Revealing legacy effects of extreme droughts on tree growth of oaks across the Northern Hemisphere.

Forests are undergoing increasing risks of drought-induced tree mortality. Species replacement patterns following mortality may have a significant impact on the global carbon cycle. Among major hardwoods, deciduous oaks (Quercus spp.) are increasingly reported as replacing dying conifers across the Northern Hemisphere. Yet, our knowledge on the growth responses of these oaks to drought is incomplete, especially regarding post-drought legacy effects. The objectives of this study were to determine the occurrence, duration, and magnitude of legacy effects of extreme droughts and how that vary across species, sites, and drought characteristics. The legacy effects were quantified by the deviation of observed from expected radial growth indices in the period 1940-2016. We used stand-level chronologies from 458 sites and 21 oak species primarily from Europe, north-eastern America, and eastern Asia. We found that legacy effects of droughts could last from 1 to 5 years after the drought and were more prolonged in dry sites. Negative legacy effects (i.e., lower growth than expected) were more prevalent after repetitive droughts in dry sites. The effect of repetitive drought was stronger in Mediterranean oaks especially in Quercus faginea. Species-specific analyses revealed that Q. petraea and Q. macrocarpa from dry sites were more negatively affected by the droughts while growth of several oak species from mesic sites increased during post-drought years. Sites showing positive correlations to winter temperature showed little to no growth depression after drought, whereas sites with a positive correlation to previous summer water balance showed decreased growth. This may indicate that although winter warming favors tree growth during droughts, previous-year summer precipitation may predispose oak trees to current-year extreme droughts. Our results revealed a massive role of repetitive droughts in determining legacy effects and highlighted how growth sensitivity to climate, drought seasonality and species-specific traits drive the legacy effects in deciduous oak species.

Climate sensitivity and drought seasonality determine post-drought growth recovery of Quercus petraea and Quercus robur in Europe.

Recent studies have identified strong relationships between delayed recovery of tree growth after drought and tree mortality caused by subsequent droughts. These observations raise concerns about forest ecosystem services and post-drought growth recovery given the projected increase in drought frequency and extremes. For quantifying the impact of extreme droughts on tree radial growth, we used a network of tree-ring width data of 1689 trees from 100 sites representing most of the distribution of two drought tolerant, deciduous oak species (Quercus petraea and Quercus robur). We first examined which climatic factors and seasons control growth of the two species and if there is any latitudinal, longitudinal or elevational trend. We then quantified the relative departure from pre-drought growth during droughts, and how fast trees were able to recover the pre-drought growth level. Our results showed that growth was more related to precipitation and climatic water balance (precipitation minus potential evapotranspiration) than to temperature. However, we did not detect any clear latitudinal, longitudinal or elevational trends except a decreasing influence of summer water balance on growth of Q. petraea with latitude. Neither species was able to maintain the pre-drought growth level during droughts. However, both species showed rapid recovery or even growth compensation after summer droughts but displayed slow recovery in response to spring droughts where none of the two species was able to fully recover the pre-drought growth-level over the three post-drought years. Collectively, our results indicate that oaks which are considered resilient to extreme droughts have also shown vulnerability when droughts occurred in spring especially at sites where long-term growth is not significantly correlated with climatic factors. This improved understanding of the role of drought seasonality and climate sensitivity of sites is key to better predict trajectories of post-drought growth recovery in response to the drier climate projected for Europe.

The within-population variability of leaf spring and autumn phenology is influenced by temperature in temperate deciduous trees.

Leaf phenology is a major driver of ecosystem functioning in temperate forests and a robust indicator of climate change. Both the inter-annual and inter-population variability of leaf phenology have received much attention in the literature; in contrast, the within-population variability of leaf phenology has been far less studied. Beyond its impact on individual tree physiological processes, the within-population variability of leaf phenology can affect the estimation of the average budburst or leaf senescence dates at the population scale. Here, we monitored the progress of spring and autumn leaf phenology over 14 tree populations (9 tree species) in six European forests over the period of 2011 to 2018 (yielding 16 site-years of data for spring, 14 for autumn). We monitored 27 to 512 (with a median of 62) individuals per population. We quantified the within-population variability of leaf phenology as the standard deviation of the distribution of individual dates of budburst or leaf senescence (SDBBi and SDLSi, respectively). Given the natural variability of phenological dates occurring in our tree populations, we estimated from the data that a minimum sample size of 28 (resp. 23) individuals, are required to estimate SDBBi (resp. SDLSi) with a precision of 3 (resp. 7) days. The within-population of leaf senescence (average SDLSi = 8.5 days) was on average two times larger than for budburst (average SDBBi = 4.0 days). We evidenced that warmer temperature during the budburst period and a late average budburst date were associated with a lower SDBBi, as a result of a quicker spread of budburst in tree populations, with a strong species effect. Regarding autumn phenology, we observed that later senescence and warm temperatures during the senescence period were linked with a high SDLSi, with a strong species effect. The shares of variance explained by our models were modest suggesting that other factors likely influence the within-population variation in leaf phenology. For instance, a detailed analysis revealed that summer temperatures were negatively correlated with a lower SDLSi.

Which oak provenances for the 22nd century in Western Europe? Dendroclimatology in common gardens.

The current distribution area of the two sympatric oaks Quercus petraea and Q. robur covers most of temperate Western Europe. Depending on their geographic location, populations of these trees are exposed to different climate constraints, to which they are adapted. Comparing the performances of trees from contrasting populations provides the insight into their expected resilience to future climate change required for forest management. In this study, the descendants of 24 Q. petraea and two Q. robur provenances selected from sites throughout Europe were grown for 20 years in three common gardens with contrasting climates. The 2420 sampled trees allowed the assessments of the relationship between radial growth and climate. An analysis of 15-year chronologies of ring widths, with different combinations of climate variables, revealed different response patterns between provenances and between common gardens. As expected, provenances originating from sites with wet summers displayed the strongest responses to summer drought, particularly in the driest common garden. All provenances displayed positive significant relationships between the temperature of the previous winter and radial growth when grown in the common garden experiencing the mildest winter temperatures. Only eastern provenances from continental cold climates also clearly expressed this limitation of growth by cold winter temperatures in the other two common gardens. However, ecological distance, calculated on the basis of differences in climate between the site of origin and the common garden, was not clearly related to the radial growth responses of the provenances. This suggests that the gradient of genetic variability among the selected provenances was not strictly structured according to climate gradients. Based on these results, we provide guidelines for forest managers for the assisted migration of Quercus petraea and Q. robur provenances.

Climatically controlled reproduction drives interannual growth variability in a temperate tree species.

Climatically controlled allocation to reproduction is a key mechanism by which climate influences tree growth and may explain lagged correlations between climate and growth. We used continent-wide datasets of tree-ring chronologies and annual reproductive effort in Fagus sylvatica from 1901 to 2015 to characterise relationships between climate, reproduction and growth. Results highlight that variable allocation to reproduction is a key factor for growth in this species, and that high reproductive effort ('mast years') is associated with stem growth reduction. Additionally, high reproductive effort is associated with previous summer temperature, creating lagged climate effects on growth. Consequently, understanding growth variability in forest ecosystems requires the incorporation of reproduction, which can be highly variable. Our results suggest that future response of growth dynamics to climate change in this species will be strongly influenced by the response of reproduction.

Stand density, tree social status and water stress influence allocation in height and diameter growth of Quercus petraea (Liebl.).

Even-aged forest stands are competitive communities where competition for light gives advantages to tall individuals, thereby inducing a race for height. These same individuals must however balance this competitive advantage with height-related mechanical and hydraulic risks. These phenomena may induce variations in height-diameter growth relationships, with primary dependences on stand density and tree social status as proxies for competition pressure and access to light, and on availability of local environmental resources, including water. We aimed to investigate the effects of stand density, tree social status and water stress on the individual height-circumference growth allocation (Δh-Δc), in even-aged stands of Quercus petraea Liebl. (sessile oak). Within-stand Δc was used as surrogate for tree social status. We used an original long-term experimental plot network, set up in the species production area in France, and designed to explore stand dynamics on a maximum density gradient. Growth allocation was modelled statistically by relating the shape of the Δh-Δc relationship to stand density, stand age and water deficit. The shape of the Δh-Δc relationship shifted from linear with a moderate slope in open-grown stands to concave saturating with an initial steep slope in closed stands. Maximum height growth was found to follow a typical mono-modal response to stand age. In open-grown stands, increasing summer soil water deficit was found to decrease height growth relative to radial growth, suggesting hydraulic constraints on height growth. A similar pattern was found in closed stands, the magnitude of the effect however lowering from suppressed to dominant trees. We highlight the high phenotypic plasticity of growth in sessile oak trees that further adapt their allocation scheme to their environment. Stand density and tree social status were major drivers of growth allocation variations, while water stress had a detrimental effect on height in the Δh-Δc allocation.

Life strategies in intra-annual dynamics of wood formation: example of three conifer species in a temperate forest in north-east France.

We investigated whether timing and rate of growth are related to the life strategies and fitness of three conifer species. Intra-annual dynamics of wood formation, shoot elongation and needle phenology were monitored over 3 years in five Norway spruces (Picea abies (L.) Karst.), five Scots pines (Pinus sylvestris L.) and five silver firs (Abies alba Mill.) grown intermixed. For the three species, the growing season (delimited by cambial activity onset and cessation) lasted about 4 months, while the whole process of wood formation lasted 5-6 months. Needle unfolding and shoot elongation followed the onset of cambial activity and lasted only one-third of the season. Pines exhibited an 'extensive strategy' of cambial activity, with long durations but low growth rates, while firs and spruces adopted an 'intensive strategy' with shorter durations but higher growth rates. We estimated that about 75% of the annual radial increment variability was attributable to the rate of cell production, and only 25% to its duration. Cambial activity rates culminated at the same time for the three species, whereas shoot elongation reached its maximal rate earlier in pines. Results show that species-specific life strategies are recognizable through functional traits of intra-annual growth dynamics. The opposition between Scots pine extensive strategy and silver fir and Norway spruce intensive strategy supports the theory that pioneer species are greater resource expenders and develop riskier life strategies to capture resources, while shade-tolerant species utilize resources more efficiently and develop safer life strategies. Despite different strategies, synchronicity of the maximal rates of cambial activity suggests a strong functional convergence between co-existing conifer species, resulting in head-on competition for resources.

Simulating phenological shifts in French temperate forests under two climatic change scenarios and four driving global circulation models.

After modeling the large-scale climate response patterns of leaf unfolding, leaf coloring and growing season length of evergreen and deciduous French temperate trees, we predicted the effects of eight future climate scenarios on phenological events. We used the ground observations from 103 temperate forests (10 species and 3,708 trees) from the French Renecofor Network and for the period 1997-2006. We applied RandomForest algorithms to predict phenological events from climatic and ecological variables. With the resulting models, we drew maps of phenological events throughout France under present climate and under two climatic change scenarios (A2, B2) and four global circulation models (HadCM3, CGCM2, CSIRO2 and PCM). We compared current observations and predicted values for the periods 2041-2070 and 2071-2100. On average, spring development of oaks precedes that of beech, which precedes that of conifers. Annual cycles in budburst and leaf coloring are highly correlated with January, March-April and October-November weather conditions through temperature, global solar radiation or potential evapotranspiration depending on species. At the end of the twenty-first century, each model predicts earlier budburst (mean: 7 days) and later leaf coloring (mean: 13 days) leading to an average increase in the growing season of about 20 days (for oaks and beech stands). The A2-HadCM3 hypothesis leads to an increase of up to 30 days in many areas. As a consequence of higher predicted warming during autumn than during winter or spring, shifts in leaf coloring dates appear greater than trends in leaf unfolding. At a regional scale, highly differing climatic response patterns were observed.

Variation in cold hardiness and carbohydrate concentration from dormancy induction to bud burst among provenances of three European oak species.

Although cold hardiness is known to be a major determinant of tree species distribution, its dynamics and the factors that regulate it remain poorly understood. Variation in cold hardiness and carbohydrate concentration, from dormancy induction until bud burst, were investigated in populations of two deciduous (Quercus robur L. and Quercus pubescens Willd.) and one evergreen (Quercus ilex L.) European oak. Mean cold hardiness values in January were -56, -45 and -27 degrees C for Q. robur, Q. pubescens and Q. ilex, respectively. Soluble carbohydrate concentrations were closely related to instantaneous cold hardiness, estimated by the electrolyte leakage method, whereas total carbohydrate concentration was related to maximum cold hardiness. Both cold hardiness and carbohydrate concentration showed a close linear relationship with temperatures at the location of the sampled population. Our results show that temporal variation in both the inter- and intraspecific cold hardiness in European oaks can be related to variations in the concentrations of soluble carbohydrates and that these relationships appear to be driven by temperature.