A Pine Is a Pine and a Spruce Is a Spruce – The Effect of Tree Species and Stand Age on Epiphytic Lichen Communities

With an increasing demand for forest-based products, there is a growing interest in introducing fast-growing non-native tree species in forest management. Such introductions often have unknown consequences for native forest biodiversity. In this study, we examine epiphytic lichen species richness and species composition on the trunks of non-native Pinus contorta and compare these to the native Pinus sylvestris and Picea abies in managed boreal forests in northern Sweden across a chronosequence of age classes. Overall, we recorded a total of 66,209 lichen occurrences belonging to 57 species in the 96 studied forest stands. We found no difference in species richness of lichens between stands of P. contorta and P. sylvestris, but stands of P. abies had higher total species richness. However, species richness of lichens in stands of P. abies decreased with increasing stand age, while no such age effect was detected for P. contorta and P. sylvestris. Lichen species composition progressively diverged with increasing stand age, and in 30-year-old stands all three tree species showed species-specific assemblages. Epiphytic lichen assemblages in stands of 30-year-old P. contorta were influenced by greater basal area, canopy closure, and average diameter at breast height, P. abies stands by higher branch density and canopy closure, and stands of P. sylvestris by greater bark crevice depth. Differences in lichen species richness and composition were mainly explained by canopy closure and habitat availability, and the greater canopy closure in mature P. abies stands promoted the colonization and growth of calicioid lichen species. Our results indicate that the non-native P. contorta have similar species richness as the native P. sylvestris. The main difference in lichen species richness and composition is between P. abies and Pinus spp. in managed forests of boreal Sweden.     

Potential contribution of the seed bank in coniferous plantations to the restoration of native deciduous forest vegetation

In this study, we compared the soil seed bank and current vegetation under coniferous plantations and adjacent native deciduous forests. The objective was to assess how much of the initial plant diversity is retained in such plantations, and the potential to restore this initial plant community from seed bank in case of reversion to broadleave stands. Four stands growing side by side and with different dominant species were selected at two locations (site of Haye: Quercus petraea, Pseudotsuga menziesii, Pinus sylvestris and Picea abies; site of La Petite-Pierre: Quercus petraea, Fagus sylvatica, Pinus sylvestris and Picea abies). In each stand, ground vegetation was surveyed and soil seed bank was sampled. Composition of ground flora and seed bank of stands were quite different: only 11 to 30 % of the species were in both the ground flora and the seed bank. Composition of the seed bank was mainly influenced by site location and sylvicultural practices such as the type of afforestation or the tree cover. Species richness of seed banks and vegetation were higher in the site of Haye than in the site of La Petite-Pierre. Seedling density strongly decreased with stand age. Whereas between 65 and 86 % of species found in the ground vegetation of native deciduous stand were also present in the understory or the seed bank of mature coniferous stands, this was only about 50 % in young coniferous stands. Species of deciduous stands which were absent from coniferous stands were typical of old forests. In contrast, species mainly found in the coniferous stands were often ruderal. In the studied areas, it would be possible to restore up to 86 % of the native deciduous forest vegetation, but some plant species typical of ancient forests may have disappeared during the coniferous stage.     

Neottia cordata (Orchidaceae) at its southernmost distribution border in Europe: Threat status and effectiveness of Natura 2000 Network for its conservation

Neottia cordata is an orchid species that exclusively occurs in Picea abies subsp. abies and Pinus sylvestris stands and forms its southernmost distribution limits in Greece. Over the last decade, it has been recorded in numerous sites, in which it forms small colonies, both in terms of population and area. This research re-evaluates the threat status of Neottia cordata based on updated chorological and population data; it also checks the effectiveness of the Natura 2000 Network as far as its conservation is concerned by applying the MaxEnt model. The recently collected data justify the classification of N. cordata as "Vulnerable". The most significant variables that affect its distribution were found to be the vegetation type and the precipitation of the warmest quarter. The results of the MaxEnt model suggest that: (i) the Natura 2000 Network is characterised by areas of higher habitat suitability values when compared to the areas that fall outside this network; and, (ii) Picea abies subsp. abies forests are more suitable for its conservation than those of Pinus sylvestris. It is concluded that the most appropriate management measure for the species conservation is the maintenance of the tree layer canopy closed.     

The structure and dynamics of old-growth Pinus sylvestris (L.) stands in the Wigry National Park,north-eastern Poland

The paper describes the structure and the developmental trends of old-growth Pinus sylvestris stands in the Wigry National Park, in north-eastern Poland. The stands represent a transitional zone between deciduous forests of Central Europe and boreal, coniferous forests of north-eastern Europe. Besides P. sylvestris, the most important tree species are Picea abies and Quercus robur. Among the subcanopy species, Corylus avellana and Sorbus aucuparia occur most frequently. On the basis of the data from 6 permanent sample plots (total size: 1.90 ha), several parameters and stand indices are analysed including species composition of the canopy and the regeneration, diameter distribution, age structure of main tree species, and the relationship between canopy and spatial dispersion of woody regeneration. The most striking feature of the stands studied is the almost complete absence of natural regeneration of P. sylvestris. This seems to be in contradiction with the apparently natural origin of this species in the stands, and a common occurrence of natural disturbances resulting in openings and gaps in forest canopy. The main tree species replacing P. sylvestris in the canopy are P. abies and Q. robur. Also increasing are some broad-leaved species typical of high fertility sites: Acer platanoides, Tilia cordata, Ulmus glabra, and Fraxinus excelsior. A shrub C. avellana occurs extensively competing with tree species and delaying tree replacement processes. While no direct data on the changes in the site conditions can be provided and the recovery hypothesis appears to be the most straightforward explanation of the changes in P. sylvestris stands, the possible role of the allogenic changes in environmental conditions (climate warming, nitrogen deposition) is also discussed.     

Ecological and methodological drivers of non-stationarity in tree growth response to climate

Radial tree growth is sensitive to environmental conditions, making observed growth increments an important indicator of climate change effects on forest growth. However, unprecedented climate variability could lead to non-stationarity, that is, a decoupling of tree growth responses from climate over time, potentially inducing biases in climate reconstructions and forest growth projections. Little is known about whether and to what extent environmental conditions, species, and model type and resolution affect the occurrence and magnitude of non-stationarity. To systematically assess potential drivers of non-stationarity, we compiled tree-ring width chronologies of two conifer species, Picea abies and Pinus sylvestris, distributed across cold, dry, and mixed climates. We analyzed 147 sites across the Europe including the distribution margins of these species as well as moderate sites. We calibrated four numerical models (linear vs. non-linear, daily vs. monthly resolution) to simulate growth chronologies based on temperature and soil moisture data. Climate–growth models were tested in independent verification periods to quantify their non-stationarity, which was assessed based on bootstrapped transfer function stability tests. The degree of non-stationarity varied between species, site climatic conditions, and models. Chronologies of P. sylvestris showed stronger non-stationarity compared with Picea abies stands with a high degree of stationarity. Sites with mixed climatic signals were most affected by non-stationarity compared with sites sampled at cold and dry species distribution margins. Moreover, linear models with daily resolution exhibited greater non-stationarity compared with monthly-resolved non-linear models. We conclude that non-stationarity in climate–growth responses is a multifactorial phenomenon driven by the interaction of site climatic conditions, tree species, and methodological features of the modeling approach. Given the existence of multiple drivers and the frequent occurrence of non-stationarity, we recommend that temporal non-stationarity rather than stationarity should be considered as the baseline model of climate–growth response for temperate forests.     

Effects of forest continuity and tree age on epiphytic lichen biota in coniferous forests in Estonia

Epiphytic lichen biota on Picea abies and Pinus sylvestris in Estonia was studied. Twenty-one spruce and 21 pine sample plots were located in old forests with long forest continuity, and 12 spruce and 12 pine sample plots in young first-generation forests (<100 years). Altogether 103 lichen species were recorded on the 330 sampled trees. Lichen species richness per plot was significantly higher in old forests in case of both tree species; 31 lichen species, including red-listed and protected species, were found only in old forests. Tree age had a positive effect on lichen species richness on tree stem in old and young spruce forests and in young pine forests. Tree age also had an effect on the presence of several species. Both tree age and forest continuity affected lichen species composition. Arthonia leucopellaea, Chrysothrix spp. and Lecanactis abietina were found in at least every third old spruce or pine forest and in no young forests, and can be regarded as good indicators of old coniferous forests with long continuity in Estonia.     

Nitrogen-addition effects on leaf traits and photosynthetic carbon gain of boreal forest understory shrubs

Boreal coniferous forests are characterized by fairly open canopies where understory vegetation is an important component of ecosystem C and N cycling. We used an ecophysiological approach to study the effects of N additions on uptake and partitioning of C and N in two dominant understory shrubs: deciduous Vaccinium myrtillus in a Picea abies stand and evergreen Vaccinium vitis-idaea in a Pinus sylvestris stand in northern Sweden. N was added to these stands for 16 and 8 years, respectively, at rates of 0, 12.5, and 50 kg N ha^?1 year^?1. N addition at the highest rate increased foliar N and chlorophyll concentrations in both understory species. Canopy cover of P. abies also increased, decreasing light availability and leaf mass per area of V. myrtillus. Among leaves of either shrub, foliar N content did not explain variation in light-saturated CO₂ exchange rates. Instead photosynthetic capacity varied with stomatal conductance possibly reflecting plant hydraulic properties and within-site variation in water availability. Moreover, likely due to increased shading under P. abies and due to water limitations in the sandy soil under P. sylvestris, individuals of the two shrubs did not increase their biomass or shift their allocation between above- and belowground parts in response to N additions. Altogether, our results indicate that the understory shrubs in these systems show little response to N additions in terms of photosynthetic physiology or growth and that changes in their performance are mostly associated with responses of the tree canopy.     

Tree population dynamics,stand structure and species composition in the montane virgin forest of Vallibäcken,northern Sweden

The population dynamics of Betula pubescens and Picea abies in a boreal forest near Kvikkjokk, northern Sweden, are governed by a process of storm gap regeneration similar to the gap regeneration described for boreo-nemoral forests. Cumulative age distribution curves, interpreted as static survivorship curves, lead to a simple theory of differential survival based on properties of the species, i.e. shade tolerance and relative growth rate. The theory is sustained by diameter and height distributions and by the spatial distributions of logs and of trees in different life-phases. Species of the field and ground layers respond differentially to gap formation and the ensuing successional stages. Browsing by moose (Alces alces) may prevent tree species, mainly Sorbus aucuparia, Betula pubescens and Pinus sylvestris, from developing into a tree layer. The regeneration ability for tree species growing in a stand at 460 m a.s.l. is limited compared with the regeneration at 330 m a.s.l., and typical storm gap formation involving more than one tree seems to occur rarely if at all, while overthrown trees with exposed rootplates are uncommon. Spruce at 460 m a.s.l. shows also a lower growth rate and a lower height/diameter ratio compared to the lower situated stands.     

Belowground interspecific competition in mixed boreal forests: fine root and ectomycorrhiza characteristics along stand developmental stage and soil fertility gradients

We studied fine roots and ectomycorrhizas in relation to aboveground tree and stand characteristics in five mixed Betula pendula Roth, Picea abies (L.) H. Karst., and Pinus sylvestris L. stands in Southern Finland. The stands formed gradients of developmental stage (15-, 30-, and 50-year-old stands) in the stands of medium fertility, and of site fertility in the young stands (30-year-old fertile, medium fertile, and least fertile stands). The biomass of the external hyphae of ectomycorrhizas (ECM) was the highest, and the diversity of the fungal community the lowest, in the most fertile stand. The vertical distributions of fine roots of the three tree species were mostly overlapping, indicating high inter-specific belowground competition in the stands. We did not find any clear trends in the fine root biomass (FRB) or length across the stand developmental stages. The FRB of the conifers varied with site fertility, whereas in B. pendula it was almost constant. In contrast to the conifers, the specific root length (SRL) of B. pendula clearly increased from the most fertile to the least fertile stand. This indicates differences in the primary nutrient acquisition strategy between conifers and B. pendula.     

Comparing growth phenology of co-occurring deciduous and evergreen conifers exposed to drought

Plant phenological events are influenced by climate factors such as temperature and rainfall. To evaluate phenological responses to water availability in a Spring Heath-Pine wood (Erico-Pinetum typicum), the focus of this study was to determine intra-annual dynamics of apical and lateral growth of co-occurring early successional Larix decidua and Pinus sylvestris and late successional Picea abies exposed to drought. The effect of reduced plant water availability on growth phenology was investigated by conducting a rainfall exclusion experiment. Timing of key phenological dates (onset, maximum rate, end, duration) of growth processes were compared among species at the rain-sheltered and control plot during 2011 and 2012. Shoot and needle elongation were monitored on lateral branches in the canopy at c. 16 m height and radial growth was recorded by automatic dendrometers at c. 1.3 m height of >120 yr old trees. Different sequences in aboveground growth phenology were detected among the three species under the same growing conditions. While onset of radial growth in April through early May was considerably preceded by onset of needle growth in Larix decidua (5–6 weeks) and shoot growth in Pinus sylvestris (c. 3 weeks), it occurred quite simultaneously with onset of shoot growth in Picea abies. Low water availability had a minor impact on onset of aboveground growth, which is related to utilization of stored water, but caused premature cessation of aboveground growth. At the control plot mean growing season length was 130 days in Pinus sylvestris, 95 days in Larix decidua and 73 days in Picea abies, supporting the hypothesis that early successional species are resource expenders, while late successional species are more efficient in utilizing resources and develop safer life strategies. High synchronicity found in culmination of radial growth in late spring (mid-May through early June) prior to occurrence of more favourable environmental conditions in summer might indicate sink competition for carbohydrates to belowground organs. This is supported by completion of apical growth in mid June in all species, except for needle growth of Pinus sylvestris which lasted until early August. Phenological observations of conifers exposed to drought revealed that tree water status early during the growing season determines total annual aboveground growth and, besides temperature, species-specific endogenous and/or environmental factors (most likely photoperiod and/or different threshold temperatures) are involved in controlling apical and lateral growth resumption after winter dormancy.     

DOES TREE AGE PREDICT THE OCCURRENCE AND ABUNDANCE OF USNEA LONGISSIMA IN MULTI-AGED SUBMONTANE PICEA ABIES STANDS?

In boreal Scandinavia Usnea longissima is a rare epiphytic lichen associated with old spruce (Picea abies) forests. To test whether tree age was an important determinant of the occurrence and abundance of U. longissima at the scale of a forest stand we estimated thallus numbers on 1008 trees within three submontane spruce stands in south-central Norway. The stands were of all ages and multi-layered and had been selectively cut previous to the 1940s. There was no relationship between number of U. longissima thalli and tree age among trees with dbh of ≥10 cm. The most consistent pattern was a positive relationship between occurrence and number of thalli and stem diameter. After controlling the covariance of stem diameter, there was a tendency for a negative relationship between number of thalli and tree age. The results suggest that in old-growth resembling stands, with already established U. longissima populations, tree size (here measured as stem diameter) is a better predictor of thallus number than tree age. This does not contradict the possibility that other environmental characteristics, associated with stands of older age, may be important for the establishment and growth of U. longissima populations.     

Contrasting acclimation abilities of two dominant boreal conifers to elevated CO2 and temperature

High latitude forests will experience large changes in temperature and CO₂ concentrations this century. We evaluated the effects of future climate conditions on 2 dominant boreal tree species, Pinus sylvestris L. and Picea abies (L.) H. Karst, exposing seedlings to 3 seasons of ambient (430 ppm) or elevated CO₂ (750 ppm) and ambient temperatures, a + 4 °C warming or a + 8 °C warming. Pinus sylvestris responded positively to warming: seedlings developed a larger canopy, maintained high net CO₂ assimilation rates (A_net), and acclimated dark respiration (R_dark). In contrast, carbon fluxes in Picea abies were negatively impacted by warming: maximum rates of A_net decreased, electron transport was redirected to alternative electron acceptors, and thermal acclimation of R_dark was weak. Elevated CO₂ tended to exacerbate these effects in warm‐grown Picea abies, and by the end of the experiment Picea abies from the +8 °C, high CO₂ treatment produced fewer buds than they had 3 years earlier. Treatments had little effect on leaf and wood anatomy. Our results highlight that species within the same plant functional type may show opposite responses to warming and imply that Picea abies may be particularly vulnerable to warming due to low plasticity in photosynthetic and respiratory metabolism.     

Exploring the growth response of Norway spruce (Picea abies) along a small-scale gradient of soil water supply

The climate-growth response of specific sites and species is one of the main research subjects in classic tree ring studies. Traditional sampling approaches therefore aim at maximizing the climate signal of the analyzed tree ring series, which is typically achieved by focusing on dominant trees or on sites located in particularly temperature or moisture limited environments. However, there is increasing evidence that these selective sampling strategies cannot yield chronologies that are representative for entire populations. One promising approach to gain a deeper understanding of forest dynamics and climate-growth responsiveness is the analysis of climate signal ranges among trees. This individualistic approach requires random sampling and the integration of information on small-scale heterogeneities in site and tree characteristics. Here, we analyze the climate-growth response of 144 Norway spruce trees (Picea abies Karst.) on difference levels of data aggregation. The aim of our study is to investigate the relevance of small-scale heterogeneities in site conditions, particularly in soil water supply, for the detected climate-growth signal. We identify soil water supply and site characteristics, which indirectly modify the water availability for trees, as dominating growth factors across scales. The driest sites show the strongest climate-growth reaction, while the growth response of wetter sites is weak or even insignificant. Therefore, we conclude that integrating small-scale information on site characteristics, particularly on soil water supply, can help to gain a deeper understanding of species specific growth limitations.     

Challenges for growth of beech and co-occurring conifers in a changing climate context

Given recent climatic trends, it is crucial to understand the plasticity and adaptiveness of forest trees in order to evaluate their current and future responses to changing climatic conditions. We investigated inter- and intra-annual xylem growth and its relation to weather factors in European beech (Fagus sylvatica) and a co-occurring conifer, either Norway spruce (Picea abies) or Scots pine (Pinus sylvestris), at five sites with different elevations and climatic conditions, in Spain, Slovenia and the Czech Republic. The selected sites were located in central and marginal parts across the distribution range of the investigated species. The results showed that climate-growth relationships vary between different species at the same site and within the same species at different sites, indicating diverse survival strategies among the observed species but also high plastic capacity within the same species. Moreover, xylogenesis analysis revealed the capacity of trees to adapt the beginning and cessation of wood formation depending on conditions. In case of beech, shortening growth duration in dryer and warmer environments and the opposite in the case of conifers, with which the growing period was extended in such conditions. Consequently, the plasticity capacity of beech was limited due to short growth duration, while conifers (especially pine) proved to be able to compensate climatic constrictions by growing over a longer period and becoming more adapted to survival in drought-prone environments.     

Tree roots as self-similar branching structures: axis differentiation and segment tapering in coarse roots of three boreal forest tree species

We applied a fractal root model to the 3D architecture of the coarse root systems of Betula pendula Roth, Picea abies (L.) H. Karst., and Pinus sylvestris L. in mixed boreal forests. Our dataset consisted of 60 root systems excavated in five different mixed forest stands. We analyzed the variability of the model parameters with respect to species, site type, and different root axes. According to our results, the cross-sectional area of root segments (i.e. second power of diameter) was a suitable variable for analyzing the values of parameters of the fractal model. The parameter values varied with generation and order of root segments; the roots thus did not follow the simple fractal branching. The variation of parameters along the root axes showed the existence of a zone of rapid tapering in all tree species. The model was, with parameter values analyzed from the data, moderately capable of accounting for the main coarse root characteristics. It was important for model predictions to take into account the tapering of root segments. We conclude that, in boreal forests, tree root systems are the output of the axis-specific morphogenetic branching rules and functional adaptation to spatial heterogeneity in the soil.     

Among‐tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide‐ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short‐ and long‐term tree growth responses, focusing on among‐tree variability and potential feedback effects. Although among‐tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth–climate relationships. We compiled tree‐ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among‐tree variability, we employed information‐theoretic model selections based on linear mixed‐effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long‐term trends in ring‐width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among‐tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L. decidua) and current year's spring (L. decidua, P. abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L. decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P. abies, P. cembra, and P. mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long‐suspected negative and positive feedback of climate change on stand dynamics at treeline.     

The Finnish forest site type approach: ordination and classification studies of mesic forest sites in southern Finland

A.K. Cajander's forest site type classification system is based on definition of plant communities typical to certain climatical and edaphical site conditions, but the structure and composition of the tree stands in Finland are considered sensitive to random variation and are therefore not used as primary classification criteria. The system has often received criticism, usually that the effects of the tree stand and successional stage of the stand have been underestimated. Most of the present-day forest stands in Finland represent young successional stages and are subjected to intensive management. This should result in an additional difficulty in the application of the forest site types in the field.The present study is based on three independent data sets representing forests on mineral soil in southern part of Finland. TWINSPAN classification, DCA ordination and canonical correspondence analysis (CCA) techniques were applied in successive stages of the data analysis. It was found that the definition of the intermediately fertile, mesic site types was clearly confused by the effects of the tree species and age of the stand. The analyses also revealed that the succession pathways on mesic forest sites are largely determined by the tree species composition. In stands dominated by Pinus sylvestris, the succession follows the competitive hierarchy model, whereas in stands dominated by Picea abies, severe shading of the tree canopy governs the development of understorey vegetation.Abbreviations CCA Canonical correspondence Analysis - DCA Detrended correspondence Analysis - TWINSPAN Two-way indicator species analysis     

Aleppo pine vulnerability to climate stress is independent of site productivity of forest stands in southeastern Spain

Key message

Clim ate is the main driver of Aleppo pine radial growth variability irrespective of site pro ductivity, with the climate effects on tree growth more limiting from 1970 onwards.

Abstract

Forest management adaptation to climate change requires identifying the previously most vulnerable stands and the possible climate impacts on forests. This study evaluates whether site index, as an indicator of forest productivity, is related to climate–growth responses and assesses the way in which local site factors modulate climate–tree growth relationships. Tree-ring width series and soil characteristics were obtained from six Pinus halepensis stands with different site indices and similar climate. Dendrochronological methods were used to compare tree climate–growth responses among sites and to study temporal trends in inter-annual growth variability and climate–growth relationships (before and after 1970). The influence of topographic and soil features on tree growth was assessed by means of partial least squares. Stands with low site indices tended to present higher mean sensitivities and greater percentages of missing rings, this relation being modulated mainly by clay percentage and nutrient status in soil. Climate is the major Aleppo pine radial growth driver in the study area with similar growth–climate relationship among sites. Radial growth was mainly influenced by spring temperature and precipitation and previous autumn–winter precipitation. This relationship was stronger after 1970 than before this year, showing also a 2-month advancement of the most influential climate variables after 1970, from May to March. These results and the increasing temporal trend found in mean sensitivity after 1970 highlight the vulnerability of these stands to climate change. Site index was not found to be directly related to stand vulnerability, although local site factors modulate in part the tree-growth response.     

Topographic influences on radial growth of Scots pine (Pinus sylvestris L.) at small spatial scales

Dendroecological and numerical methods were used to study the influence of topographic position on radial growth of Scots pine (Pinus sylvestris) stands exposed to soil dryness. The correlation structure of total tree-ring width and latewood width of eight scattered populations representing various topographic habitats (steep south-facing slopes, plateaus and hollows) within a rock-slide area (750 m a.s.l.) of about 1 km² was investigated by principal component analysis. Scatter plots of component loadings indicated that (i) total ring width and latewood width are influenced by various climatic factors, (ii) stands growing at similar topographic position show a high agreement in year-to-year variability of radial growth, and (iii) distinct effects of topographic features (slope aspect, slope magnitude) on tree growth are modified by local disturbances (erosion, grazing) and the age structure of stands. Furthermore, both the time series of component scores and non-metric multidimensional scaling of chronologies indicated years where extremely limiting or favorable climate conditions prevailed throughout the study area (pointer years). The influence of climate on tree growth in various topographic habitats was mediated through the influence of climatically stressful years. Because stands are located at sites with different levels of water stress, growth differences between chronologies are considered to be caused by site-specific susceptibility of tree growth to soil dryness. Significant correlations between precipitation in April to June and ring-width confirm that water availability is the primary growth-limiting factor within the study area. These small-scale variations in growth-climate relationships have significant implications for dendroclimatological studies. So paleoclimatic reconstructions based on tree rings will have to assure that an unbiased data set is used, which compensates for local growth-variabilities due to site related environmental stresses.