Drought sensitivity of Norway spruce is higher than that of silver fir along an altitudinal gradient in southwestern Germany

Context

For Central Europe, climate projections foresee an increase in temperature combined with decreasing summer precipitation, resulting in drier conditions during the growing season. This might negatively affect forest growth, especially at sites that are already water-limited, i.e., at low elevation. At higher altitudes trees might profit from increasing temperatures.

Aims

We analyzed variations in radial growth of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) along an altitudinal gradient from 400 until 1,140 m a.s.l. in the Black Forest, to assess climate responses with increasing elevation.

Methods

Climate–growth relationships were analyzed retrospectively using tree-ring and climate data. In total, we sampled stem discs of 135 trees to build 27 species- and site-specific chronologies (n _fir?=?13, n _spruce?=?14).

Results

Our results indicate distinct differences in climate–growth relations between fir and spruce along the gradient. Growth of high-altitude fir was positively related to temperature from January till March. Growth of low-altitude fir and spruce at all elevations was positively related to precipitation and negatively to temperature during the growing season, particularly in July. A self-calibrating Palmer drought severity index underlined summer drought sensitivity of these trees.

Conclusion

Overall, we found that climatic control of tree growth changes over altitude for fir. For spruce, a remarkable synchrony in growth variation and climate response was shown, which indicates that this species is drought sensitive at all studied elevations. In a future warmer climate, the growth of low-altitude fir and spruce along the entire studied gradient may be negatively affected in the Black Forest, if an increased evaporative demand cannot be compensated by increased water supply.     

Does tree seedling growth and survival require weeding of Himalayan balsam (Impatiens glandulifera)?

The effect of Himalayan balsam (Impatiens glandulifera) on survival and growth of naturally regenerated silver birch (Betula pendula) and planted Norway spruce (Picea abies) and silver fir (Abies alba) seedlings was studied in a weeding experiment over 3 years. Three different treatments were applied: control, mowing, and hand weeding by pulling out the entire plant. There were no consistent treatment effects on height and diameter of the tree seedlings. The coverage of Rubus fruticosus had a negative impact on diameter increment of Norway spruce and silver fir. As opposed to growth, treatment effects on seedling survival could be found for planted Norway spruce and silver fir. However, it is very likely that these effects, namely higher seedling survival after mowing, have to be attributed to the control of bramble (Rubus fruticosus) rather than to that of Himalayan balsam. It is concluded that Himalayan balsam is not able to seriously affect the growth of already established seedlings.     

Tree rings indicate different drought resistance of a native (Abies alba Mill.) and a nonnative (Picea abies (L.) Karst.) species co-occurring at a dry site in Southern Italy

Climate changes induced by the anthropogenic alteration of the atmospheric radiative balance are expected to change the productivity and composition of forest ecosystems. In Europe, the Mediterranean is considered one of the most vulnerable regions according to climatic forecasts and simulations. However, although modifications in the inter-specific competition are envisaged, we still lack a clear understanding of the ability of the Mediterranean vegetation to adapt to climate changes. We investigated how two co-occurring tree species commonly used in afforestation programmes, the native Abies alba Mill. and the nonnative Picea abies L. Karst., adapt to climate change by assessing their growth performance and physiological responses in relation to past climate variability. Growth was addressed by analysing tree-ring width and carbon and oxygen stable isotopes. Statistical relationships between isotopic value and monthly climate data suggest that the two species underwent ecophysiological adaptation to Mediterranean climatic constraints. These adaptations are also expressed in the ring-width data. Based on the carbon isotope ratio reflecting the stomatal response to drought, we found that the precipitation in the first period of the growing season, i.e. early spring, is a major factor influencing the annual growth of A. alba, which although native, proved to be sensitive to drought. P. abies, on the other hand, showed a higher tolerance to summer drought stress. These findings should help define criteria for sustainability and effective forest conservation in the Mediterranean region.     

Growth responses of three coexisting conifer species to climate across wide geographic and climate ranges in Yukon and British Columbia

This work aimed to compare radial growth–climate relationships among three coexisting coniferous tree species across a wide geographic and climate range from southern British Columbia (BC) to central Yukon, Canada. Tree-ring data were collected from 20 mature stands of white spruce (Picea glauca), lodgepole pine (Pinus contorta var. latifolia), and subalpine fir (Abies lasiocarpa). Linear relationships between annual growth variation and monthly and seasonal climate were quantified with correlation and regression analyses, and variation in climate–growth responses over a climatic gradient were quantified by regressing growth responses against local mean climatic conditions. Temperatures had more consistent and stronger correlations with growth for all three species than precipitation, but growth–climate responses varied among species and among sites. In particular, pine and fir populations showed different responses between BC and Yukon, whereas spruce showed a more consistent response across the study domain. Results indicate that (1) the response and sensitivity of trees to seasonal climate variables vary among species and sites and (2) winter temperatures prior to growth may have significant impacts on pine and fir growth at some sites. The capacity to adapt to climate change will likely vary among the study species and across climatic gradients, which will have implications for the future management of mixed-species forests in Yukon and BC.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

Establishment of the Second Rotation in the Jutland Heath Plantations

The paper describes briefly the silvicultural methods used toregenerate stands of pure Norway spruce (Picea abies Karst.)or Norway spruce mixed with a small proportion of Silver fir(Abies alba Mill.) on the poor heath soils of Jutland. Regenerationafter clear felling, uniform shelterwood, and shelterwood stripfelling is described. The last method has advantages over theother two.     

Variation in the incidence and severity of drought crack in three conifer species in North East Scotland

The incidence of longitudinal drought cracks in coniferous trees may increase as a consequence of climate change. While larger diameter trees are more prone to splitting, it is unclear why only some of the larger trees within the same stand typically suffer damage. Matched pairs of trees of the same size with and without cracks were studied in two adjacent pole-stage mixed stands of Sitka spruce/grand fir (Picea sitchensis/Abies grandis) and Sitka spruce/noble fir (Picea sitchensis/Abies procera) located in North-East Scotland. A range of attributes were measured on 15 damaged and undamaged trees of each species. Length and position of cracks on stems were also recorded, and annual ring width and latewood percentage measured on cores. Noble fir had significantly longer cracks and these were located higher in the stem than the other species. Fewer cracks formed on east-facing side of stems in all species suggesting that prevailing westerly winds may place greater tensile forces on the damaged sides of stems. A higher proportion of latewood (associated with higher tangential shrinkage) was found in the firs and may explain the greater incidence of cracking on grand and noble fir stems (13% and 16%, respectively) in comparison with Sitka spruce (3%).     

Long-term dynamics of a mixed conifer stand in Slovenia managed with a farmer selection system

The single-tree selection system is an important option for management of Norway spruce (Picea abies (L.) Karst.) and silver fir (Abies alba Mill.) forests because it provides continuous cover, requires low investments for tending, and promotes natural regeneration as well as high stand resistance and elasticity. It is often regarded as a very conservative system that usually results in only minor spatiotemporal changes in forest structure and composition. We studied management history, structural changes, regeneration dynamics, and light climate of a traditional single-tree farmer selection silver fir-Norway spruce forest (site typology Bazzanio-Abietetum). Stand structure was analyzed on five 0.25 ha permanent plots in 1994, 2001, and 2008. Regeneration density and height growth, forest floor vegetation, and light climate were also assessed on 1.5 × 1.5 m regeneration subplots in 2001 and 2008. Tree cores extracted from dominant trees from both species in two plots were used for reconstructing stand history and age structure of the canopy layer. We documented the forest response to three types of selection management regimes: excessive, normal, and conservative. Excessive management with harvest intensity significantly above the increment was documented until the late 1950s, including two peaks of heavy fellings (diameter limit cut) in the 1880s and 1930s, which favoured establishment of Norway spruce and released regeneration. The period that followed was characterized by normal selection management, but was nevertheless marked by a decline of silver fir as a result of air pollution and several droughts. This led to sanitary fellings that were carried out from the late 1970s to the early 1990s. In the last two decades conservative management followed, which led to suppression and decline of regeneration, especially of Norway spruce, and loss of selection structure. Although we recorded lower regeneration potential of silver fir compared with Norway spruce within the seedling category, silver fir outcompeted Norway spruce within the small-sized tree category (1 cm < dbh ? 10 cm) because of its superior height growth in low light levels (diffuse light <6%) and occupied a greater share of the canopy. Nevertheless, we anticipate that over the long-term the low light regime will also cause regeneration decline of silver fir and broadleaves. Our research revealed significant structural changes in a single-tree farmer selection forest during the last 150 years. These were a result of variable management regime and environment. A farmer single-tree selection system could better mimic the natural disturbance regime if spatiotemporal combinations of diverse felling regimes would be used.     

Size-mediated climate-growth relationships in temperate forests: A multi-species analysis

In most dendrochronological studies, climate-growth relationships are established on dominant trees to minimize non-climatic signals. However, response to environmental factors may be affected by tree-size, which begs the question of the representativeness of dominant trees on the stand level. To highlight the variations in climate-growth relationships among sizes and species, under a wide range of ecological conditions (climate and soil properties), 61 pure even-aged stands were sampled across France. At each stand, two tree-ring chronologies were established from 10 big- to 10 small-diameter trees. Our objectives were, (1) to assess variations in climate sensitivity between the two size-diameter classes, and (2) to investigate the role of species and ecological conditions on these variations. The climate-growth relationships were evaluated from 122 tree-ring chronologies (1 220 trees) through extreme growth years and correlation function analyses. Sensitivity to climate of shade-intolerant and moderately shade-tolerant species (Picea abies (L.) Karst., Pinus sylvestris L. and Quercus petraea (Matt.) Liebl.) remained constant between the size-diameter classes for both temperature and hydric balance, while the shade-tolerant species Abies alba Mill. and Fagus sylvatica L. displayed significant differences, with larger trees being more sensitive to summer drought than smaller trees. This difference increased with increasing climatic xericity. Our results suggest that, for shade-tolerant species, (1) big trees could be more sensitive to climatic change especially under xeric climate, and (2) future tree ring studies should include trees stratified by size to produce unbiased estimation of sensitivity to climate.     

Dependence of winter water relations of mature high-elevation Picea engelmannii and Abies lasiocarpa on summer climate

Water relations of Engelmann spruce (Picea engelmannii Parry) and subalpine fir (Abies lasiocarpa (Hook.) Nutt.) trees growing at an elevation of 3230 m on Mt. Evans, Colorado, USA, were studied during the winters of 1995-1996 and 1996-1997. During both winters, current-year and 1-year-old shoots were collected weekly and their relative water contents (RWC) determined. Measured meteorological parameters were used in a conifer winter water relations model, WINWAT, to simulate changes in shoot RWC of P. engelmannii and A. lasiocarpa during the winter. The model failed to predict shoot RWCs in 1996-1997 when calibrated with 1995-1996 data. The cold early summer of 1995 inhibited xylem formation, which appears to have caused lower rates of water recharge to the needles during the 1995-1996 winter than during the 1996-1997 winter. We conclude that summer climate strongly affects winter water relations in these subalpine species, and that changes in both summer and winter climate must be considered when predicting future ranges of these species.     

An increase in the upper tree-limit of silver fir (Abiesalba Mill.) in the Alps since the mid-20th century: A land-use change phenomenon

Global environmental changes observed during recent decades are likely to have had an impact on the distribution of species. Currently, silver fir (Abies alba) is becoming established in the subalpine forests of the west central Alps at elevations higher than 2000 m a.s.l.; prior to the 1970s its upper altitudinal limit was 2000 m. Several hypotheses could explain this recent expansion of the upper tree-limit. Silver fir regeneration could be linked (1) to land-use changes or (2) to current climatic warming. Using dendrochronology, the age structure of 31 forest plots containing at least one silver fir was examined in order to elucidate the population dynamics of subalpine communities. This allowed us to decipher the timing of fir regeneration in relation to the mean age of the stands examined and of the other tree-canopy species present. The majority of the firs germinated sporadically since 1950, before the regional temperature increase. The pattern of fir recruitment did not appear to relate to altitude, but followed a pattern characteristic of secondary succession. The age structures identified showed an exponential increase in tree-density during the 20th century; the stands were first dominated by Larix decidua during the 18th and 19th centuries, and then by Pinus cembra during the 20th century. In most stands, fir regeneration occurred after Larix and before P. cembra dominated, following a similar pattern to Picea abies regeneration. The number of local inhabitants and temperature both exhibited a negative relationship with fir tree recruitment, thus supporting the land-use change hypothesis. There has been a significant upward shift of the altitudinal range of fir, amounting to an increase of about 300 m since 1950. This followed the abandonment of low-productivity land. This trend is likely to continue during the 21st century, because of new agricultural and forestry practices which involve limited intervention in low-productivity areas and may be because of the effects of global warming.     

Impact of bark wounds on sapwood in Norway spruce and silver fir

Bark wounds by damage during harvesting are a serious problem in forestry due to fungi infection and wood deterioration. This paper presents results of an investigation about the influence of the wounds on the internal structure of such injured stems. In an experiment, bark wounds were artificially created at the stem base of Norway spruce [Picea abies (L.) Karst.] and silver fir (Abies alba Mill.). 2 years later, the injured stems along with undamaged controls were cut and the trunk portion below breast height subjected to computer tomographic (CT) analysis. Analysis of the CT-images revealed a substantial impact of wounding on sapwood properties in spruce: directly adjacent to the wound surface in all examined trees, a large disturbance zone was detected affecting on average 17% of the potential sapwood area. With increasing distance from the wound, the size of this disturbance zone diminished, but was still detectable in all trees at breast height ca. 1 m above the bark wound.     

Response to climate changes in radial growth of Picea crassifolia in the Qilian mountains of northwestern China

In order to investigate the response to climate changes in radial growth of Picea crassifolia at the lower tree line in the middle Qilian mountains in northwestern China, relationships of standardized chronologies of annua ring, earlywood and latewood widths with mean monthly temperature and total monthly precipitation were analyzed by ways of correlation and pointer year analyses. The results show that annual ring, earlywood and latewood widths are significantly negatively correlated with mean monthly temperature in June and Juy. Annual ring and eadywooc widths are significantly and positively correlated with tOtal monthly precipitation in March, May and June and negatively correlated with total monthly precipitation in September. Latewood width is less sensitive to climate changes than the width of earlywood and insignificantly sensitive to precipitation. The results of pointer year analysis revealed that whet summer temperatures are higher than the mean summer temperature synchronization and the summer precipitatior lower than mean summer precipitation synchronization, narrow annual rings are formed. Wide annual rings are formec~ when summer temperatures are lower than the mean summer temperature synchronization and summer precipitation higher than mean summer precipitation synchronization. The results indicate that more precipitation in the spring and summer is helpful for radial growth while warmer summer restricts radial growth of P. crassifolia at the lower tree line in the middle Qilian mountains.     

Relationships of climate and cell features in stems and roots of black spruce and balsam fir

? The anatomical differences of mature black spruces and balsam firs were examined at stem and root level in order to characterize their wood properties at cellular level and link these differences to climate.

? Anatomical variability of these species was evaluated in relation to climate data gathered from 2001 to 2004 during the cell enlargement (CE) and wall thickening and lignification (WTL) phases. Lumen area, single cell wall thickness and total tracheid radial diameter were analyzed and regrouped into earlywood and latewood.

? Results from a principal component analysis (PCA) indicated that both first eigenvectors account for 82% and 90% of total variance for CE and WTL respectively. These component factors revealed that precipitation, humidity and number of days with precipitation significantly influence the lumen area (p = 0.0168) and radial cell diameter (p = 0.0222) in earlywood. Significant differences were registered between species and tree parts (stem and root) for the lumen area, radial cell diameter and cell wall thickness in both earlywood and latewood.

? In our study, black spruce exhibited smaller tracheid size in both stem and roots compared to balsam fir. Furthermore, the lower amount of tracheids produced during the growing season and higher proportion of latewood ensure a higher wood density of black spruce. The influence of temperature on earlywood formation is significant, whereas no influence was observed on latewood.

     

Growth patterns and sensitivity to climate predict silver fir decline in the Spanish Pyrenees

Patterns in radial growth have often been used to predict forest decline since they are regarded as indicators of the tree responses to long-term stressors. However, the sensitivity of tree growth to climate, as a proxy of the trees’ adaptive capacity to short-term climatic stress, has received less attention. Here, we used retrospective tree-ring analyses to determine whether growth patterns and sensitivity to climate are related to Abies alba (silver fir) decline in the Spanish Pyrenees. We used regional climatic data to calculate normalised temperatures and drought indexes. Basal-area increment (BAI) was measured for declining (defoliation >50%) and non-declining (defoliation <50%) silver firs in four stands with contrasting decline levels. A dynamic factor analysis (DFA) was applied to test the hypothesis that declining and non-declining trees have experienced different long-term growth trends. Growth sensitivity to climate was computed as the average change in BAI per unit of change in a given climate variable. Declining trees showed a negative growth trend during the last 20?years. Trees with lower relative BAI and negative BAI trends showed stronger growth sensitivity to climate and higher defoliation than trees with the opposite characteristics. Our findings underscore the idea that long-term climatic warming seems to be a major driving factor of growth decline in Pyrenean silver fir forests. Ongoing growth reduction and enhanced growth sensitivity to climate may promote vegetation shifts in these declining forests located near the xeric edge of the species distribution area.     

Estimating Scots pine (Pinus sylvestris L.) log diameter under bark by using shape and image data

The sawmilling industry stores and measures logs in bark in order to maximize efficiency, quality conservation and preservation. However, billing is based on the diameter under bark, which it is necessary to estimate based on manual or automatic bark detection. Recently, an approach for automatic determination of diameter under bark based on a multi-sensor approach, including shape data, colour image data and tracheid effect data has been presented, including promising results for logs of the species Norway spruce (Picea abies (L.) H. Karst) and Silver fir (Abies alba Mill.). This paper extends this approach to Scots pine (Pinus sylvestris L.). The comparison of the estimated diameters under bark of 270 pine logs with the respective diameters after debarking shows that the method works well and reliably. Estimation errors are in general close to zero and are below ±10?mm for 98% of the logs. In comparison with manual bark detection, the automatic approach is clearly an improvement. Influences of season or characteristics like discolouration are mostly small. Applying a bark detection algorithm trained on spruce to the pine logs leads to acceptable results, but using a separate algorithm for pine leads to an even better performance.     

The fracture behaviour of single wood fibres is governed by geometrical constraints: in situ ESEM studies on three fibre types

In situ tensile tests were performed in an environmental scanning electron microscope (ESEM) on earlywood, transition wood and latewood cells of Norway spruce (Picea abies [L.] Karst.). In order to examine the single wood fibres in a wet state, a specially designed tensile testing stage with a cooling device was built. The fracture behaviour of the cell types was studied at high resolution while straining. Different failure mechanisms were observed for the three tissue types. The thin-walled earlywood fibres showed tension buckling which gave rise to crack initiation and resulted in low tensile strength, whereas thick-walled latewood fibres predominately failed by transverse crack propagation without fibre folding.     

Terpene response of Picea abies and Abies alba to infection with Heterobasidion s.l.

The monoterpene composition of Picea abies and Abies alba resin was analysed in relation to growth by Heterobasidion spp. Fifteen‐year‐old P. abies and A. alba trees were inoculated on branches with three species of Heterobasidion annosum s.l. After 4 months of incubation, each host was colonized to a significantly greater degree by the pathogen specific to that host (H. parviporum on P. abies, H. abietinum on A. alba) than by the other fungi. Analysis of the enantiomeric monoterpene profiles in the spruce and fir showed that the response in terms of the relative proportions of the monoterpene compounds in the resin differed between tree species. Following challenge with Heterobasidion spp., A. alba trees did not show changes in monoterpene composition in addition to those in the wounding response (increase in (−)‐α‐pinene and (−)‐camphene, and decrease in β‐phellandrene). In P.abies, (−)‐α‐pinene, (+)‐α‐pinene and δ‐3‐carene increased following Heterobasidion attack but not after wounding alone.     

Climatic significance of tree-ring width and intra-annual density fluctuations in Pinus pinea from a dry Mediterranean area in Portugal

In Mediterranean climates trees may go through two periods of dormancy, resulting in special anatomical features such as false rings and other intra-annual density fluctuations (IADFs). In this paper, ring growth and the presence of IADFs were studied in Pinus pinea L. growing in the coastal and inland regions of Alentejo (southern Portugal). In order to identify the triggering factors associated with the IADFs, a new classification was proposed for the IADFs in P. pinea: Type E (latewoodlike cells within earlywood); Type E+ (transition cells between earlywood and latewood); Type L (earlywoodlike cells within latewood) and Type L+ (earlywoodlike cells between latewood and earlywood of the next tree ring). Response function analyses showed that radial growth of P. pinea was strongly correlated with precipitation in southern Portugal. The climatic response of P. pinea was higher in the inland area where the summer drought is more severe, the winter temperatures are lower and the soils have low water-holding capacity, in comparison with the coastal area. IADFs were frequent in P. pinea and most of the IADFs were observed in latewood. The presence of IADFs was correlated with fluctuations in climate parameters during the growing season. The IADF type E+ was linked to precipitation events early in summer. The IADF type L and L+ were associated with above-average precipitation in early autumn.