic data of ∼2,300 seedlings grown in a common garden. We focused on 11 potentially adaptive traits with significant heritabilities (Gauzere et al. 2016) and tested for signature of local selection on quantitative trait differentiation. We then identified the offspring likely originating from local or distant pollen immigration events and quantified the role of gene flow in increasing locally the additive variance of traits under selection. We found a significant signal of adaptive differentiation among plots separated by less than one kilometre, with local selection acting on growth and phenological traits. We found that trees in the plots at high elevation, experiencing the lowest temperature conditions, flushed earlier and had a higher height and diameter growth in our common garden than trees from the plot at low elevation. Beech populations originating from higher longitude or elevation have also been shown to be genetically earlier in provenance tests, suggesting that these populations evolved phenological traits promoting a longer vegetation period. At this southern margin of the species, the reduced allocation to stem growth at the low elevation plot is likely an adaptive response to drought, which has previously been described by comparing marginal vs central beech populations. Consistently with theoretical expectations, our results suggest a beneficial effect of pollen dispersal by increasing the genetic diversity for these locally differentiated traits. These effects were quantitatively high, with more than twice higher genetic variance for immigrant than local offspring, although with large standard errors around estimates. Our results highlight that local selection is an important evolutionary force in natural tree populations, and provide a strong evidence that adaptive genetic differentiation can occur despite high gene flow. For the two genetically differentiated traits, our analyses suggested a beneficial effect of pollen dispersal by increasing genetic diversity after one episode of reproduction. The findings suggest that conservation and management interventions to facilitate movement of gametes along short ecological gradients would boost genetic diversity of individual tree populations, and thereby enhance their adaptive potential

Where is the optimum? Predicting the variation of selection along climatic gradients and the adaptive value of plasticity. A case study on tree phenology

International audienceMany theoretical models predict when genetic evolution and phenotypic plasticity allow adaptation to changing environmental conditions. These models generally assume stabilizing selection around some optimal phenotype. We however often ignore how optimal phenotypes change with the environment, which limit our understanding of the adaptive value of phenotypic plasticity. Here, we propose an approach based on our knowledge of the causal relationships between climate, adaptive traits, and fitness to further these questions. This approach relies on a sensitivity analysis of the process-based model Phenofit, which mathematically formalizes these causal relationships, to predict fitness landscapes and optimal budburst dates along elevation gradients in three major European tree species. Variation in the overall shape of the fitness landscape and resulting directional selection gradients were found to be mainly driven by temperature variation. The optimal budburst date was delayed with elevation, while the range of dates allowing high fitness narrowed and the maximal fitness at the optimum decreased. We also found that the plasticity of the budburst date should allow tracking the spatial variation in the optimal date, but with variable mismatch depending on the species, ranging from negligible mismatch in fir, moderate in beech, to large in oak. Phenotypic plasticity would therefore be more adaptive in fir and beech than in oak. In all species, we predicted stronger directional selection for earlier budburst date at higher elevation. The weak selection on budburst date in fir should result in the evolution of negligible genetic divergence, while beech and oak would evolve counter-gradient variation, where genetic and environmental effects are in opposite directions. Our study suggests that theoretical models should consider how whole fitness landscapes change with the environment. The approach introduced here has the potential to be developed for other traits and species to explore how populations will adapt to climate change

Drought-induced decline and mortality of silver fir differ among three sites in Southern France

Abstract : Context : In the Mediterranean area, numerous decline and mortality processes have been reported during recent decades, affecting forest dynamics. They are likely due to increases in summer drought severity and therefore especially affect drought-sensitive species, such as silver fir (Abies alba Mill.). Aims and methods : To understand the relationships between tree growth, crown condition and mortality probability, radial growth trends of healthy, declining (showing crown damages) and dead trees were compared using tree-ring analysis. Factors involved in determining this mortality were also examined at the plot and tree level using altitudinal gradients on three contrasted sites in southeastern France. Results : Individuals with higher inter-annual variability in growth were more prone to dieback. At two sites, dead trees displayed lower growth rates over their entire lifetime, while, on the last site, their juvenile growth rate was higher. Trees with crown damage had higher growth rates than healthy trees on one site, and their radial growth trends over time always differed from those of dead trees. Mortality and crown damage were little related to altitude, but strongly differed between sites and among plots underlining the importance of local edaphic and topographic conditions. Conclusion : These results suggest that the relationships among mortality probability, crown condition and growth can differ among sites, and highlight the impact of soil conditions and the need to assess them in tree mortality studies

A study of karst hydrosystem recharge at the parcel scale, using modeling and correlation analysis - Low noise underground laboratory of Rustrel site

La caractérisation des flux d’eaux qui rechargent réellement les hydrosystèmes souterrains reste un frein à la compréhension du fonctionnement hydrogéologique des milieux souterrains. Lors d’événements pluvieux, quelle part de l’eau est évapo-transpirée ? Quelle part est temporairement stockée dans le sol ? Ces incertitudes sont particulièrement fortes dans le cas de la recharge des milieux hétérogènes tel que le karst. En général, les calculs de recharge des hydrosystèmes karstiques se basent sur une représentation simplifiée de l’évapotranspiration qui considère seulement le climat et pas le fonctionnement de la végétation. Dans cette étude, un modèle de végétation permettant de simuler les transferts d’eaux entre le sol et l’atmosphère en contexte forestier (le modèle CASTANEA), a été appliqué à une parcelle de Chêne vert. L’infiltration efficace (un indicateur de la recharge) estimé avec CASTANEA a été comparée à celle estimée par des approches classiques ainsi qu’à des séries long terme de flux d’eaux souterraines (9 années). Les résultats de cette analyse révèlent que l’infiltration efficace modélisée à partir d’un modèle de végétation comme CASTANEA est plus satisfaisante que les approches classiques ne tenant pas compte du fonctionnement de la végétation. Ce travail ouvre des perspectives intéressantes pour mieux tenir compte du fonctionnement de la végétation et de l’usage du sol sur la recharge des hydrosystèmes karstiques.Assessing the recharge of underground hydrosystems remains an obstacle to understand their hydrologeological functioning. During a rain event, which part of the rain is evapotranspired ? And how much is temporarily stored within the soil ? These questions are particularly relevant in heterogeneous media such as karst hydrosystems. Currently, the models used to compute recharge of karst hydrosystems, rely on simplistic formulations of evapotranspiration that do not account for vegetation functioning. In this study, we used the vegetation process based model CASTANEA, which is designed to compute water transfer between soil, plant and atmosphere. We computed effective infiltration (an index of recharge) with CASTANEA and with other classical approach (based on precipitation minus ETP), and for a welldocumented holm oak site in Provence. Our results provide evidences that effective infiltration computed with CASTANEA yield more satisfactory correlation with measured outflow than simulations based on the classical approach. Our results provide a promising way to improve the simulation of karst hydrosystem recharge

Towards a common methodology for developing logistic tree mortality models based on ring-width data

Tree mortality is a key process shaping forest dynamics. Thus, there is a growing need for indicators of the likelihood of tree death. During the last decades, an increasing number of tree-ring based studies have aimed to derive growth-mortality functions, mostly using logistic models. The results of these studies, however, are difficult to compare and synthesize due to the diversity of approaches used for the sampling strategy (number and characteristics of ‘alive’ and ‘death’ observations), the type of explanatory growth variables included (level, trend, etc.), and the length of the time-window (number of years preceding the alive/death observation) that maximized the discrimination ability of each growth variable. Here, we assess the implications of key methodological decisions when developing tree-ring based growth-mortality relationships using logistic mixed-effects regression models. As examples we use published tree-ring datasets from Abies alba (13 different sites), Nothofagus dombeyi (one site) and Quercus petraea (one site). Our approach is based on a constant sampling size and aims at (1) assessing the dependency of growth-mortality relationships on the statistical sampling scheme used; (2) determining the type of explanatory growth variables that should be considered; and (3) identifying the best length of the time window used to calculate them. The performance of tree-ring based mortality models was reasonably high for all three species (Area Under the receiving operator characteristics Curve: AUC > 0.7). Growth level variables were the most important predictors of mortality probability for two species (A. alba, N. dombeyi), while growth-trend variables need to be considered for Q. petraea. In addition, the length of the time window used to calculate each growth variable was highly uncertain and depended on the sampling scheme, as some growth-mortality relationships varied with tree age. The present study accounts for the main sampling-related biases to determine reliable species-specific growth-mortality relationships. Our results highlight the importance of using a sampling strategy that is consistent with the research question. Moving towards a common methodology for developing reliable growth-mortality relationships is an important step towards improving our understanding of tree mortality across species and its representation in dynamic vegetation models

Early-Warning Signals of Individual Tree Mortality Based on Annual Radial Growth

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter- annual growth variability and a decrease in growth synchrony in the last similar to 20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.Peer reviewe

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery

International audienceThe relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential