A density management diagram for Norway spruce in the temperate European montane region

Norway spruce is one of the most important conifer tree species in Europe, paramount for timber provision, habitat, recreation, and protection of mountain roads and settlements from natural hazards. Although natural Norway spruce forests exhibit diverse structures, even-aged stands can arise after disturbance or as the result of common silvicultural practice, including off-site afforestation. Many even-aged Norway spruce forests face issues such as senescence, insufficient regeneration, mechanical stability, sensitivity to biotic disturbances, and restoration. We propose the use of Density Management Diagrams (DMD), stand-scale graphical models designed to project growth and yield of even-aged forests, as a heuristic tool for assessing the structure and development of even-aged Norway spruce stands. DMDs are predicated on basic tree allometry and the assumption that self-thinning occurs predictably in forest stands. We designed a DMD for Norway spruce in temperate Europe based on wide-ranging forest inventory data. Quantitative relationships between tree- and stand-level variables that describe resistance to selected natural disturbances were superimposed on the DMD. These susceptibility zones were used to demonstrate assessment and possible management actions related to, for example, windfirmness and effectiveness of the protective function against rockfall or avalanches. The Norway spruce DMD provides forest managers and silviculturists a simple, easy-to-use, tool for evaluating stand dynamics and scheduling needed density management actions.     

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.     

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

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

Individual-tree diameter growth models for black spruce and jack pine plantations in northern Ontario

Individual-tree distance independent diameter growth models were developed for black spruce and jack pine plantations. Data used in this study came via stem analysis on 1170 black spruce (Picea mariana [Mill.] B.S.P.) and 800 jack pine (Pinus banksiana Lamb.) trees sampled from 75 stands of 25 even-aged monospecific plantations for each species in the Canadian boreal forest region of northern Ontario. Of the 75 stands, 50 were randomly selected for each species and all trees from these stands were used for model development. Trees from the remaining stands were used for model evaluation.A nonlinear mixed-effects approach was applied in fitting the diameter growth models. The predictive accuracy of the models was improved by including random effects coefficients. Four selection criteria - random, dominant or codominant, tree size close to quadratic mean diameter, and small sized - were evaluated for accuracy in predicting random effects for a new stand using the developed models. Random effects predicted based on trees selected using the random selection criterion provided more accurate diameter predictions than those using trees obtained via other selection criteria for both species. The models developed here are very important to forest managers as the diameters predicted by these models or, their stand-level summaries (i.e., basal area, average diameter), are used as inputs in any forest growth and yield models. In addition, individual-tree diameter growth models can be used to directly forecast changes in diameter distribution of stands.     

Assessing the impacts of climate change and nitrogen deposition on Norway spruce (Picea abies L. Karst) growth in Austria with BIOME-BGC

The aim of this paper is to determine whether a detectable impact of climate change is apparent in Austrian forests. In regions of complex terrain such as most of Austria, climatic trends over the past 50 years show marked geographic variability. As climate is one of the key drivers of forest growth, a comparison of growth characteristics between regions with different trends in temperature and precipitation can give insights into the impact of climatic change on forests. This study uses data from several hundred climate recording stations, interpolated to measurement sites of the Austrian National Forest Inventory (NFI). Austria as a whole shows a warming trend over the past 50 years and little overall change in precipitation. The warming trends, however, vary considerably across certain regions and regional precipitation trends vary widely in both directions, which cancel out on the national scale These differences allow the delineation of 'climatic change zones' with internally consistent climatic trends that differ from other zones. This study applies the species-specific adaptation of the biogeochemical model BIOME-BGC to Norway spruce (Picea abies (L.) Karst) across a range of Austrian climatic change zones, using input data from a number of national databases. The relative influence of extant climate change on forest growth is quantified, and compared with the far greater impact of non-climatic factors. At the national scale, climate change is found to have negligible effect on Norway spruce productivity, due in part to opposing effects at the regional level. The magnitudes of the modeled non-climatic influences on aboveground woody biomass increment increases are consistent with previously reported values of 20-40 kg of added stem carbon sequestration per kilogram of additional nitrogen deposition, while climate responses are of a magnitude difficult to detect in NFI data.     

Evaluation of forest growth simulators with NFI permanent sample plot data from Finland

Tree-level and stand-level forest growth simulators and their combination were evaluated using data from a large network of permanent sample plots of the National Forest Inventory covering the whole of Southern Finland. The simulators were built up with the SIMO framework. The evaluation was carried out both at the stand-level and separately for Scots pine (Pinus sylvestris), Norway spruce (Picea abies), silver birch (Betula pendula) and white birch (Betula pubescens) strata within the plots. Effects of different factors, e.g. age, soil type, stand density and geographical location on the results were also analysed.     

A variance-covariance structure to take into account repeated measurements and heteroscedasticity in growth modeling

This study proposes a within-subject variance-covariance (VC) structure to take into account repeated measurements and heteroscedasticity in a context of growth modeling. The VC structure integrates a variance function and a continuous autoregressive covariance structure. It was tested on a nonlinear growth model parameterized with data from permanent sample plots. Using a stand-level approach, basal area growth was independently modeled for red spruce (Picea rubens Sarg.) and balsam fir [Abies balsamea (L.) Mill.] in mixed stands. For both species, the implementation of the VC structure significantly improved the maximum likelihood of the model. In both cases, it efficiently accounted for heteroscedasticity and autocorrelation, since the normalized residuals no longer exhibited departures from the assumptions of independent error terms with homogeneous variances. Moreover, compared with traditional nonlinear least squares (NLS) models, models parameterized with this VC structure may generate more accurate predictions when prior information is available. This case study demonstrates that the implementation of a VC structure may provide parameter estimates that are consistent with asymptotically unbiased variances in a context of nonlinear growth modeling using a stand-level approach. Since the variances are no longer biased, the hypothesis tests performed on the estimates are valid when the number of observations is large.     

Effects of initial stand states on optimal thinning regime and rotation of Picea abies stands

Abstract

This study analysed the effects of young stand characteristics on optimal thinning regime and length of rotation periods for even-aged Norway spruce [Picea abies (L.) Karst.] stands. Stand development was based on a distance-independent, individual-tree growth model. The young stand data were collected from 12 well-stocked Norway spruce stands in southern Finland. Results showed that optimal thinning regimes and rotation period depend on site quality and initial stand characteristics. At the first thinning, optimal thinning type depended on initial density. Thinning from both ends of the diameter distribution turned out to be optimal for initially dense stands. At the second and subsequent thinnings, thinning from above was clearly superior. At a low interest rate, thinning from below was optimal for the first thinning regardless of stocking level. For the study data, optimal rotation periods varied from 61 to 92 years at 3% interest rate. The high variation in length of rotation period was due to the sensitivity of optimal length of rotation period to site qualities, initial stand structure and density.     

Damage by pine weevil Hylobius abietis to conifer seedlings after shelterwood removal

Abstract

Pine weevil (Hylobius abietis L.) damage to seedlings after overstorey removal was investigated in a survey study in six shelterwoods in the south–central part of Sweden. The shelterwoods predominantly consisted of Scots pine, except at one site where the shelter trees mainly consisted of Norway spruce. Before final cutting, 10 plots were laid out at each site and measurements of shelter trees and marked seedlings were taken. The seedlings were examined during the 2 years after final cutting. The study showed that removal of shelter trees increases the risk of severe damage by pine weevil and the variable that was most strongly correlated with the risk was the seedling root collar diameter. Both Scots pine and Norway spruce seedlings were severely damaged by pine weevil, and most of the feeding occurred during the first year after cutting. The amount of debarked area was significantly larger for Scots pine than for Norway spruce seedlings. Vitality (growth of the leading shoot before final cutting) of the seedlings also affected the probability of damage. Seedlings with high vitality were less damaged by pine weevil than seedlings with low vitality. For Scots pine the shelterwood density before final cutting was correlated to the intensity of pine weevil feeding after cutting. In conclusion, after the final cutting of a pine or spruce shelterwood, pine weevils will probably invade the area. To avoid serious damage, Norway spruce and Scots pine seedlings should have reached a diameter of at least 10–12 mm.     

The genetic variation of inter‐provenance hybrids of picea abies and possible breeding consequences

The variance of general combining ability (GCA) and specific combining ability (SCA) for height and diameter growth were estimated from a 13 years old provenance hybridization study in Norway spruce. The GCA variance component for both height and diameter growth was significant, but there was no significant SCA variance. Hybrid progenies outperformed the progenies from stands, but this superiority was not interpreted as a provenance heterosis effect. Estimated family heritabilities for height and diameter growth were moderately high (0.40 for both traits). A consequence of the low SCA variance for Norway spruce may be that clonal testing should replace progeny testing for estimating GCA.     

Forest growth response to changing climate between 1961 and 1990 in Austria

Using 30 years of climate records from 20 weather stations, we investigate the magnitude of temperature and precipitation change, and change in the length of the growing season between 1961 and 1990. Special attention is paid to the period between 1981 and 1990, because recent research suggests that, during this time span, forest productivity may have increased in the northern latitudes. In order to understand the importance of changes in climate on forest growth, we use the ecosystem model FOREST-BGC as a diagnostic tool to predict the annual net primary production (NPP). The results of our study indicate: no change in precipitation between 1961 and 1990; a significant (α = 0.05) increase in mean annual temperature of 0.72°C, mean annual minimum temperature (0.80°C), winter temperature (2.36°C) as well as an increase in the length of the temperature-controlled growing season by 11 days, resulting in a significant increase in diameter increment obtained from 1179 cores of Norway spruce across Austria. The trends in NPP are consistent with observed increment rates validating the use of biogeochemical modeling as a diagnostic tool to search for possible causes on changing environmental conditions.     

Factors affecting the probability of windthrow at stand level as a result of Gudrun winter storm in southern Sweden

A storm in January 2005 in southern Sweden (Gudrun) caused large damage to forests, archaeological sites, buildings, power supply, telecommunications, and infrastructure. In this work, we analyzed the factors affecting the probability of windthrow at stand level (i.e. tree- and stand characteristics) in southern Sweden. We selected a sub-sample (1721 plots) of existing plots from the Swedish National Forest Inventory (inventoried in 2003 and 2004). To assess whether these plots had suffered damage or not, aerial photographs were taken in summer 2005. In order to analyze the effects of measured characteristics on damage, field data for each sub-sample plot were merged with the corresponding result of the aerial photo interpretation.We found that the plots damaged were typically well stocked mature stands, dominated by Norway spruce (Picea abies (L.) Karst.), and often also recently thinned. Probability of damage increased with increase in stand height. In mixed stands, decrease of proportion of Norway spruce decreased the probability of damage. An admixture of deciduous trees, leafless in this winter storm, showed larger effect in this sense than that of Scots pine.     

The impact of Sirococcus shoot blight on radial and height growth of Norway spruce (Picea abies) in young plantations

The impact of Sirococcus shoot blight on the radial and height growth of Norway spruce in a young plantation (approximately 20 years old) was investigated by examining the increment losses for four infection intensities (classes). The average diameter at breast height of trees in the lightly damaged class was 72% when compared with the average diameter of the healthy trees, whereas moderately and severely damaged tree classes were 67 and 57%, respectively. Using tree ring analysis, the development of radial growth over time due to intensity of infection was studied. Height growth of affected trees was also significantly reduced (up to 43%) compared with the healthy trees, thus indicating a dramatic impact of Sirococcus conigenus on the growth of young Norway spruce.     

Sources of within- and between-stand variability in specific leaf area of three ecologically distinct conifer species

Specific leaf area (SLA) is an important ecophysiological variable, but its variability within and between stands has rarely been simultaneously examined and modeled across multiple species. Extensive datasets on SLA in coastal Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco), hybrid spruce (Picea engelmannii Parry × Picea glauca (Moench) Voss × Picea sitchensis (Bong.) Carr.), and ponderosa pine (Pinus ponderosa Dougl. ex P. & C. Laws.) were used to estimate variability of SLA within a canopy and its relationship to tree- and stand-level covariates, and to predict SLA at various locations in tree crowns. Also, in the case of hybrid spruce, variation in SLA due to different relative horizontal lengths from the bole was examined. In all species, SLA systematically increased from tree tip to crown base and decreased with foliage age class. Cardinal direction did not have a highly significant influence in either Douglas-fir or hybrid spruce, but SLA did significantly decrease from branch tip to bole in hybrid spruce. Tree- and stand-level (e.g. density, site index) factors had relatively little influence on SLA, but stand age did have a significant positive influence. For ponderosa pine, a significant relationship between canopy mean current-year SLA and carbon isotope discrimination was also found, suggesting the importance of water stress in this species. An equation was fitted to estimate SLA at various points in the canopy for each species and foliage age class using absolute height in the canopy, relative vertical height in the tree, and stand age.     

Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> L.) regeneration and growth of understory trees under single-tree selection silviculture in Finland

This study presents empirical data on regeneration and growth of understory trees and constructs simple models for predicting these characteristics at various stand structure and post-thinning standing volume levels. The field experiment was established on a grass/herb mineral soil site in central Finland. Regeneration and understory tree growth data were collected from 24 Norway spruce (Picea abies L.)-dominated mixed substands. Seedlings with heights from 5 to 130 cm were measured in 1996 and 2007. In addition, the annual height growth of Norway spruce seedlings was measured in 2007. The spatial pattern of the substands varied from clustered to regular with a decreasing standing volume. Stand complexity was uniform across the range of standing volume. In the 2007 survey, the amount of Norway spruce regeneration increased from approximately 400 to 5,000 seedlings ha⁻¹ when the post-thinning standing volume level was reduced from 230 to 90 m³ ha⁻¹, respectively. Nearly no seedlings were found when the standing volume was over 300 m³ ha⁻¹. The annual diameter increment in Norway spruce understory (dbh < 5 cm) trees decreased, on the average, from 2.3 to 0.3 mm with an increase in the standing volume level from 90 to 340 m³ ha⁻¹, respectively; their height growth showed also a decreasing trend when the standing volume increased. The results indicated that a post-thinning standing volume lower than 150 m³ ha⁻¹ with a regular overstory spatial structure provides a suitable environment for regeneration and growth of Norway spruce understory trees in the studied forest type.     

Effect of planting stocktype and cultivation treatment on the establishment of Norway spruce on cutaway peatlands

In order to determine the effect of stocktype and cultivation treatment on the field performance (survival and growth) and physiological status of Picea abies in cutaway peatlands, small bare-root, large bare-root and containerised seedlings were planted in a deep ploughed and a control site. Survival after 2 years was good across all treatment (>90%) except for the large bare-root seedlings growing in the control site (84%). For all the morphological characteristics assessed in this study, there was no significant interaction between stocktype and cultivation treatment indicating that the growth response to site cultivation was not stocktype dependent. After two growing seasons, all Norway spruce seedlings performed better in the deep ploughed site and displayed also better nutritional and physiological status. Regardless of cultivation treatment, mean height, diameter and volume increment were significantly smaller for the large bare-root seedlings while the small bare-root seedlings displayed the greatest growth rates. In order to promote early height growth in container and small bare-root stock, large diameter is important. Other initial characteristics such as foliar nitrogen content may also have a strong influence on first year field performance. The physiological status of the seedlings during the first year after outplanting was assessed using chlorophyll fluorescence (CF) measurements. CF measurements detected a higher level of stress for the large-bare root stock (low Fv/Fm). On the other hand, small bare-root stock displayed highest maximum potential photochemical activity which corresponded to greatest growth rates. Container seedlings demonstrated higher capacity for photosynthetic electron transport during the first five months after planting suggesting that they recovered from planting stress quicker and optimised better light interception and utilization than bare-root stock. It can be concluded that intensive management systems including deployment of best-adapted stocktype and site cultivation can be used to enhance early height growth of Norway spruce on cutaway peatlands.     

Prediction of individual tree growth in managed stands of mixed Picea abies (L) Karst,and Betula pendula Roth & Betula pubescens Ehrh.

The individual tree growth in stands of mixed Norway spruce (Picea abies (L.) Karst.) and birch (Betula pendula Roth & Betula pubescens Ehrh.) is estimated using basal area and height growth functions for each species separately. The individual tree growth models are distant dependent and the number, size and proximity of neighbours are expressed as size‐ratio competition indices. The competition indices were calculated using a basal area factor gauge to define competitors. The tree growth functions are based on data from nine mixed stands of Norway spruce and birch. The recursive multivariate regression approach is used. The growth functions have standard deviation about the function/standard deviation about the mean (sf/sm) values between 31 and 61% and the evaluations made with root mean square error (rmse) give estimates which vary between 8 and 45 % of the observed mean value. These values are comparable with the precision reported in other studies. In the present investigation the distant dependent indices are important independent variables. It seems suitable to describe the change in growth conditions from retarded to released growth by means of size ratio competition indices. For birch, a positive effect on growth is obtained the more the total competition consists of Norway spruce. For Norway spruce a negative effect on growth is obtained the more the total competition consists of Norway spruce. The lower competitors have a positive effect on the growth of the spruce trees. The interpretation should be that it is better to have a small competitor than a larger one, not that small competitors as such have a positive effect on growth of Norway spruce.     

A Basal Area Increment Model for Norway Spruce in Mixed Stands in Estonia

An individual tree basal area increment model was developed for Norway spruce [Picea abies (L.) Karst] in mixed stands of spruce and birch in Estonia. Different regression equations were fitted for different combinations of variables to obtain biologically tractable interactions between growth and factors affecting it. The best fit index of the regression model was pursued in trials with variable combinations. The basal area increment was chosen to describe the tree growth and both the diameter and age of the tree were included as independent variables. The logical growth relationships were obtained. The basal area increment has a culmination introduced by the simultaneous influence of tree size and age explicitly included in the model. The stand level attributes contributed modestly to the explanatory power of the model because of the narrow range of stand conditions sampled. The present model is applicable to Estonian conditions.