Prediction of yield in the Rothamsted Park Grass Experiment by Ellenberg indicator values

Abstract. The Rothamsted Park Grass Experiment was established in 1856, with experimental plots subjected to annual applications of fertilizer and twice-yearly cutting of hay. There were two major responses to fertilizer, one reflecting high ammonium-nitrogen and increased acidity and the other reflecting high herbage yield without increased acidity. We calculated mean Ellenberg indicator values for N (nitrogen) and R (soil reaction) for the hay harvested between 1948 and 1975, using both unweighted and abundance-weighted means. Plot Ellenberg values were compared with herbage yield and with fertilizer application rates and published soil data. Annual yield of hay varied from 1.5 to 7.4 t/ha and was well predicted by the unweighted mean Ellenberg N-values (r = 0.91). Relatively large negative residuals from the relationship were found in plots whose soil combined low K and low pH. Soil pH was poorly predicted by the unweighted mean R-value, but showed a moderately good relation with weighted mean R (r = 0.73). The fact that Ellenberg N-values correlated better with yield than with applied nitrogen suggests that they might rather be called productivity values.     

The TRM Model of Potential Natural Vegetation in Mountain Forests

Due to advances in spatial modeling and improved availability of digital geodata, traditional mapping of potential natural vegetation (PNV) can be replaced by ecological modeling approaches. We developed a new model to map forest types representing the potential natural forest vegetation in the Bavarian Alps. The TRM model is founded on a three-dimensional system of the ecological gradients temperature (T), soil reaction (R), and soil moisture (M). Within such a “site cube” forest types are defined as homogenous site units that give rise to forest communities with comparable species composition, structure, production and protective functions. The three gradients were modeled using regression algorithms with area-wide, high resolution geodata on climate, relief and soil as predictors and average Ellenberg indicator values for temperature, acidity and moisture of vegetation plots as dependent variables summarizing plant responses to ecological gradients. The resulting predictor-response relationships allowed us to predict gradient positions of each raster cell in the region from geodata layers. The three-dimensional system of gradients was partitioned into 26 forest types, which can be mapped for the whole region. TRM-based units are supplemented by 22 forest types of special sites defined by other ecological factors such as geomorphology, for which individual GIS rules were developed. The application of our model results in an intermediate-scale map of potential natural forest vegetation, which is based on an explicit function of temperature, reaction and moisture and is therefore consistent and repeatable in contrast to traditional PNV maps.     

An application case of ecological indicator values (Zeigerwerte) calculated with a simple algorithmic approach

Abstract

The vegetation of the study site near Rome (Castelporziano Estate), where different woodland types occur, was analysed on the basis of ecological indicator values (Zeigerwerte) for light, temperature, continentality of climate, soil moisture, soil pH and nitrogen. Indicator values were estimated with Hill's reprediction algorithm for the flora of Central-Southern Italy relying on a database of 4,207 original relevés representing a balanced survey of the vegetation of this and surrounding areas. It was possible to obtain indicator values for an important fraction of the Italian Mediterranean flora. Results are ecologically reasonable, and it was possible to find strong correlation between the recalculated values and a few environmental variables. These correlations were not significant in an analogous test with subjectively derived scores of Ellenberg indicator values.     

Reliability of Ellenberg indicator values for moisture,nitrogen and soil reaction: a comparison with field measurements

Abstract. Ellenberg indicator values for moisture, nitrogen and soil reaction were correlated with measured soil and vegetation parameters. Relationships were studied through between‐species and between‐site comparisons, using data from 74 roadside plots in 14 different plant communities in The Netherlands forming a wide range. Ellenberg moisture values correlated best with the average lowest moisture contents in summer. Correlations with the annual average groundwater level and the average spring level were also good. Ellenberg N‐values appeared to be only weakly correlated with soil parameters, including N‐mineralization and available mineral N. Instead, there was a strong relation with biomass production. We therefore endorse Hill & Carey's (1997) suggestion that the term N‐values be replaced by ‘productivity values'. For soil reaction, many species values appeared to need regional adjustment. The relationship with soil pH was unsatisfactory; mean indicator values were similar for all sites at pH > 4.75 because of wide species tolerances for intermediate pH levels. Site mean reaction values correlated best (r up to 0.92) with the total amount of calcium (exchangeable Ca²⁺ plus Ca from carbonates). It is therefore suggested that reaction values are better referred to as ‘calcium values'. Using abundance values as weights when calculating mean indicator values generally improved the results, but, over the wide range of conditions studied, differences were small. Indicator values for bryophytes appeared well in line with those for vascular plants. It was noted that the frequency distributions of indicator values are quite uneven. This creates a tendency for site mean values to converge to the value most common in the regional species pool. Although the effect on overall correlations is small, relationships tended to be less linear. Uneven distributions also cause the site mean indicator values at which species have their optimum to deviate from the actual Ellenberg values of these species. Suggestions for improvements are made. It is concluded that the Ellenberg indicator system provides a very valuable tool for habitat calibration, provided the appropriate parameters are considered.     

Regionalizing Indicator Values for Soil Reaction in the Bavarian Alps – from Averages to Multivariate Spectra

We present an approach to produce maps of Ellenberg values for soil reaction (R-value) in the Bavarian Alps. Eleven meaningful environmental predictors covering GIS-derived information on climatic, topographic and soil conditions were used to predict R-values. As dependent variables, Ellenberg indicator values for soil reaction were queried from plot records in the vegetation database WINALPecobase. We used an additive georegression model, which combines complex prediction models and the increased prediction accuracy of a boosting algorithm. In addition to environmental predictors we included spatial effects into the model to account for spatial autocorrelation. As we were particularly interested in the usefulness of averaged R-values for spatial prediction, we applied two different models: (1) a geo-additive regression model that estimates mean R-values and (2) a proportional odds model predicting the probability distribution over R-values 1 to 9. We found meaningful dependencies between the R-value and our predictors. Both models produced the same spatial pattern of predictions. Spatial effects had an impact only in the first model. The main drawback of mean R-values is the oversimplification of complex conditions of soil reaction, which is entailed by averaging and regression to mean values. Therefore, regionalized average indicator values provide only limited information on site-ecological characteristics. Model 1 failed to predict the range and shapes of original indicator spectra precisely. In contrast, the second model provided a more sophisticated picture of soil reaction. To make the multivariate output of model 2 comparable to that of model 1, we propose to plot the distribution in a three-dimensional color-space. In addition, comparison of both models based on a multiple linear regression model resulted in a R ² of 0.93. The proportional odds model is a promising approach also for other indicator values and different regions as well as for other ordinal-scaled ecological parameters.     

Long‐term impacts of nitrogen and sulphur deposition on forest floor vegetation in the Northern limestone Alps,Austria

Question: Are there effects of long‐term deposition of airborne nitrogen and sulphur on the forest floor vegetation from permanent plots collected in 1993 compared to 2005. Location: Northern limestone Alps in Austria. Methods: Single species responses were analysed by correlating trends in cover‐abundance values, as derived from marginal models, with Ellenberg indicator values. Changes in the species composition of plots were analysed by correlating changes in mean Ellenberg indicator values with the displacement of plots within a multidimensional scaling ordination. Results: Trends in single species abundance were positively correlated with indicator values of soil pH but were independent of nutrient availability. A general trend towards the homogenisation of vegetation, due to convergent time vectors of the relevés, became obvious. Oligotrophic sites previously situated at the distal ends of ordination axes shifted towards the centre since they were enriched by species preferring mesotrophic conditions. The bulk of plots with intermediate site conditions hardly showed any trends. A concomitant analysis demonstrated that temporal changes in species composition exceed the variation in cover abundance estimates among different field botanists. Conclusions: N deposition can lead to a homogenisation of forest floor vegetation. Larger limestone areas with diverse soil conditions, such as the Northern limestone Alps in Austria, as a whole are thus negatively affected by airborne N deposition. Nevertheless, the vegetation was at least as strongly affected by an increase of basiphilous species as a result of decreasing S deposition.     

Application of the Ancient Forest Concept to Potential Natural Vegetation Mapping in Flanders, A Strongly Altered Landscape in Northern Belgium

Construction of potential natural vegetation (PNV) poses particular challenges in landscapes heavily altered by human activity and must be based on transparent, repeatable methods. We integrated the concept of ancient forest (AF) and ancient forest species (AFS) into a four-step procedure of PNV mapping: 1) classification of forest vegetation relevés; 2) selection of those vegetation types that can serve as PNV units, based on AF and AFS; 3) merging of selected vegetation types into five PNV units that can be predicted from a digital morphogenetic soil map; 4) mapping of three additional PNV units based on additional environmental data. The second step, concerning the selection of reference forest vegetation, is of particular interest for PNV construction in Flanders (northern Belgium), where forest cover has been subject to temporal disruption and spatial fragmentation. Among the variety of extant forest recovery states, we chose as PNV units those vegetation types for which a high proportion of relevés had been located in AF and that contained many AFS. As the frequency of AFS depends on site conditions, we only compared and selected vegetation types that are found on similar sites according to average Ellenberg indicator values. While succession is irrelevant for the definition of PNV, colonization rates of AFS can be used to estimate the time required for PNV to be restored in a site.     

Long-term changes in lowland calcareous grassland plots using <Emphasis Type="Italic">Tephroseris integrifolia</Emphasis> subsp. <Emphasis Type="Italic">integrifolia</Emphasis> as an indicator species

We investigated the changes to calcareous grassland plots within protected sites, and whether Tephroseris integrifolia subsp. integrifolia can act as a useful indicator species for re-visitation studies within vegetation predicted to remain relatively stable. Twenty-two plots located across lowland England and all formerly containing T. integrifolia were re-surveyed following the methodology used in the original survey undertaken in the 1960s. Pseudo-turnover and between-observer bias were minimised by sampling replicate quadrats at each fixed plot using a single surveyor and at a similar time of year as the original survey. Qualitative details concerning grazing management were obtained for all sites. In contrast to other long-term re-visitation studies, all our study plots were intact and retained diverse, herb-rich vegetation, demonstrating the value of site protection. However, there were clear shifts in vegetation composition, most notably where T. integrifolia was absent, as shown by an increase in Ellenberg fertility and moisture signifying nutrient enrichment, and a decrease in the cover of low-growing, light-demanding specialists, with a change likely to be associated predominantly with grazing management. Whereas in the mid-20th century the greatest threat to calcareous grassland was habitat loss, undergrazing or temporary neglect now appears to pose the principal threat. Distinctive species such as T. integrifolia with marked sensitivity to habitat change provide a potentially useful tool for rapid assessment and monitoring of site quality. Focusing monitoring on such species allows non-expert observers to recognise the early stages of habitat degradation, providing, in effect, a “health check” on individual sites and groups of sites.     

The forest-cerrado transition zone in southern Amazonia: Results of the 1985 Projeto Flora Amazônica expedition to Mato Grosso

An account of the history of collecting in the Mato Grosso is followed by a discussion of the current views on the nature of the vegetation along the transition from Amazonian forest to cerrado. The purpose of the 1985 Projeto Flora Amazônica expedition was to sample the vegetation in the transition region in northern Mato Grosso. In order to characterize the vegetation at the various localities, the identified collections were assigned to one of five broad distributional categories: Planaltine, Transitional, Amazonian, Wide-spread, or Other. In most cases, as expected, the collecting sites in the north and the west were characterized by Amazonian forest species while those in the south and east comprised mainly cerrado elements. Our collections show that the transition zone is a complex mosaic of Amazonian forest and cerrado formations.     

Shift in Plant Species Composition Reveals Environmental Changes During the Last Decades: A Long-Term Study in Beech (Fagus sylvatica) Forests in Bavaria, Germany

Changes in vegetation composition due to the increasing temperatures in the past few decades have already been reported from several parts of Europe. It has been shown that single species move either northwards or to higher elevations. We expected that the species composition of forest stands should also have changed, i.e., an increase of thermophilous species. Another site factor changing for decades is nitrogen availability; we therefore also expected an increase of nitrophilous species, which was one main result in former long-term studies. We studied the species composition of beech (Fagus sylvatica) forests in southern Germany (Bavaria), comparing old (from 1949 to 1985) and young (2010) phytosociological relevés. Ellenberg indicator values representing plant species specific environmental factors combined with climatic data were used in a partial canonical correspondence analysis (pCCA) for vegetation comparisons. Changes in plant species composition were analyzed considering species frequency, distribution of Ellenberg indicator values, shares of i) non-native plants and ii) tree, shrub and herbaceous species. Contrary to our expectations, global warming in Bavaria during the past decades resulted only in an explained dispersion of 5 % in the species composition. On the species level, an overall increase of thermophilous walnut tree saplings (Juglans regia) was conspicuous in some study areas. Nitrophilous species, however, generally increased in frequency throughout the study areas. Throughout Bavaria the most significant change was a striking increase of juvenile tree species and a decrease of herbaceous species. Up to now the increased nitrogen input into forests had a stronger provable influence on species composition shift than the global warming of the last decades. Additionally general changes in forest management also had effects on forest species. Therefore community reorganization mirroring temperature factors in beech forests in central Europe seems to be only at its very beginning.     

Short-term and long-term variation in seed bank/vegetation relations along an environmental and successional gradient

The seed bank along a successional and environmental gradient was analysed. Soil was collected in 3-cm thick horizons from permanent plots along two transects across a land uplift seashore, spanning several centuries of succession from shoreline to mature forest. Vegetation in the plots was recorded when the soil was sampled and also 9 and 15 yr before that. Within- and between-plot effects on seed bank./vegetation relationships were analysed using estimates of seed longevity. Sorensen's similarity index and mean Ellenberg indicator values.
A seed bank longevity index was constructed by using the database by Thompson et al, (1997 The soil seed banks of north west Europe. Methodology, density and longevity, Cambridge Univ Press), for all species with more than one entry in the database. For species with one or no entry, an internal Index was constructed. The two indices were correlated and it was suggested that the internal index should be used where the Thompson database is insufficient.
There were small differences between the upper three soil horizons in seed density, in similarity with the vegetation and in mean Ellenberg values. The highest seed densities and seed bank/vegetation similarities were found at the shoreline, after that the density and the similarity decreased with increasing successional age, with the mature forest having very low seed density and similarity values. Weighted mean Ellenberg indicator values for light, nitrogen, salt and moisture differed between vegetation and seed bank. For the seed bank, the mean Ellenberg values for light, moisture and nitrogen and weighted mean of seed bank longevity indices showed a trend along one of the transects.     

Comparative life history of aspidiaceous ferns in northern Japan with reference to fertility during sporophyte development in relation to habitats

The life-cycle characteristics of ferns, in particular reproductive characteristics in relation to alternation of generations, have been studied in populations of 19 species of aspidiaceous ferns native to Hokkaido. The developmental age of the sporophyte was quantitatively expressed by the number of midrib branches of the leaf (NV; number of venation). The maximum developmental age (MNV) of the aspidiaceous ferns in Hokkaido ranged from 49 to 113. The initiation of fertility (IF) in the relative developmental age (RDA), percentages of any NV to the maximum NV of the leaf population) ranged from 15 to 65%. Ferns occurring on rocky cliffs initiated fertility at 15–21%, while ferns occurring on the forest floor initiated fertility at 41–65% in the RDA of a leaf population. Further observations were undertaken on 26 ferns of other families and revealed that there were correlations between habitat and maximum NV, and habitat and IF in the RDA. Ferns occurring on cliffs, on mossy rocks and on tree trunks, had lower maximum NV of the leaf and earlier initiation of fertility in the RDA, than those of ferns occurring on the forest floor, edge of forests and on grassland in Hokkaido. Ferns occurring in the exposed habitat seemed to behave as a r-strategist, while ferns occurring in the forest floor seemed to behave as a K-strategist.     

Predicting forest site productivity in temperate lowland from forest floor, soil and litterfall characteristics using boosted regression trees

Aims

The aim of this study is on the one hand to identify the most determining variables predicting the site productivity of pedunculate oak, common beech and Scots pine in temperate lowland forests of Flanders; and on the other hand to test whether the accuracy of site productivity models based exclusively on soil or forest floor predictor variables is similar to the accuracy achieved by full ecosystem models, combining all soil, vegetation, humus and litterfall composition related variables.

Methods

Boosted Regression Trees (BRT) were used to model in a climatically homogeneous region the relationship between environmental variables and site productivity. A distinction was made between soil (soil physical and chemical), forest floor (vegetation and humus) and ecosystem (soil, forest floor and litterfall composition jointly) predictors.

Results

Our results have illustrated the strength of BRT to model the non-linear behaviour of ecological processes. The ecosystem models, based on all collected variables, explained most of the variability and were more accurate than those limited to either soil or forest floor variables. Nevertheless, both the soil and forest floor models can serve as good predictive models for many forest management practices.

Conclusions

Soil granulometric fractions and litterfall nitrogen concentrations were the most effective predictors of forest site productivity in Flanders. Although many studies revealed a fertilising effect of increased nitrogen deposition, nitrogen saturation seemed to reduce species’ productivity in this region.     

Simulation of multidimensional community patterns: towards a comprehensive model

Ecological gradients in the field layer of southern boreal forests in South Finland were studied in relation to the dominant tree species and the age of forest stands. The data are from a systematic sample of 529 plots from an area of 150 × 200 km, collected in the Third National Forest Inventory in 1951–53. Detrended correspondence analysis (DCA) was applied to log-transformed species cover values. It revealed three main gradients: fertility, moisture, and the effect of cattle grazing in forests (still extensive in the early 1950's). The fertility gradient dominated the first axis and the two latter sources of variation confounded with it in a complex manner in the first two axes of DCA. The second DCA axis was associated with canopy effects on understory pattern, with Pinus and Picea having opposite and Betula intermediate effects. These results were compared with an ordination model of Cajander's forest site types, based on DCA of independent, ideal data of 107 indicator species. The fertility gradient recovered by the model was almost identical to that obtained from the field data. The gradient was also stable from intermediate-age (40–69 yrold) to older forests. The forest site types showed rather large overlaps with main neighbouring types in composition of ground vegetation or nutrient status of the humus. Competitively efficient feather-mosses, which are dependent on nutrients released from the tree crowns, are considered important regulators of the understory vegetation. Accordingly, alternative approaches to the forest site type classification to be used in boreal forests treated by modern intensive forestry should give more weight to the effect of the canopy trees.     

The potential of generalized additive modelling for the prediction of radial growth of Norway spruce from Central Germany

During the past decades managed forest ecosystems in Central Europe underwent vast changes, induced by extreme climate conditions and occasionally adverse forest management. Tree ring width patterns mirror these changes and thus have been widely examined as environmental archives and reliable empirical data sources in ‘tree growth modelling’. Dendrochronologists often suppose linear co-variation among the covariates, variable independence and homoscedasticity. Conventionally, these assumptions were achieved by eliminating biological age trends (detrending) and removing the autocorrelation from the time series (pre-whitening). Particularly detrending might be biased according to the scientific problem and sometimes inflexible age models. In this study, we tackle these issues and examine the suitability of a flexible Generalized Additive Model (GAM) on recently developed tree ring width time series of 30 Norway spruce stands (Picea abies [L.] H. Karst) from Central Germany.The model was established to simultaneously cope with the mentioned detrending issue, to unravel nonlinear climate-growth relationships and to predict mean ring width time series for spruce stands in the region. Particularly the latter was of primary interest, since recent forest planning relies on static yield tables that often underestimate the actual growth.The model reliably captured the empirical data, indicated by a small Generalized Cross Validation criterion (GCV = 0.045) and a deviance explained of 88.6 %. The flexible additive smoothers accounted for the social status of individual trees, captured low frequency variations of changing growth conditions adequately and displayed a rather flat biological age trend. The radial increment responded positively to summer season precipitation of the current and previous year. Positive temperature responses were found during the early vegetation period, whereas high summer season temperatures negatively affected the radial growth. The seasonal transition from spring to summer in June induced a shift in the climate response of the linear predictor, leading to a distinct negative effect of temperature and a no-role of precipitation on the linear predictor.Most important, utilizing the calibrated GAM for the purely climate-driven prediction of mean ring width time series from five independent spruce sites revealed proper coherencies. Herein, the mean ring width for sites located within the climatic-optimum for spruce growth were more exactly predicted than for sites with adverse spruce growth conditions. In addition, large mean ring widths were systematically underestimated, whereas small mean ring widths were precisely predicted. Overall, we strongly recommend GAMs as a powerful tool for the investigation of nonlinear climate-growth relationships and for the prediction of radial growth in managed forest ecosystems.     

A comparison of biological characteristics and distribution between Swedish threatened and non-threatened forest vascular plants

Threatened (n = 59) and non-threatened (n = 308) Swedish forest vascular plant taxa were compared with regard to a number of variables, including distribution, site factors taxonomy, morphology and flowering time A majority of the threatened taxa occur in the southern deciduous woodlands, which only constitute c 0.5% of the total forested area in Sweden There are considerably more threatened taxa in southern than in northern Sweden, a consequence of the successively higher number of forest vascular plants from the north towards the south Threatened taxa grow in forests with significantly higher soil fertility than non-threatened taxa Significant differences were also revealed regarding light conditions, soil water conditions and month of flowering Threatened taxa grow on soils with significantly higher pH and also with slightly more available nitrogen than non-threatened taxa, as measured with Ellenberg indicator values Forest stands on fertile soils are uncommon in Sweden and they are also very species-rich Soil-type rarity in combination with high species diversity in these soil types thus partly explain why taxa are included in the Swedish Red data list     

Amelanchier alnifolia vegetation in eastern Idaho,USA and its environmental relationships

Fifteen shrub stands of Amelanchier alnifolia vegetation were studied using Braun-Blanquet procedures, tabulated, and ordered by Twinspan, Decorana, and field experience into 5 groups of stands. The stands occurred on the lower slopes of the southwestern Teton and southeastern Big Hole mountains of eastern Idaho and in-and outside a U.S. Forest Service exclosure above the Hoback River in adjacent Wyoming. The ecology of this Amelanchier vegetation as a whole and of the 5 groups of stands is discussed using Jenny's factors of soil formation (1941, 1958, 1980) as a framework. Plant indicator values (Ellenberg 1979, 1988) which rank species on their occurrences in relation to light, temperature, continentality of climate, water availability, soil reaction (pH), and soil nitrogen supply, were independently assigned to the 121 species of vascular plants encountered in the described stands.     

NIRS meets Ellenberg's indicator values: Prediction of moisture and nitrogen values of agricultural grassland vegetation by means of near-infrared spectral characteristics

Ellenberg indicator values are widely used ecological tools to elucidate relationships between vegetation and environment in ecological research and environmental planning. However, they are mainly deduced from expert knowledge on plant species and are thus subject of ongoing discussion. We researched if Ellenberg indicator values can be directly extracted from the vegetation biomass itself. Mean Ellenberg “moisture” (mF) and “nitrogen” (mN) values of 141 grassland plots were related to nutrient concentrations, fibre fractions and spectral information of the aboveground biomass. We developed calibration models for the prediction of mF and mN using spectral characteristics of biomass samples with near-infrared reflectance spectroscopy (NIRS). Prediction goodness was evaluated with internal cross-validations and with an external validation data set. NIRS could accurately predict Ellenberg mN, and with less accuracy Ellenberg mF. Predictions were not more precise for cover-weighted Ellenberg values compared with un-weighted values. Both Ellenberg mN and mF showed significant and strong correlations with some of the nutrient and fibre concentrations in the biomass. Against expectations, Ellenberg mN was more closely related to phosphorus than to nitrogen concentrations, suggesting that this value rather indicates productivity than solely nitrogen. To our knowledge we showed for the first time that mean Ellenberg indicator values could be directly predicted from the aboveground biomass, which underlines the usefulness of the NIRS technology for ecological studies, especially in grasslands ecosystems.     

Measurement errors and regression to the mean cannot explain bias in average Ellenberg indicator values

Smart & Scott (2004, this is sue) criticized our paper (Wamelink et al. 2002) about the bias in average Ellenberg indicator values. Their main criticism concerns the method we used, regression analysis. They state the bias can be mimicked by the construction of an artificial data set and that regression analysis is not a suited tool to investigate underlying phenomena. Moreover they claim that the present bias is caused by the distribution of Ellenberg indicator values between syntaxa, instead of a bias in average Ellenberg indicator values per species. We show that their criticism of the use of regression analysis does not hold. We selected average Ellenberg values per vegetation group for several pH classes and applied an F‐test to determine whether or not the vegetation groups within each pH class differed significantly from each other. This was the case for all tested classes (P < 0.001). Moreover we simulated an artificial data set, of which the F‐test for varying measurement error could not explain the magnitude of the F‐value we found earlier. This indicates that the bias we found in average Ellenberg indicator values cannot be explained by measurement errors or by regression to the mean. In the end, Smart & Scott, as we did, come to the conclusion that there is a bias present and that separate regression lines per vegetation type are necessary, but the debate remains open on whether or not this is caused by the bias in Ellenberg indicator values per species.     

Simulating forest succession along ecological gradients in southern Central Europe

Forest stand development was simulated using a forest succession model of the JABOWA/FORET type. The environmental conditions are representative for a wide spectrum of Swiss forest sites ranging from 220 m to 1 700 m a.s.l. Each model run covers a period of 1 200 yr and is based on the averaged successional characteristics of 50 forest plots with an individual size of 1/12 ha. These small forest plots serve as basic units to simulate establishment, growth, and death of individual trees of 29 species. Existing light in the forest stand, climatic conditions, soil properties, and other environmental factors control the growth of each individual tree. Compared with previous simulation studies, some major modifications were made, including the incorporation of the indicator values of Ellenberg (1979) to describe the ecophysiological behaviour of the species considered. As a test, the simulated species composition through time was compared with the actual vegetation and the potentially natural species composition on the corresponding site types. The extensive comparison revealed that approximately 80% of the simulations match the expected species configurations. Thus, it was concluded that the model is valid for the purpose of evaluating impacts of natural and human disturbances on forest communities.