Species distribution models may misdirect assisted migration: insights from the introduction of Douglas-fir to Europe.

Species distribution models (SDMs), which statistically relate species occurrence to climatic variables, are widely used to identify areas suitable for species growth under future climates and to plan for assisted migration. When SDMs are projected across times or spaces, it is assumed that species climatic requirements remain constant. However, empirical evidence supporting this assumption is rare, and SDM predictions could be biased. Historical human-aided movements of tree species can shed light on the reliability of SDM predictions in planning for assisted migration. We used Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), a North American conifer introduced into Europe during the mid-19th century, as a case-study to test niche conservatism. We combined transcontinental data sets of Douglas-fir occurrence and climatic predictors to compare the realized niches between native and introduced ranges. We calibrated a SDM in the native range and compared areas predicted to be climatically suitable with observed presences. The realized niches in the native and introduced ranges showed very limited overlap. The SDM calibrated in North America had very high predictive power in the native range, but failed to predict climatic suitability in Europe where Douglas-fir grows in climates that have no analogue in the native range. We review the ecological mechanisms and silvicultural practices that can trigger such shifts in realized niches. Retrospective analysis of tree species introduction revealed that the assumption of niche conservatism is erroneous. As a result, distributions predicted by SDM are importantly biased. There is a high risk that assisted migration programs may be misdirected and target inadequate species or introduction zones.

Does long-term drought or repeated defoliation affect seasonal leaf N cycling in young beech trees?

Forest trees adopt effective strategies to optimize nitrogen (N) use through internal N recycling. In the context of more recurrent environmental stresses due to climate change, the question remains of whether increased frequency of drought or defoliation threatens this internal N recycling strategy. We submitted 8-year-old beech trees to 2 years of either severe drought (Dro) or manual defoliation (Def) to create a state of N starvation. At the end of the second year before leaf senescence, we labeled the foliage of the Dro and Def trees, as well as that of control (Co) trees, with 15N-urea. Leaf N resorption, winter tree N storage (total N, 15N, amino acids, soluble proteins) and N remobilization in spring were evaluated for the three treatments. Defoliation and drought did not significantly impact foliar N resorption or N concentrations in organs in winter. Total N amounts in Def tree remained close to those in Co tree, but winter N was stored more in the branches than in the trunk and roots. Total N amount in Dro trees was drastically reduced (-55%), especially at the trunk level, but soluble protein concentrations increased in the trunk and fine roots compared with Co trees. During spring, 15N was mobilized from the trunk, branches and twigs of both Co and Def trees to support leaf growth. It was only provided through twig 15N remobilization in the Dro trees, thus resulting in extremely reduced Dro leaf N amounts. Our results suggest that stress-induced changes occur in N metabolism but with varying severity depending on the constraints: within-tree 15N transport and storage strategy changed in response to defoliation, whereas a soil water deficit induced a drastic reduction of the N amounts in all the tree organs. Consequently, N dysfunction could be involved in drought-induced beech tree mortality under the future climate.

Short-term nitrogen dynamics are impacted by defoliation and drought in Fagus sylvatica L. branches.

The predicted recurrence of adverse climatic events such as droughts, which disrupt nutrient accessibility for trees, could jeopardize the nitrogen (N) metabolism in forest trees. Internal tree N cycling capacities are crucial to ensuring tree survival but how the N metabolism of forest trees responds to intense, repeated environmental stress is not well known. For 2 years, we submitted 9-year-old beech (Fagus sylvatica L.) trees to either a moderate or a severe prolonged drought or a yearly removal of 75% of the foliage to induce internal N cycling changes. During the second year of stress, in spring and summer, we sprayed 15N-urea on the leaves (one branch per tree). Then, for 14 days, we traced the 15N dynamics through the leaves, into foliar proteins and into the branch compartments (leaves and stems segments), as well as its long-distance transfer from the labeled branches to the tree apical twigs. Defoliation caused a short- and mid-term N increase in the leaves, which remained the main sink for N. Whatever the treatment and the date, most of the leaf 15N stayed in the leaves and was invested in soluble proteins (60-68% of total leaf N). 15N stayed more in the proximal part of the branch in response to drought compared with other treatments. The long-distance transport of N was maintained even under harsh drought, highlighting efficient internal N recycling in beech trees. Under extreme constraints creating an N and water imbalance, compensation mechanisms operated at the branch level in beech trees and allowed them (i) to maintain leaf N metabolism and protein synthesis and (ii) to ensure the seasonal short- and long-distance transfer of recycled leaf N even under drastic water shortage conditions.

Modeling climate change effects on the potential production of French plains forests at the sub-regional level.

We modeled the effects of climate change and two forest management scenarios on wood production and forest carbon balance in French forests using process-based models of forest growth. We combined data from the national forest inventory and soil network survey, which were aggregated over a 50 x 50-km grid, i.e., the spatial resolution of the climate scenario data. We predicted and analyzed the climate impact on potential forest production over the period 1960-2100. All models predicted a slight increase in potential forest yield until 2030-2050, followed by a plateau or a decline around 2070-2100, with overall, a greater increase in yield in northern France than in the south. Gross and net primary productivities were more negatively affected by soil water and atmospheric water vapor saturation deficits in western France because of a more pronounced shift in seasonal rainfall from summer to winter. The rotation-averaged values of carbon flux and production for different forest management options were estimated during four years (1980, 2015, 2045 and 2080). Predictions were made using a two-dimensional matrix covering the range of local soil and climate conditions. The changes in ecosystem fluxes and forest production were explained by the counterbalancing effect of rising CO2 concentration and increasing water deficit. The effect of climate change decreased with rotation length from short rotations with high production rates and low standing biomasses to long rotations with low productivities and greater standing biomasses. Climate effects on productivity, both negative and positive, were greatest on high fertility sites. Forest productivity in northern France was enhanced by climate change, increasingly from west to east, whereas in the southwestern Atlantic region, productivity was reduced by climate change to an increasing degree from west to east.

Estimation of deciduous forest leaf area index using direct and indirect methods.

This study evaluated one semi-direct and three indirect methods for estimating leaf area index (LAI) by comparing these estimates with direct estimates derived from litter collection. The semi-direct method uses a thin metallic needle to count a number of contacts across fresh litter layers. One indirect method is based on the penetration of diffuse global radiation measured over the course of a day. The second indirect method uses the LAI-2000 plant canopy analyser (PCA) which measures diffuse light penetration from five different sky sectors simultaneously. The third indirect method uses the "Demon" portable light sensor to measure the penetration of direct beam sunlight at different zenith angles over the course of half a day. The Poisson model of gap frequency was applied to estimate plant area index (PAI) from observed transmittances using the second and third methods. Litter collection from 11 temperate decidous forests gave values of LAI ranging from 1.7 to 7.5. Estimates based on the needle method showed a significant linear relationship with LAI values obtained from litter collections but were systematically lower (by 6-37%). PAI estimates using all three indirect techniques (fixed light sensor system, LAI-2000 and Demon) showed a strong linear relationship with LAI derived from litter collection. Differences, averaged over all forest stands, between PAI estimates from each of the three indirect methods and LAI from litter collections were below 2%. If we consider that LAI=PAI-WAI (wood area index) then, all three indirect methods underestimated LAI by an additional factor close to the value of WAI. One reason could be a local clumping of architectural canopy components: in particular, the spatial dispositions of branchlets and leaves are not independent, leading to a non-random relationship between the distributions of these two canopy components.

Comparison of water-use efficiency of seedlings from two sympatric oak species: genotype x environment interactions.

Seedlings of two sympatric oak species, Quercus robur L. and Quercus petraea (Matt.) Liebl., were grown in common garden conditions to test for potential interspecific differences in intrinsic water-use efficiency (WUE). Intrinsic water-use efficiency was estimated based on carbon isotope composition of shoots (delta13C) and on gas exchange measurements (ratio of net CO2 assimilation rate to stomatal conductance (A/g(sw))). In addition, genotype x environment interactions were tested by subjecting the seedlings to four irradiance treatments (8, 18, 48 and 100% of incident solar irradiance) imposed by neutral shading nets, and, in the 100% irradiance treatment, two watering regimes. In all treatments, initial growth of Q. robur was faster than that of Q. petraea. In both species, there was a tight correlation between delta13C and A/g(sw). Intrinsic water-use efficiency increased with increasing irradiance (almost doubling from 8 to 100% irradiance), and this effect paralleled the increase in A with increasing irradiance. In full sun, WUE of Q. petraea seedlings was 10-15% higher than in Q. robur seedlings, with the difference attributable to a difference between the species in g(sw). The interspecific difference in WUE was maintained during drought, despite the appreciable increase in WUE and decrease in growth imposed by drought. No interspecific differences in WUE were observed at low irradiances, suggesting a strong genotype x environment interaction for WUE. These findings confirm the existence of interspecific genetic differences in WUE, but also show that there is large intraspecific variability and plasticity in WUE. The initially greater height and biomass increments in Q. robur seedlings illustrate the ability of this species to out-compete Q. petraea in the early stages of forest regeneration. For adult trees growing in closed canopies, the high WUE of Q. petraea may contribute significantly to its survival during dry years, whereas the low WUE of Q. robur may account for the frequently observed declines in adult trees of this species following drought.

Nitrogen sources for current-year shoot growth in 50-year-old sessile oak trees: an in situ (15)N labeling approach.

We used long-term in situ (15)N labeling of the soil to investigate the contribution of the two main nitrogen (N) sources (N uptake versus N reserves) to sun shoot growth from bud burst to full leaf expansion in 50-year-old sessile oaks. Recovery of (15)N by growing compartments (leaves, twigs and buds) and presence of (15)N in phloem sap were checked weekly. During the first 2 weeks following bud burst, remobilized N contributed ~90% of total N in growing leaves and twigs. Nitrogen uptake from the soil started concomitantly with N remobilization but contributed only slightly to bud burst. However, the fraction of total N due to N uptake increased markedly once bud burst had occurred, reaching 27% in fully expanded leaves and 18% in developed twigs. In phloem sap, the (15)N label appeared a few days after the beginning of labeling and increased until the end of bud burst, and then decreased at full leaf expansion in June. Of all the shoot compartments, leaves attracted most of the absorbed N, which accounted for 68% of new N in shoots, whereas twigs and new buds accounted for only 28 and 3%, respectively. New N allocated to leaves increased from unfolding to full expansion as total N concentration in the leaves decreased. Our results underline the crucial role played by stored N in rapid leaf growth and in the sustained growth of oak trees. Any factors that reduce N storage in autumn may therefore impair spring shoot growth.

Ground-based measurements of leaf area index: a review of methods, instruments and current controversies.

Leaf area index (LAI) is the total one-sided area of leaf tissue per unit ground surface area. It is a key parameter in ecophysiology, especially for scaling up the gas exchange from leaf to canopy level. It characterizes the canopy-atmosphere interface, where most of the energy fluxes exchange. It is also one of the most difficult to quantify properly, owing to large spatial and temporal variability. Many methods have been developed to quantify LAI from the ground and some of them are also suitable for describing other structural parameters of the canopy. This paper reviews the direct and indirect methods, the required instruments, their advantages, disadvantages and accuracy of the results. Analysis of the literature shows that most cross-validations between direct and indirect methods have pointed to a significant underestimation of LAI with the latter techniques, especially in forest stands. The two main causes for the discrepancy, clumping and contribution of stem and branches, are discussed and some recent theoretical or technical solutions are presented as potential improvements to reduce bias or discrepancies. The accuracy, sampling strategy and spatial validity of the LAI measurements have to be assessed for quality assurance of both the measurement and the modelling purposes of all the LAI-dependent ecophysiological and biophysical processes of canopies.