Seasonality matters—The effects of past and projected seasonal climate change on the growth of native and exotic conifer species in Central Europe

Norway spruce is one of the economically most important tree species in Central European forestry. However, its high susceptibility to droughts poses a strong challenge to its cultivation under future conditions with likely more frequent and prolonged droughts and shifts in the seasonal climate. To compensate for expected losses of forest areas suitable for the cultivation of spruce, more drought-tolerant species are required. Silver fir and Douglas fir are two potential candidates, which promise lower drought susceptibility and equal or even higher yield when compared to Norway spruce.Using the Black Forest as a regional case study, we assessed the effects of seasonal climate change, including drought stress, on tree-ring width formation of these three economically relevant conifer species over the last 60 years. In addition, we projected potential species-specific growth changes under different climate change scenarios until 2100.Our results suggest that both silver fir and Douglas fir will possibly experience growth increases in a warmer future climate, as predicted under the 4.5 and 8.5 Representative Concentration Pathway (RCP) climate change scenarios, whereas growth of spruce is expected to decline. Moreover, drought susceptibility in silver fir and Douglas fir is lower than in spruce, as shown for past drought events, and their ability to benefit from milder winters and springs could play a major role in their capacity to compensate for drier summers in the near to mid-term future. This study highlights the need to advance our understanding of the processes that drive drought resistance and resilience in tree species to guide management strategies in the face of climate change.     

Silver fir and Douglas fir are more tolerant to extreme droughts than Norway spruce in south‐western Germany

Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long‐term alternatives for susceptible yet economically important tree species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought‐tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed‐species stands along an altitudinal gradient (400–1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population‐level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change.     

Patterns of drought tolerance in major European temperate forest trees: climatic drivers and levels of variability

The future performance of native tree species under climate change conditions is frequently discussed, since increasingly severe and more frequent drought events are expected to become a major risk for forest ecosystems. To improve our understanding of the drought tolerance of the three common European temperate forest tree species Norway spruce, silver fir and common beech, we tested the influence of climate and tree‐specific traits on the inter and intrasite variability in drought responses of these species. Basal area increment data from a large tree‐ring network in Southern Germany and Alpine Austria along a climatic cline from warm‐dry to cool‐wet conditions were used to calculate indices of tolerance to drought events and their variability at the level of individual trees and populations. General patterns of tolerance indicated a high vulnerability of Norway spruce in comparison to fir and beech and a strong influence of bioclimatic conditions on drought response for all species. On the level of individual trees, low‐growth rates prior to drought events, high competitive status and low age favored resilience in growth response to drought. Consequently, drought events led to heterogeneous and variable response patterns in forests stands. These findings may support the idea of deliberately using spontaneous selection and adaption effects as a passive strategy of forest management under climate change conditions, especially a strong directional selection for more tolerant individuals when frequency and intensity of summer droughts will increase in the course of global climate change.     

Sensitivity of French temperate coniferous forests to climate variability and extreme events (Abies alba,Picea abies and Pinus sylvestris)

Questions: (1) How do extreme climatic events and climate variability influence radial growth of conifers (silver fir, Norway spruce, Scots pine)? (2) How do elevation and soil water capacity (SWC) modulate sensitivity to climate? Location: The sampled conifer stands are in France, in western lowland and mountain forests, at elevations from 400 to 1700 m, and an SWC from 50 to 190 mm. Methods: We established stand chronologies for total ring width, earlywood and latewood width for the 33 studied stands (985 trees in total). Responses to climate were analysed using pointer years and bootstrapped response functions. Principal component analysis was applied to pointer years and response function coefficients in order to elucidate the ecological structure of the studied stands. Results: Extreme winter frosts are responsible for greater growth reductions in silver fir than in Norway spruce, especially at the upper elevation, while Scots pine was the least sensitive species. Exceptional spring droughts caused a notable growth decrease, especially when local conditions were dry (altitude<1000 m and SWC<100 mm for silver fir, western lowlands for Scots pine). Earlywood of silver fir depended on previous September and November and current‐year February temperature, after which current June and July water supply influenced latewood. Earlywood of Norway spruce was influenced by previous September temperature, after which current spring and summer droughts influenced both ring components. In Scots pine, earlywood and latewood depended on the current summer water balance. Local conditions mainly modulated latewood formation. Conclusions: If the climate becomes drier, low‐elevation dry stands or trees growing in western lowlands may face problems, as their growth is highly dependent on soil moisture availability.     

Tree allometry of Douglas fir and Norway spruce on a nutrient-poor and a nutrient-rich site

Research related to the allometric relationships of tree height and projected tree crown area to diameter at breast height was conducted to look at the biological suitability and timber production potential of Douglas fir under the conditions present in central Europe. The dependence of allometric relationships on soil nutrient conditions were described in forest stands of Douglas fir and Norway spruce. The studied sites were climatically similar but differed in soil nutrient availability. A significant difference was found in the allometric relationships of Norway spruce trees from the nutrient poor and nutrient rich site. In contrast to the Norway spruce, there was no significant effect of site fertility on allometric relationships for Douglas fir suggesting that its allocation patterns were less sensitive to site nutrient conditions. Stem growth increment, which was measured weekly during two consecutive seasons for both species, was related to the weather conditions and available soil moisture. Stem growth of Douglas fir began about 2 weeks earlier than in the Norway spruce at both sites. At the nutrient rich site, most of the stem growth of both species occurred at the beginning of the season, while growth at the other site was more evenly distributed throughout the season. Data obtained in this study will be useful for modeling stem growth and analysis of water use efficiency of these two tree species.     

Contrasting resistance and resilience to extreme drought and late spring frost in five major European tree species

Extreme climate events (ECEs) such as severe droughts, heat waves, and late spring frosts are rare but exert a paramount role in shaping tree species distributions. The frequency of such ECEs is expected to increase with climate warming, threatening the sustainability of temperate forests. Here, we analyzed 2,844 tree‐ring width series of five dominant European tree species from 104 Swiss sites ranging from 400 to 2,200 m a.s.l. for the period 1930–2016. We found that (a) the broadleaved oak and beech are sensitive to late frosts that strongly reduce current year growth; however, tree growth is highly resilient and fully recovers within 2 years; (b) radial growth of the conifers larch and spruce is strongly and enduringly reduced by spring droughts—these species are the least resistant and resilient to droughts; (c) oak, silver fir, and to a lower extent beech, show higher resistance and resilience to spring droughts and seem therefore better adapted to the future climate. Our results allow a robust comparison of the tree growth responses to drought and spring frost across large climatic gradients and provide striking evidence that the growth of some of the most abundant and economically important European tree species will be increasingly limited by climate warming. These results could serve for supporting species selection to maintain the sustainability of forest ecosystem services under the expected increase in ECEs.     

The Origin and Dynamics of Subalpine White Spruce and Balsam Fir Stands in Boreal Eastern North America

Associations among the few tree species in the North American boreal landscape are the result of complex interactions between climate, biota, and historical disturbances during the Holocene. The closed-crown boreal forest of eastern North America is subdivided into two ecological regions having distinct tree species associations; the balsam fir zone and the black spruce zone, south and north of 49°N, respectively. Subalpine old-growth stands dominated by trees species typical of the balsam fir forest flora (either balsam fir or white spruce) are found on high plateaus, some of which are isolated within the black spruce zone. Here we identified the ecological processes responsible for the distinct forest associations in the subalpine belt across the eastern boreal landscape. Extensive radiocarbon dating, species composition, and size structure analyses indicated contrasted origin and dynamics of the subalpine forests between the two ecological regions. In the black spruce zone, the subalpine belt is a mosaic of post-fire white spruce or balsam fir stands coexisting at similar elevation on the high plateaus. With increasing time without wildfire, the subalpine forests become structurally similar to the balsam fir forest of the fir zone. These results concur with the hypothesis that the subalpine forests of this area are protected remnants of an historical northern expansion of the fir zone. Its replacement by the fire-prone black spruce forest flora was caused by recurrent fires. In the subalpine belt of the fir zone, no fire was recorded for several millennia. Harsh climate at high altitude is the primary factor explaining white spruce dominance over balsam fir forming a distinct subalpine white spruce belt above the balsam fir dominated forest.     

Drought induces lagged tree mortality in a subalpine forest in the Rocky Mountains

Extreme climatic events are key factors in initiating gradual or sudden changes in forest ecosystems through the promotion of severe, tree-killing disturbances such as fire, blowdown, and widespread insect outbreaks. In contrast to these climatically-incited disturbances, little is known about the more direct effect of drought on tree mortality, especially in high-elevation forests. Therefore projections of drought-induced mortality under future climatic conditions remain uncertain. For a subalpine forest landscape in the Rocky Mountains of northern Colorado (USA), we quantified lag effects of drought on mortality of Engelmann spruce Picea engelmannii , subalpine fir Abies lasiocarpa , and lodgepole pine Pinus contorta . For the period 1910–2004, we related death dates of 164 crossdated dead trees to early-season and late-season droughts. Following early-season droughts, spruce mortality increased over five years and fir mortality increased sharply over 11 years. Following late-season droughts, spruce showed a small increase in mortality within one year, whereas fir showed a consistent period of increased mortality over two years. Pine mortality was not affected by drought. Low pre-drought radial growth rates predisposed spruce and fir to drought-related mortality. Spruce and fir trees that died during a recent drought (2000–2004) had significantly lower pre-drought growth rates than live neighbour trees. Overall, we found large interspecific differences in drought-related mortality with fir showing the strongest effect followed by spruce and pine. This direct influence of climatic variability on differential tree mortality has the potential for driving large-scale changes in subalpine forests of the Rocky Mountains.     

Size‐dependence of tree growth response to drought for Norway spruce and European beech individuals in monospecific and mixed‐species stands

• Climate anomalies have resulted in changing forest productivity, increasing tree mortality in Central and Southern Europe. This has resulted in more severe and frequent ecological disturbances to forest stands. This study analysed the size‐dependence of growth response to drought years based on 384 tree individuals of Norway spruce [Picea abies (L.) Karst.] and European beech [Fagus sylvatica ([L.)] in Bavaria, Germany.
• Samples were collected in both monospecific and mixed‐species stands. To quantify the growth response to drought stress, indices for basal area increment, resistance, recovery and resilience were calculated from tree ring measurements of increment cores. Linear mixed models were developed to estimate the influence of drought periods.
• The results show that ageing‐related growth decline is significant in drought years. Drought resilience and resistance decrease significantly with growth size among Norway spruce individuals. Evidence is also provided for robustness in the resilience capacity of European beech during drought stress. Spruce benefits from species mixing with deciduous beech, with over‐yielding spruce in pure stands.
• The importance of the influence of size‐dependence within tree growth studies during disturbances is highlighted and should be considered in future studies of disturbances, including drought.

     

Silver fir stand productivity is enhanced when mixed with Norway spruce: evidence based on large‐scale inventory data and a generic modelling approach

Questions: How to evaluate the mixture effect on basal area increment in two‐species forest stands? Is a mixed Norway spruce–silver fir stand more productive than pure adjacent stands of either species? How to develop generic modelling approaches to assess mixture effects in forest stands? Location: In addition to a case study on Norway spruce–silver fir stands in French mountain forests, the generic approach used goes beyond local applications. Methods: We took advantage of National Forest Inventory data to develop a unique stand basal‐area‐increment model for pure and mixed stands of Norway spruce and silver fir that responds to ecological site conditions. The database was made up of 284 pure Norway spruce stands, 196 pure silver fir stands, and 323 mixed stands of these species. Results: Pure silver fir basal area increment is strongly influenced by spring climatic conditions, whereas pure Norway spruce is more influenced by soil conditions. The mixture of these species has a positive effect on silver fir, which decreases as the proportion of fir increases. In contrast, the mixture has no noticeable effect on Norway spruce. Conclusion: We developed a stand basal‐area‐increment model evidencing an advantage of the mixture on silver fir basal area increment, but not on Norway spruce. The mathematical formulation of the model developed is generic and can be used in all two‐species mixture situations. It also makes it possible to compare different mixture situations with each other.     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.     

Resilience capacity of Araucaria araucana to extreme drought events

Ongoing climate change has induced modification in the frequency and intensity of extreme climatic events, with consequent impact on tree and forest growth resilience. Araucaria araucana is an endangered Patagonian conifer, which provides several ecosystem services to local human societies and plays fundamental ecological roles in natural communities. These woodlands have historically suffered different types of anthropogenic disturbance, such as fire, logging and grazing, nevertheless the species resilience to extreme drought events remains still poorly understood. To fill this gap of knowledge, we applied dendrochronological methods to several A. araucana stands distributed along a steep bioclimatic gradient in order to reconstruct resilience capacity, in term of stem growth resistance and recovery, to three successive extreme spring-early summer droughts which occurred during the 20th century. Results showed an increase in the species recovery along the considered dry spells, whereas no clear trend emerged for resistance, suggesting no cumulative effect of drought upon resilience. Both resistance and recovery presented different values depending on bioclimatic settings, being xeric stands more sensitive to extreme episodes with respect to mesic woodlands, particularly during the more recent drought event when trees growing in drier environments were not able to reach pre-drought stem growth rates. Tree-level characteristics, such as age and growth trends prior to drought, modulated the species resilience, suggesting that future dry spells would possibly induce shifts in population dynamics, and furthermore be detrimental for fast-growing trees. Our analysis highlighted the response of a key Patagonian tree species to extreme drought events, providing bioclimatic-specific useful information for conservation plans of this natural resource.     

Tree ring isotopic composition, radial increment and height growth reveal provenance-specific reactions of Douglas-fir towards environmental parameters

In the search of timber species being tolerant towards summer droughts, which are expected to be more frequent in future, Douglas-fir is often discussed as a potential alternative for spruce in Central Europe. To assess physiological and growth reactions of Douglas-fir provenances towards climate- and weather-related environmental conditions we took advantage of a provenance trial with three sites in south-western Germany located along an elevation gradient. We examined six different provenances of Douglas-fir from North America for oxygen (δ¹⁸O) and carbon (δ¹³C) stable isotope composition in tree rings as well as for radial increment for a 7 year period and long-term height growth. Our results show that different Douglas-fir provenances clearly vary in their drought sensitivity at the driest and warmest site in the valley as shown by the radial growth decline in the extreme dry and hot year 2003. The growth decline in the provenances Pamelia Creek, Cameron Lake, Duncan Paldi and Conrad Creek could be clearly attributed to a reduction in stomatal conductance as assessed by the relations between δ¹⁸O and δ¹³C in the tree rings. These responses were not related to the long-term average climate at the places of origin of the provenances and the provenances with the lowest long-term (height) growth potential were the ones least affected in radial increment by the extreme drought of 2003. When selecting suitable Douglas-fir provenances, which are adapted to the climatic conditions projected for the future we thus might need to take into account the trade-off between the adaptation to extreme drought periods and the long-term growth performance. Site-specific evaluations of the probability of extreme drought events are thus needed to select the appropriate provenances.     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

The symmetry of competitive interactions in mixed Norway spruce,silver fir and European beech forests

Questions

We aim for a better understanding of the different modes of intra‐ and inter‐specific competition in two‐ and three‐species mixed‐forests. How can the effect of different modes of competitive interactions be detected and integrated into individual tree growth models? Are species interactions in spruce–fir–beech forests more associated with size‐symmetric or size‐asymmetric competition? Do competitive interactions between two of these species change from two‐ to three‐species mixtures?

Location

Temperate mixed‐species forests in Central Europe (Switzerland).

Methods

We used data from the Swiss National Forest Inventory to fit basal area increment models at the individual tree level, including the effect of ecological site conditions and indices of size‐symmetric and size‐asymmetric competition. Interaction terms between species‐specific competition indices were used to disentangle significant differences in species interactions from two‐ to three‐species mixtures.

Results

The growth of spruce and fir was positively affected by increasing proportions of the other species in spruce–fir mixtures, but negative effects were detected with increasing presence of beech. We found that competitive interactions for spruce and fir were more related to size‐symmetric competition, indicating that species interactions might be more associated with competition for below‐ground resources. Under constant amounts of stand basal area, the growth of beech clearly benefited from the increasing admixture of spruce and fir. For this species, patterns of size‐symmetric and size‐asymmetric competitive interactions were similar, indicating that beech is a strong self‐competitor for both above‐ground and below‐ground resources. Only for silver fir and beech, we found significant changes in species interactions from two‐ to three‐species mixtures, but these were not as prominent as the effects due to differences between intra‐ and inter‐specific competition.

Conclusions

Species interactions in spruce–fir–beech, or other mixed forests, can be characterized depending on the mode of competition, allowing interpretations of whether they occur mainly above or below ground level. Our outcomes illustrate that species‐specific competition indices can be integrated in individual tree growth functions to express the different modes of competition between species, and highlight the importance of considering the symmetry of competition alongside competitive interactions in models aimed at depicting growth in mixed‐species forests.

     

Control of Nitrification by Tree Species in a Common-Garden Experiment

We studied the effect of tree species on nitrification in five young plantations and an old native beech coppice forest at the Breuil experimental site in central France. The potential net nitrification (PNN) of soil was high in beech, Corsican pine, and Douglas fir plantations (high nitrifying stands denoted H) and low in spruce and Nordmann fir plantations as well as in native forest stands (low nitrifying stands denoted L). We hypothesized that tree species would stimulate or inhibit nitrification in transplanted soil cores within a few years after the cores were transplanted between stands. We first initiated a transplant experiment where soil cores were exchanged between all stands. The PNN remained high in soil cores from H transferred to H and low in soil cores from L transferred to L. The PNN increased considerably after 16 months in soil cores transferred from L to H, whereas the transfer of soil cores from H to L decreased the PNN only slightly after 28 months. In a second transplant experiment, forest floor material was exchanged between the Douglas fir (H) and the native forest (L) stand. Six months later, the forest floor from the native forest had increased the PNN of the Douglas fir soil considerably, whereas the forest floor from Douglas fir did not affect the PNN of the soil in the native forest stand. It was concluded that beech, Corsican pine, and Douglas fir rapidly stimulate soil nitrification by either activation of suppressed nitrifier communities and/or colonization by new nitrifier communities. Conversely, the slow and irregular reduction of nitrification in spruce, Nordmann fir, and native forest was probably due to the low and heterogeneously distributed flux of inhibiting substances per volume of soil. Our experiments suggest that the inhibition of nitrification is not tightly connected to forest floor leachates, but that the forest floor both reflects and maintains the major ongoing processes. In the long term, humus build up and the production of inhibiting substances may completely block the nitrification activity.     

Tree-ring growth of silver fir (Abies alba Mill.) in two stands under different silvicultural systems in central Italy

A preliminary analysis was made of growth trends in two silver fir forests in the Apennines in central Italy. The forests are closely located but were managed differently in the past. The aim was to verify the climatic variation over time, the influence of extreme events on radial growth and the role of silvicultural systems in defining growth response. The two forests have distinct structures due to differing past silvicultural management. The Camaldoli Forest, containing one-storeyed, evenaged, pure silver fir stands, was essentially managed in the past for wood production and featured high plantation density, low to moderate thinning, and clear cutting every 80–100 years. The La Verna Forest includes multi-layered, unevenaged stands that are mixed in with beech and other broadleaves. Increment cores were collected from aged dominant and healthy trees in the two forests. Climatic data were taken from the Camaldoli station and covers the period 1885–1994. All the analysed dendroecological parameters displayed similar patterns and indicated suitable site conditions for the silver fir. Silvicultural systems played an important role in defining growth pattern and rate. In the evenaged system there were higher growth rates and a greater number of pointer years, whereas there were lower growth rates and long, homogeneous development stages in the semi-natural conditions of La Verna. A rising growth trend recorded in the last three decades in both series could be explained by the consistent extension of the growing season due to a significant increase of mean spring and autumn temperatures. No correlation was found with precipitation while significant correlations were found between tree ring growth and February, April and August temperatures. Preliminary results indicated that the silvicultural system does not bias, but just enhances the ability of the species to record the effects of disturbing factors.     

Tree‐ring stable isotopes record the impact of a foliar fungal pathogen on CO2 assimilation and growth in Douglas‐fir

Swiss needle cast (SNC) is a fungal disease of Douglas‐fir (Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree‐rings of Douglas‐fir and a non‐susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ¹³C) of treated Douglas‐fir tree‐rings was greater than that of untreated Douglas‐fir tree‐rings during the fungicide treatment period. Both annual growth and tree‐ring Δ¹³C increased with treatment such that treated Douglas‐fir had values similar to co‐occurring western hemlock during the treatment period. There was no difference in the tree‐ring oxygen stable isotope ratio between treated and untreated Douglas‐fir. Tree‐ring Δ¹³C of diseased Douglas‐fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years.