Will the CO2 fertilization effect in forests be offset by reduced tree longevity?

Experimental studies suggest that tree growth is stimulated in a greenhouse atmosphere, leading to faster carbon accumulation (i.e., a higher rate of gap filling). However, higher growth may be coupled with reduced longevity, thus leading to faster carbon release (i.e., a higher rate of gap creation). The net effect of these two counteracting processes is not known. We quantify this net effect on aboveground carbon stocks using a novel combination of data sets and modeling. Data on maximum growth rate and maximum longevity of 141 temperate tree species are used to derive a relationship between growth stimulation and changes in longevity. We employ this relationship to modify the respective parameter values of tree species in a forest succession model and study aboveground biomass in a factorial design of growth stimulation×reduced maximum longevity at multiple sites along a climate gradient from the cold to the dry treeline. The results show that (1) any growth stimulation at the tree level leads to a disproportionately small increase of stand biomass due to negative feedback effects, even in the absence of reduced longevity; (2) a reduction of tree longevity tends to offset the growth-related biomass increase; at the most likely value of reduced longevity, the net effect is very close to zero in most multi- and single-species simulations; and (3) when averaging the response across all sites to mimic a "landscape-level” response, the net effect is close to zero. Thus, it is important to consider ecophysiological responses with their linkage to demographic processes in forest trees if one wishes to avoid erroneous inference at the ecosystem level. We conclude that any CO2 fertilization effect is quite likely to be offset by an associated reduction in the longevity of forest trees, thus strongly reducing the carbon mitigation potential of temperate forest

Daily Maximum Temperatures Induce Lagged Effects on Leaf Unfolding in Temperate Woody Species Across Large Elevational Gradients

The timing of leaf unfolding in temperate woody species is predominantly controlled by the seasonal course of temperature in late winter and early spring. However, quantifying lagged temperature effects on spring phenology is still challenging. Here, we aimed at investigating lagged and potentially non-linear effects of daily maximum temperatures on the probability of leaf unfolding in temperate woody species growing across large elevational gradients. We analyzed 5280 observations of leaf-out time of four tree species (European beech, horse chestnut, European larch, Norway spruce) and one shrub species (common hazel) that were recorded by volunteers over 40 years at 42 locations in Switzerland. We used a case-crossover sampling design to match leaf-out dates with control dates (i.e., dates before or after leaf-out), and analyzed these data with conditional logistic regression accounting for lagged temperature effects over 60 days. Multivariate meta-analyses were used to synthesize lagged temperature and elevational effects on leaf unfolding across multiple phenological stations. Temperature effects on the probability of leaf unfolding were largest at relatively short lags (i.e., within ca. 10 days) and decreased with increasing lags. Short- to mid-term effects (i.e., within ca. 10 to 20 days) were larger for late-leafing species known to be photoperiod-sensitive (beech, Norway spruce). Temperature effects increased for the broadleaved species (horse chestnut, hazel, beech) with decreasing elevation, particularly within ca. 10 to 40 days, i.e., leaf unfolding occurs more rapidly at low elevations for a given daily maximum temperature. Our novel findings provide evidence of cumulative and long-term temperature effects on leaf unfolding, whereby the efficiency of relatively high temperatures to trigger leaf-out becomes higher shortly before bud burst. These lagged associations between temperature and leaf unfolding improve our understanding of phenological responses across temperate woody species with differing ecological requirements that occur along elevational gradients

Effects of growth rates, tree morphology and site conditions on longevity of Norway spruce in the northern Swiss Alps

Longevity of trees is known to be associated with growth rates, but also with tree morphology and spatial influences. However, very little quantitative information is available on the effects of these biotic and abiotic influences on maximum ages of trees. The objectives of this study were to investigate the trade-off between longevity and growth rates of Norway spruce (Picea abies) and to quantify the effects of tree morphology and abiotic site conditions on longevity of this species. Data were collected along different topographical and climatic gradients in a 20×25km study area in the northern part of the Swiss Alps (Glarus). The ages of the more than 100 sampled dead Norway spruces ranged between 50 and 367years. Longevity of these trees was negatively related to tree growth, i.e. slow-growing trees tended to grow older than fast-growing trees. Tree height was positively associated with longevity for both upper and lower storey trees. Longevity of lower storey trees was increased with large crown diameter, but decreased with long crown length. Upper storey trees growing at higher altitude tended to get older than at lower altitude. We conclude that the combined effects of growth rates, variability in site conditions and different traits of tree morphology determine tree longevity of Norway spruce in the Swiss Alps. Because longevity is tightly linked to mortality rates of tree populations, our study may improve our understanding of long-term processes of forest dynamics under current and future climat

Analysis and modelling of tree succession on a recent rockslide deposit

Dendroecological methods were combined with vegetation and soil mapping to study recolonisation of European larch (Larix decidua), silver birch (Betula pendula) and Norway spruce (Picea abies) on a recently formed rockslide cone (deposit of 30×106m3) in the Valais Alps (Switzerland). Tree density and tree height were predicted with regression models that we derived using an information-theoretic model selection approach. Results demonstrate that the deposits of the 1991 rockslide have been colonised relatively rapidly with larch (ecesis time 2years), birch (5years) and spruce (2years). Most of the colonisation occurred 5-11years following the rockslide. Clast size was the primary factor driving tree colonisation with the highest tree densities found on plots with mainly smaller (<30cm) clast sizes. Tree height was affected by a combination of different influences, with tree age and tree density showing the most obvious effects. This study demonstrates how dendroecological methods allow reconstruction of spatio-temporal patterns of tree succession on rockslides, which may ultimately facilitate a more accurate dating of similar landforms of unknown ag

Precision and accuracy of tree-ring-based death dates of mountain pines in the Swiss National Park

Key message : Mountain pines in the Swiss National Park show evidence of partial cambial mortality, which affects the precision of tree-ring-based death dates, followed by lagged crown mortality. Abstract: The time of tree death is commonly reconstructed by dating the outermost ring of tree-ring series. However, due to the occurrence of partial cambial mortality, the date of the outermost tree ring may vary between different locations on the tree stem. Furthermore, a tree may continue to live following the formation of the most recent tree ring. In this study, we quantified precision and accuracy of tree-ring-based death dates from 229 dead mountain pines (Pinus montana) from a 28km2 study area in the Swiss National Park. For almost two-thirds of the trees, a maximum difference of just 0-4years between the dates of cambial mortality from three increment cores was observed, however, for a few trees the difference reached 30-65years. Higher maximum differences between the dates of cambial mortality are expected for trees on steep slopes, for old trees or for trees that died a long time ago. For 84% of dead mountain pines, which were sampled in a permanent sample plot with 2-year remeasurement intervals, the difference between the date of observed crown mortality and the death date determined from three cores was 0-5years. Sampling two or just one core per tree decreases the accuracy of tree-ring-based death dates. Based on the findings of our study, we recommend a prior assessment of the precision and accuracy of tree-ring-based death dates for any dendroecological study dealing with the reconstruction of tree mortalit

Do small-grain processes matter for landscape scale questions? Sensitivity of a forest landscape model to the formulation of tree growth rate

Process-based forest landscape models are valuable tools for testing basic ecological theory and for projecting how forest landscapes may respond to climate change and other environmental shifts. However, the ability of these models to accurately predict environmentally-induced shifts in species distributions as well as changes in forest composition and structure is often contingent on the phenomenological representation of individual-level processes accurately scaling-up to landscape-level community dynamics. We use a spatially explicit landscape forest model (LandClim) to examine how three alternative formulations of individual tree growth (logistic, Gompertz, and von Bertalanffy) influence model results. Interactions between growth models and landscape characteristics (landscape heterogeneity and disturbance intensity) were tested to determine in what type of landscape simulation results were most sensitive to growth model structure. We found that simulation results were robust to growth function formulation when the results were assessed at a large spatial extent (landscape) and when coarse response variables, such as total forest biomass, were examined. However, results diverged when more detailed response variables, such as species composition within elevation bands, were considered. These differences were particularly prevalent in regions that included environmental transition zones where forest composition is strongly driven by growth-dependent competition. We found that neither landscape heterogeneity nor the intensity of landscape disturbances accentuated simulation sensitivity to growth model formulation. Our results indicate that at the landscape extent, simulation results are robust, but the reliability of model results at a finer resolution depends critically on accurate tree growth function

Drought as an Inciting Mortality Factor in Scots Pine Stands of the Valais, Switzerland

During the 20th century, high mortality rates of Scots pine (Pinus silvestris L.) have been observed over large areas in the Rhône valley (Valais, Switzerland) and in other dry valleys of the European Alps. In this study, we evaluated drought as a possible inciting factor of Scots pine decline in the Valais. Averaged tree-ring widths, standardized tree-ring series, and estimated annual mortality risks were related to a drought index. Correlations between drought indices and standardized tree-ring series from 11 sites showed a moderate association. Several drought years and drought periods could be detected since 1864 that coincided with decreased growth. Although single, extreme drought years had generally a short-term, reversible effect on tree growth, multi-year drought initiated prolonged growth decreases that increased a tree's long-term risk of death. Tree death occurred generally several years or even decades after the drought. In conclusion, drought has a limiting effect on tree growth and acts as a bottleneck event in triggering Scots pine decline in the Valai

Estimating the age-diameter relationship of oak species in Switzerland using nonlinear mixed-effects models

Tree growth plays a key role in forest dynamics, yet little attention has been paid to quantifying tree age-diameter relationships. Predicting diameter growth of oaks is especially important due to their role in nature conservation and adaptive forest management under climate change. Thus, we (1) identified environmental variables that shape age-diameter relationships of oaks and (2) quantified the accuracy of predictions based on these variables. We determined the age-diameter relationship of 243 oaks (Quercus spp.) growing in Switzerland by using tree-ring samples. Nonlinear mixed-effects models based on a modified Chapman-Richards equation were fitted with environmental variables included as covariates. The fixed effects elevation, slope and water-holding capacity were most important in shaping the age-diameter relationships. Lower elevations, steeper slopes, north-facing aspects, higher water-holding capacities and moister summers resulted in larger maximum diameters. For 75% of the oaks, age-diameter relationships predicted by the fixed effects matched fairly well the observations (root mean square error between predictions and observations <6cm); the inclusion of random effects reduced root mean square errors for 86% of the trees. These results suggest that water runoff plays a key role for the age-diameter relationships, accompanied by limiting temperature effects at higher elevations. The fixed effects covered variability in site quality, whereas the random effects included tree-specific deviations from expected age-diameter relationships, potentially due to neighbourhood effects such as stand density and competitio

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