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.     

Validity of Ellenberg indicator values judged from physico‐chemical field measurements

Abstract. The relationship between mean Ellenberg indicator values (IV) per vegetation relevé and environmental parameters measured in the field usually shows a large variation. We tested the hypothesis that this variation is caused by bias dependent on the phytosociological class. For this purpose we collected data containing vegetation relevés and measured soil pH (3631 records) or mean spring groundwater level (MSL, 1600 records). The relevés were assigned to vegetation types by an automated procedure. Regression of the mean indicator values for acidity on soil pH and the mean indicator values for moisture on MSL gave percentages explained variance similar to values that were reported earlier in literature. When the phytosociological class was added as an explanatory factor the explained variance increased considerably. Regression lines per vegetation type were estimated, many of which were significantly different from each other. In most cases the intercepts were different, but in some cases their slopes differed as well. The results show that Ellenberg indicator values for acidity and moisture appear to be biased towards the values that experts expect for the various phytosociological classes. On the basis of the results, we advise to use Ellenberg IVs only for comparison within the same vegetation type.     

Shifts in ecological behaviour of herbaceous forest species along a transect from northern central to North Europe

We investigated the ecological behaviour (the response to environmental factors in the field, synonymous to the term realized niche) of four closely related species pairs (Melica nutans, M. uniflora; Primula veris, P. elatior; Veronica chamaedrys, V. montana; Viola riviniana, V. reichenbachiana) across a transect from northern Central to North Europe. The second-mentioned species of each pair is confined in its geographical distribution to the southern parts of the studied transect. Sample plot data of deciduous forests were compiled from (1) Germany, S Niedersachsen, (2) Germany, northern Schleswig-Holstein, (3) Denmark, and (4) Boreo-nemoral Sweden. We compared the ecological optima and amplitudes of the response curves of species along the gradients for moisture, pH and nitrogen by means of phytosociological data, detrended correspondence analysis (DCA) and Ellenberg indicator values. pH measurements from Sweden were significantly correlated with the corresponding DCA sample plot scores and the plot averages of the Ellenberg values for reaction (pH). In accordance with our main hypothesis, the wide range species appeared to have broader ecological amplitudes on the northern margins of their distributional ranges, especially in Boreo-nemoral Sweden, than in the southern parts of the study area. Our findings are in contrast to theories claiming a reduced niche breadth of range-margin populations of species compared to range-centre populations. The shifts in ecological behaviour were particularly obvious with respect to soil acidity. We believe that these shifts are caused by changes in the competitive relationships between the species: in the north, the total pool of species in deciduous forests is comparatively small. The low species richness is likely to lead to reduced competition and to an expansion of the ecological amplitude, known as competitive release.     

Calibration of Ellenberg indicator values for the Faroe Islands

Abstract. Elenberg's bio‐indication system for soil moisture (F), soil nitrogen (N) and soil reaction (R) was examined, based on 559 vegetation samples and environmental characteristics (vegetation cover, soil depth, soil moisture, chemical soil properties) from four Faroe islands. The original indicator values from central Europe were used for the calculation of weighted community indicator values of F, N and R. These were regressed with respect to environmental data, applying standard curvilinear regression and generalized linear modelling (GLM) and new predicted values of community indicator values were obtained from the best model. Faroe species optima values of 162 taxa for one or more of the three EUenberg scales were derived from fitting Huisman‐Olff‐Fresco (HOF) models of species abundance with respect to predicted community indicator values and are proposed as new EUenberg species indicator values to be used in the Faroe Islands. F was best correlated with a GLM model containing soil moisture, organic soil fraction, soil depth and total vegetation cover, R with a GLM model containing pH and calcium in % organic soil fraction, N with total phosphorus in % organic soil fraction. The calibrated species indicator scales are much truncated, as compared with the original values, resulting in significantly different overall distributions of the original and new species indicator values. The recalculated community indicator values are much better correlated to environmental measurements. Several species do not have clear optima, but linear or monotone relationships to the examined indicator scales. This probably indicates that the occurrence of some species in the Faroe Islands are either determined by factors other than moisture, pH or soil nutrient status or, given the young age and environmental instability of the islands, are governed by stochastic mechanisms. Extension of Ellenberg indicator values outside central Europe should always be carefully calibrated by means of adequate environmental data and adequate statistical models, such as HOF models, should be applied.     

Factors affecting Sciomyzidae (Diptera) across a transect at Skealoghan Turlough (Co. Mayo,Ireland)

Sciomyzid flies, which have potential as bio-indicators, were sampled by sweep-net surveys at a turlough in the west of Ireland. Turloughs are ephemeral wetlands (unique to Ireland), which flood from groundwater in winter and empty in the summer, during that time, they are frequently grazed. The weekly survey consisted of ten linear sweeps (5 m × 1 m) in each of six homogeneous and contiguous vegetation zones throughout the summer of 2004. The fauna was dominated by univoltine species with Ilione albiseta being particularly abundant, though species displaying a range of phenologies and life histories were also present. Species richness and total abundance were significantly higher in the two zones with the highest hydroperiods. Mantel tests showed that the species matrix was significantly co-structured with permanent features of the physical environment, but not with stochastic sampling variables related to weather conditions. Mantel correllograms displayed typical patterns of autocorrelation for hydroperiod, soil moisture and soil pH in each zone and vegetation height, vegetation length and Ellenberg moisture index (weighted for vegetation composition) in each sweep-path. No patterns of autocorrelation were evident for distance among zones, area of patch of vegetation zone sampled, area of the vegetation zone on the whole turlough, soil mass-loss-on-ignition and Ellenberg N and reaction indices. These results provide strong evidence for high microhabitat specificity in Sciomyzidae at this site and indicate a major influence of vegetation structure and hydrological regime on their ecology.

Prediction of occurrence of vascular plants in deciduous forests of South Sweden by means of Ellenberg indicator values

Abstract. In this study we present a new method for predicting the occurrences of species using data from deciduous forests in South Sweden. Complete species lists of vascular plants were compiled from 101 stands and from representative sample plots inside the stands. Soil samples from each stand were collected for determination of pH and nitrogen mineralization. Presence-absence data for species were fitted to the values of four environmental variables - soil moisture, soil reaction (pH), soil nitrogen and light - by means of Linear (Multiple) Logistic Regression (LLR), and Gaussian (Multiple) Logistic Regression (GLR). First, these values were estimated by calculating the weighted averages of Ellenberg indicator values. Second, the estimates for reaction and nitrogen were substituted by the real measurements of pH and mineralized NH₄⁺, keeping the Ellenberg estimates for light and moisture. The models were validated by an independent test data set. In general, the models had high predictive abilities. GLR fitted the species occurrences better to the environmental variables than LLR, but had a lower accuracy of prediction of species occurrence in the stands. The use of soil measurements instead of Ellenberg indicator values did not improve the predictive abilities of the models. The environmental conditions in the stand test set were successfully estimated by using species data from the plots. When using the species lists of the stands instead of plot data, a slightly better predictive ability was obtained. The collection of plot data, however, is easier and less time-consuming. The accuracy of prediction differed considerably between species.     

pH optima for Danish forest species compared with Ellenberg reaction values

Species distribution depends on the physiological and ecological niche where a species can exist and regenerate in resource competition with other species (niche limitation). The realized niche is influenced by local biotic processes that influence species behaviour and the shape of the response curves relative to environmental gradients. Processes on larger scales also influence the species niche through source-sink mechanisms (dispersal limitation) and the species richness of an area (pool limitation). Despite the growing evidence of skewed or irregular species response curves along gradients, many ecologists still assume symmetric, unimodal response curves along gradients in ecological interpretation. Ellenberg’s indicator system is probably the most common example. However, the assumption is not ecologically or statistically valid, due to the many different processes affecting the distribution of plant species. Here I present the results of Huisman-Olff-Fresco (HOF) regressions for 209 Danish forest species. HOF modelling is chosen to avoid the classical drawbacks of assuming symmetric, unimodal response patterns. I calculate the optima for all species with unimodal responses to soil pH and compare these with the Ellenberg indicator values for reaction (R), which are often used as a substitute for soil pH measurements. I demonstrate that the assumption of symmetric, unimodal species behaviour is violated in 54% of the cases and that pH optima and R indicator values for species are not always compatible. Ellenberg reaction scale has been used byEwald (Folia Geobot. 38: 357–366, 2003) as an indicator of which species are calcicole, i.e., whether they can grow and reproduce on calcareous soils. Such affinities of species, however, are related to both local niche properties and processes on large scales and cannot be generalized from a single empirical variable such as pH, nor from Ellenberg semi-ordinal indicator scale. I conclude that while the determination of whether species are calcicole or calcifuge requires more research, it is evident that Denmark contains a fairly balanced number of calciphytic and acidophytic species. This is probably due to the nearly equal areas with acidic and alkaline soils in Denmark, which also contribute to the high species richness of more than 500 vascular plant species in Danish forests.     

Use and improvement of Ellenberg's indicator values in deciduous forests of the Boreo-nemoral zone in Sweden

Ellenberg's indicator values for light, moisture and reaction were tested in deciduous hardwood forests of the Boreo-nemoral zone m Sweden Weighted averages of indicator values were calculated and correlated with field measurements for two test sets of relevés The best, highly significant correlations were found for reaction, particularly for weighted averages based on vascular plant species Significant correlations were also found for light, whereas those for moisture were less good and only partly significant Weighted averages based on presence/absence values and those based on cover/abundance values were generally very similar to each other, but showed differences in their correlations with actual measurements
A training set of relevés was used to calculate optima and ecological amplitudes of the most common species Under certain conditions, the original indicator values were replaced by these optima in order to obtain improved indicator values for Swedish deciduous forests These were correlated with the same test sets of field measurements as the original values and clearly gave better results for light as well as for reaction regarding vascular plants No clear differences were found for moisture and reaction regarding bryophytes For the majority of species, the original indicator values also expressed the ecological optima in Swedish deciduous forests as determined by this study However, several species, most of them preferring more open habitats, showed ecological optima and amplitudes that considerably deviated from central European conditions In conclusion, Ellenberg's indicator values can successfully be used in south Swedish deciduous forests, particularly after calibration of the values according to regional deviations     

Ellenberg’s water table experiment put to the test: species optima along a hydrological gradient

An important aspect of niche theory is the position of species’ optima along ecological gradients. It is widely believed that a species’ ecological optimum takes its shape only under competitive pressure. The ecological optimum, therefore, is thought to differ from the physiological optimum in the absence of interspecific competition. Ellenberg’s Hohenheim water table experiment has been very influential in this context. However, the water table gradient in Ellenberg’s experiment was produced by varying the soil thickness above the water table, which confounded the potentially disparate impacts of water table depth (WTD) and soil depth on species growth. Accordingly, here we have re-evaluated Ellenberg’s work. Specifically, we tested the hypothesis that physiological and ecological optima are identical and unaffected by interspecific interaction. We used the same six grasses as in Ellenberg’s experiments, but in our mesocosms, WTD was varied but soil depth kept constant. The design included both an additive component (with/without plant interaction) and a substitutive component (monocultures vs. species mixtures). The results show that the physiological optima along the hydrological gradient varied greatly between species, even in the absence of interspecific interaction. Within species, however, physiological and ecological optima appeared identical in most cases, irrespective of the competition treatment. We conclude that the ‘physiological capacity’ of species largely determines where they are able to persist and that any impact of interspecific interaction is only marginal. These findings are at variance with Ellenberg’s rule, where competition is considered to shift the distribution of a species away from its physiological optimum.     

Species Richness, Community Specialization and Soil-Vegetation Relationships of Managed Grasslands in a Geologically Heterogeneous Landscape

The increasing importance of the conservation value of managed grasslands has led to many studies exploring edaphic determinants of grassland biodiversity. Most studies, however, come either from very large areas, where biogeographical factors such as dispersal limitation may play a role, or from small, but ecologically rather uniform, regions. In addition, few studies further distinguish between plant specialists and generalists in the interpretation of the observed patterns. Here we studied species richness in semi-natural, managed grasslands in the Strá?ovské vrchy Mountains in the West Carpathians, Slovakia, where there is a matrix of different bedrocks (crystalline, sandstone, claystone, limestone) on a steep altitudinal gradient. In 89 vegetation plots we sampled the species composition of vascular plants and bryophytes and measured soil chemistry, slope angle, heat index, altitude and soil depth. We further applied Ellenberg indicator values and classified species into community specialists or generalists based on the analysis of a large phytosociological database. Using cluster analysis, we delimited five vegetation types that clearly differed in response to soil characteristics. Species richness varied between 19 and 64 species per 16?m². The main compositional gradient correlated with measured soil pH and calcium, but species richness was not significantly correlated with these factors. Soil available phosphorus was not associated with species composition as has been found elsewhere, but it did correlate negatively with species richness and the richness of specialists. Overall, species richness was largely driven by the number of specialists in the plot and particular vegetation types differed conspicuously in their number. We further found significant effects of iron, potassium and sodium on species richness, species composition and the representation of specialists and generalists. Our results provide new insights into the determinants of diversity in managed grasslands as well as to the theoretical species pool concept, explaining species richness variation along a pH gradient.     

杭州市区土生苔藓植物分布与生态因子的关系

在杭州市区设立的21个样点中,记录到土生苔藓植物22科31属47种.基于样点中苔藓植物种类的生态重要值和样点生态因子数据进行典范对应分析.结果表明:人为干扰和土壤pH是影响杭州市区土生苔藓植物分布的主要生态因子.在市区平原地势的公园和绿地,人为干扰较大,土壤呈碱性,苔藓植物种类以小羽藓属、真藓属和丛藓科为主;而在海拔较高的丘陵地区,人为干扰较小,土壤呈酸性,苔藓植物种类相对丰富,侧蒴藓类和苔类较多.对47种土生苔藓植物进行生态位宽度计测,多数苔藓植物的生态位宽度较窄.其中,东亚拟鳞叶藓的生态位宽度最大(0.3510),广泛分布在西湖西南的丘陵地区;平叶毛口藓和细叶小羽藓的生态位宽度次之,分别为0.2239和0.2185,是市区平原地势的公园和绿地中的常见种.     

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.     

Empirical realised niche models for British higher and lower plants – development and preliminary testing

Question: Can useful realised niche models be constructed for British plant species using climate, canopy height and mean Ellenberg indices as explanatory variables? Location: Great Britain. Methods: Generalised linear models were constructed using occurrence data covering all major natural and semi‐natural vegetation types (n=40 683 quadrat samples). Paired species and soil records were only available for 4% of the training data (n=1033) so modelling was carried out in two stages. First, multiple regression was used to express mean Ellenberg values for moisture, pH and fertility, in terms of direct soil measurements. Next, species presence/absence was modelled using mean indicator scores, cover‐weighted canopy height, three climate variables and interactions between these factors, but correcting for the presence of each target species in training plots to avoid circularity. Results: Eight hundred and three higher plants and 327 bryophytes were modelled. Thirteen per cent of the niche models for higher plants were tested against an independent survey dataset not used to build the models. Models performed better when predictions were based only on indices derived from the species composition of each plot rather than measured soil variables. This reflects the high variation in vegetation indices that was not explained by the measured soil variables. Conclusions: The models should be used to estimate expected habitat suitability rather than to predict species presence. Least uncertainty also attaches to their use as risk assessment and monitoring tools on nature reserves because they can be solved using mean environmental indicators calculated from the existing species composition, with or without climate data.     

Using topographic wetness index in vegetation ecology: does the algorithm matter?

Questions: How important is the choice of flow routing algorithm with respect to application of topographic wetness index (TWI) in vegetation ecology? Which flow routing algorithms are preferable for application in vegetation ecology? Location: Forests in three different regions of the Czech Republic. Methods: We used vegetation data from 521 georeferenced plots, recently sampled in a wide range of forest communities. From a digital elevation model, we calculated 11 variations of TWI for each plot with 11 different flow routing algorithms. We evaluated the performance of differently calculated TWI by (1) Spearman rank correlation with average Ellenberg indicator values for soil moisture, (2) Mantel correlation coefficient between dissimilarities of species composition and dissimilarities of TWI and (3) the amount of variation in species composition explained by canonical correspondence analysis. Results: The choice of flow routing algorithm had a considerable effect on the performance of TWI. Correlation with Ellenberg indicator values for soil moisture, Mantel correlation coefficient and explained variation doubled when the appropriate algorithm was used. In all regions, multiple flow routing algorithms performed best, while single flow routing algorithms performed worst. Conclusions: We recommend the multiple flow routing algorithms of Quinn et al. and Freeman for application in vegetation ecology.     

Germination and Seedling Responses of Subalpine Plants to Different Soil Substrates

In this study we explored the regeneration niche of 18 alpine plant species by testing the germination responses and seedling growth to soil mixtures made with gypsum, ophiolite, and quartzite, and a control soil. We analyzed different phases of germination: time of radicle emergence, germination duration, and germination percentage, and thereafter seedling performance measured using seedling biomasses. Species were grouped as acidophilous, basophilous and neutrophilous according to their Ellenberg indicator values. Seeds were germinated on each soil mixture in a greenhouse, and then seedlings were selected and transplanted outdoors to grow for one season at 2,050 m a.s.l. The time of radicle emergence, length of germination, final germination percentage, seedling biomass and specific leaf area (SLA) varied according to the soil mixtures depending on the pH group. There were some consistencies between the Ellenberg groups, which were built for adult plant species, and the way these seedlings responded to a particular soil mixture. For instance, the group of species predominantly occurring in basic soils during their adult life (basophilous) had the highest germination percentage and the shortest emergence time on the soil mixture composed of gypsum, but a low germination percentage and the smallest seedling biomass on the other two soil mixtures. In addition, the decrease in SLA for seedlings of the basophilous group when they were cultivated on acidic soil (especially quartzite) seems to indicate a weaker functioning of the plants. Our results highlight, first, the significance of the regeneration niche in the establishment of plants in a given soil environment, especially by emphasizing the links between germination and soil chemistry, and secondly, the consideration that different germination phases add more information about the plant community assemblage with respect to the soil environment.     

Calibrating Ellenberg indicator values for moisture,acidity, nutrient availability and salinity in the Netherlands

A general calibration of Ellenberg indicator values for moisture, acidity, nutrient availability and salinity was carried out on a large database of relevées and environmental variables from a variety of ecosystems in the Netherlands.Satisfying relationships with Ellenberg indicator values for moisture, acidity and salinity were found for mean groundwater level in spring time, soil pH and chloride concentration in groundwater. For mean groundwater level in spring and chloride concentration in groundwater subdivision of the database led to clearer relationships with indicator values. For the Ellenberg indicator value for nutrient availability satisfying calibration results were only achieved with data on standing crops and N stock in standing crop. The relationship with soil chemical variables was less clear.Although the correlation between indicator and measured values is obvious, the variation around the regression lines is considerable. However, because of the size and composition of the database, it is unlikely that our calibration results can be much improved by adding more (Dutch) data.The calibration results will be applied in the multi-stress model SMART-MOVE, developed to predict changes in species composition due to acidification, eutrophication and the effects of lowering groundwater.     

Plant functional trait response to environmental drivers across European temperate forest understorey communities

• Functional traits respond to environmental drivers, hence evaluating trait‐environment relationships across spatial environmental gradients can help to understand how multiple drivers influence plant communities. Global‐change drivers such as changes in atmospheric nitrogen deposition occur worldwide, but affect community trait distributions at the local scale, where resources (e.g. light availability) and conditions (e.g. soil pH) also influence plant communities.
• We investigate how multiple environmental drivers affect community trait responses related to resource acquisition (plant height, specific leaf area (SLA), woodiness, and mycorrhizal status) and regeneration (seed mass, lateral spread) of European temperate deciduous forest understoreys. We sampled understorey communities and derived trait responses across spatial gradients of global‐change drivers (temperature, precipitation, nitrogen deposition, and past land use), while integrating in‐situ plot measurements on resources and conditions (soil type, Olsen phosphorus (P), Ellenberg soil moisture, light, litter mass, and litter quality).
• Among the global‐change drivers, mean annual temperature strongly influenced traits related to resource acquisition. Higher temperatures were associated with taller understoreys producing leaves with lower SLA, and a higher proportional cover of woody and obligate mycorrhizal (OM) species. Communities in plots with higher Ellenberg soil moisture content had smaller seeds and lower proportional cover of woody and OM species. Finally, plots with thicker litter layers hosted taller understoreys with larger seeds and a higher proportional cover of OM species.
• Our findings suggest potential community shifts in temperate forest understoreys with global warming, and highlight the importance of local resources and conditions as well as global‐change drivers for community trait variation.

     

Using species‐environmental amplitudes to predict pH values from vegetation

Question: Species optima or indicator values are frequently used to predict environmental variables from species composition. The present study focuses on the question whether predictions can be improved by using species environmental amplitudes instead of single values representing species optima. Location: Semi‐natural, deciduous hardwood forests of northwestern Germany. Methods: Based on a data set of 558 relevés, species responses (presence/absence) to pH were modelled with Huisman‐Olff‐Fresco (HOF) regression models. Species amplitudes were derived from response curves using three different methods. To predict the pH from vegetation, a maximum amplitude overlap method was applied. For comparison, predictions resulting from several established methods, i. e. maximum likelihood/present and absent species, maximum likelihood/present species only, mean weighted averages and mean Ellenberg indicator values were calculated. The predictive success (squared Pearson's r and root mean square error of prediction) was evaluated using an independent data set of 151 relevés. Results: Predictions based upon amplitudes defined by maximum Cohen's x probability threshold yield the best results of all amplitude definitions (R²= 0.75, RMSEP = 0.52). Provided there is an even distribution of the environmental variable, amplitudes defined by predicted probability exceeding prevalence are also suitable (R²= 0.76, RMSEP = 0.55). The prediction success is comparable to maximum likelihood (present species only) and – after rescaling – to mean weighted averages. Predicted values show a good linearity to observed pH values as opposed to a curvilinear relationship of mean Ellenberg indicator values. Transformation or rescaling of the predicted values is not required. Conclusions: Species amplitudes given by a minimum and maximum boundary for each species can be used to efficiently predict environmental variables from species composition. The predictive success is superior to mean Ellenberg indicator values and comparable to mean indicator values based on species weighted averages.     

黑龙江兴凯湖沼泽地苔藓植物群落初步研究

2009年夏在黑龙江兴凯湖沼泽地中随机选取47个1m×1m样方,进行苔藓植物及相应维管植物群落调查,并测量记录环境因子,以分析苔藓植物多样性,探讨影响苔藓植物生长与分布的主要环境因子。结果表明：所调查的沼泽地中共有苔藓植物5科、6属、7种,其中粗叶泥碳藓（Sphagnumsquarrosum）和中位泥碳藓（S.magellanicum）为优势种。CCA分析结果显示,粗叶泥炭藓和中位泥炭藓多分布于水位埋深大的生境中,且前者常出现于以细叶苔草（Carexrigescens）为主的群落中,而后者则主要出现于以芦苇（Phragmitesaustralis）-小叶章（Deyeuxiaangustifolia）为主的群落中;中位泥炭藓能生长于沼泽地pH4·4~6·5的所有生境中,粗叶泥碳藓则局限生长于pH5·3左右的生境中,而大湿原藓（Calliergonellacuspidata）、万年藓（Climaciumdendroides）、地钱（Marchantiapolymorpha）和镰刀藓（Drepanocladusaduncus）等4种苔藓植物生长于水体pH值为6·3左右的生境中。研究认为,沼泽地的水位埋深、水pH值和维管植物群落特征是影响这7种苔藓植物生长和分布的主要因子。     

Effects of Nitrogen Deposition: Results of a Temporal-Spatial Analysis of Deciduous Forests in South Sweden

Abstract: We studied permanent plots in deciduous forests in two provinces of South Sweden, Skåne (127 plots) and Småland (116 plots). Vegetation data were sampled in 1983 and 1993 and used to calculate weighted averages of Ellenberg indicator values for light, reaction (pH) and nitrogen. Soil samples were collected from all plots in 1993 for determination of pH, organic matter content and potential net nitrogen mineralization (minN). Data on minN were also available for 66 plots in Skåne from 1983. Nitrogen deposition is considerably higher in Skåne than in Småland. The same holds true for total minN, minNH₄⁺, minNO₃^-_-, and the proportion of total minN due to nitrate (nitrification ratio). According to the comparison of average Ellenberg indicator values between 1983 and 1993, the stands had become darker (not in the heavily managed plots), more acid and nitrogen rich, except at the most acid sites where only minor changes could be observed. The nitrification ratio was significantly higher in 1993 than in 1983. Mean soil nitrification ratios were determined for the most common species in both regions. The means were higher in Skåne than in Småland. It is likely that nitrogen deposition causes an increase in nitrification rate and nitrification ratio. This ratio is a significant determinant of the species composition in deciduous forests.