Vegetation of beech forests in the Rychlebské Mountains,Czech Republic,re-inspected after 60 years with assessment of environmental changes

From 1941–;1944 nearly 30 phytosociological relevés were completed by F. K. Hartmann in the Rychlebské Mountains, a typical mountainous area in northeastern Czech Republic. Of the original plots still covered with adult grown beech (Fagus sylvatica) forest, 22 were resampled in 1998 and 1999. In order to describe the recent vegetation variability of the sites 57 relevés were recorded. Changes in vegetation were estimated using relative changes in species density and ordinations (PCA, RDA). Environmental changes were assessed using Ellenberg indicator values when no direct measurements were available. A decline in species diversity has been documented, particularly, many species occurring frequently in deciduous forests with nutrient and moisture well-supplied soils around neutral have decreased. In contrast, several light-demanding, acid- and soil desiccation-tolerant species have increased. Natural succession, quantified as forest age, contributed slightly to these changes. In Ellenberg indicator values, a decline in F (soil moisture), R (soil calcium) and N (ecosystem productivity), and an increase in L (understorey light) were shown. This is interpreted as the influence of modified forestry management and of airborne pollutants. Intensified logging caused the canopy to open and soil conditions to worsen. The latter is most likely also due to acid leaching of soil cations (Ca, K, Na). This caused a decline in soil productivity, thus the effect of nitrification could not be detected. The original relevés may have differed in size influencing the results. This revised version was published online in August 2006 with corrections to the Cover Date.     

Some notes on estimation of the basic homoteneity-coefficient of sets of phytosociological relevés

The formula for calculation of the basic homotoneity-coefficient of sets of phytosociological relevés does not give adequate values for sets of 4 or 3 relevés. Therefore an empirical correction procedure is proposed for the calculation of this coefficient for such small sets of relevés. For sets of 2 relevés the calculation of the basic homotoneity-coefficient gives values equal toSørensen’s coefficient of floristic similarity. A simplification of calculation of the basic homotoneity-coefficient for sets of more than 20 relevés is proposed.     

Plant indicator values of a high-phytodiversity country (Italy) and their evidence,exemplified for model areas with climatic gradients in the southern inner Alps

We tested the new Ellenberg–Pignatti indicator values in Festucetalia valesiacae communities of three valley regions (southern inner Alps: Valle d’Aosta, Valtellina, Valle Venosta/Vinschgau). In these landscape types gradients of all value types exist: light (L), temperature (T), moisture (F/M), nitrogen (N), continentality (C) and soil reaction (R). L- and T-values were extended from 9 to 12 in the database of Pignatti due to the higher intensity of these factors in the area in question; in our dataset this is especially relevant for the L-values.In a spatial approach, we compared DCA results with indicator value results (presence–absence) and with climatic data for eight different plant communities. We found comprehensible results in all cases, also for the extension of the L-factor.In a temporal approach, we compared relevés and their indicator values from two time windows: 1930–1950 (Braun-Blanquet) and 1990–1995 (Schwabe and Kratochwil) for six of the eight different plant communities. Both relevé types were made in the same or in similar localities, but no permanent plots existed. Mostly we found in these communities, which are said to have high “stability”, no significant changes on average for any of the indicator values. In the case of N- and R-values there was a trend of increase in the different datasets, and in some cases this trend is significant.In general, the new database has shown the “robustness” of the Ellenberg indicator value model including the extended L-values, and established its usefulness for a synthetic approach in the studied valley systems. In our systems L-, T-, F/M- and C-gradients can be characterised well by means of indicator values and are an excellent tool for explaining DCA results. Caution is necessary especially when correlations of different factors are intermingled in temporal approaches.     

Bayesian classification of vegetation types with Gaussian mixture density fitting to indicator values

Question: Is it possible to mathematically classify relevés into vegetation types on the basis of their average indicator values, including the uncertainty of the classification? Location: The Netherlands. Method: A large relevé database was used to develop a method for predicting vegetation types based on indicator values. First, each relevé was classified into a phytosociological association on the basis of its species composition. Additionally, mean indicator values for moisture, nutrients and acidity were computed for each relevé. Thus, the position of each classified relevé was obtained in a three‐dimensional space of indicator values. Fitting the data to so called Gaussian Mixture Models yielded densities of associations as a function of indicator values. Finally, these density functions were used to predict the Bayesian occurrence probabilities of associations for known indicator values. Validation of predictions was performed by using a randomly chosen half of the database for the calibration of densities and the other half for the validation of predicted associations. Results and Conclusions: With indicator values, most reléves were classified correctly into vegetation types at the association level. This was shown using confusion matrices that relate (1) the number of relevés classified into associations based on species composition to (2) those based on indicator values. Misclassified relevés belonged to ecologically similar associations. The method seems very suitable for predictive vegetation models.     

Experience with using Ellenberg’s <Emphasis Type="Italic">R</Emphasis> indicator values in Slovakia: Oligotrophic and mesotrophic submontane broad-leaved forests

Ellenberg’s indicator values have been suggested as useful method of estimating site conditions using plants. We examined whether Ellenberg’s R values are suitable for indicating soil reaction and if calibration to physical pH measurements can improve bioindication in oligotrophic and mesotrophic submontane broad-leaved forests in Slovakia. Vegetation relevés and pH-H₂O and pH-CaCl₂ soil reaction were recorded for this purpose. Ellenberg’s R values (R _e) were compared to Jurko’s indicator values (R _j) and a set of species R values and tolerances (T), which were calibrated with physical pH data using the weighted averaging (R _w, T _w) and Huisman-Olff-Fresco modelling (R _h, T _h). Original R _e values were then recalibrated with measured pH data to establish new, adjusted set of scores (R _c, T _c) at Ellenberg’s scale. The Re values are significantly correlated with the other R values, and they demonstrate similar frequency distribution to R _j and R _w values for the studied species pool. The frequency distribution becomes similar across all the R values when indifferent species were excluded. The performance of all the indicator values in terms of bioindication was tested. Relevé means of the R values were regressed on the field pH measurements. The performance of bioindication varied from 36% to 49% of the explained variance for pH-CaCl₂, with the R _e and R _c values yielding 46% and 49% respectively. The bioindication slightly improved for all calibrated methods (R _w, R _h and R _c) when species were weighted inversely with their tolerances — the performance varied from 42% to 51%, and the R _c values performed most effectively. We concluded that Ellenberg’s R values represent a powerful system for bioindicating soil acidity when compared to the other alternatives, with pH-CaCl₂ showing better results than pH-H₂O. Recalibration of Ellenberg’s values to the measured data improved the indicator system.     

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.     

Towards unification of national vegetation classifications: A comparison of two methods for analysis of large data sets

Abstract. In European phytosociology, national classifications of corresponding vegetation types show considerable differences even between neighbouring countries. Therefore, the European Vegetation Survey project urgently needs numerical classification methods for large data sets that are able to produce compatible classifications using data sets from different countries. We tested the ability of two methods, TWINSPAN and COCKTAIL, to produce similar classifications of wet meadows (Calthion, incl. Filipendulenion) for Germany (7909 relevés) and the Czech Republic (1287 relevés) in this respect. In TWINSPAN, the indicator ordination option was used for classification of two national data sets, and the extracted assignment criteria (indicator species) were applied crosswise from one to the other national data set. Although the data sets presumably contained similar community types, TWINSPAN revealed almost no correspondence between the groups derived from the proper classification of the national data set and the groups defined by the assignment criteria taken from the other national data set. The reason is probably the difference in structure between the national data sets, which is a typical, but hardly avoidable, feature of any pair of phytosociological data sets. As a result, the first axis of the correspondence analysis, and consequently the first TWINSPAN division, are associated with different environmental gradients; the difference in the first division is transferred and multiplied further down the hierarchy. COCKTAIL is a method which produces relevé groups on the basis of statistically formed species groups. The user determines the starting points for the formation of species groups, and groups already found in one data set can be tested for existence in the other data set. The correspondence between the national classifications produced by COCKTAIL was fairly good. For some relevé groups, the lack of correspondence to groups in the other national data set could be explained by the absence of the corresponding vegetation types in one of the countries, rather than by methodological problems.     

Effects of environmental heterogeneity on species diversity and composition of terrestrial bryophyte assemblages in tropical montane forests of southern Ecuador

Background: Most studies on tropical bryophytes deal with epiphytic species. This is the first ecological study of tropical forests that focuses specifically on terrestrial bryophytes.

Aim: To investigate the differences between slope and ridge environments in upper montane forests of southern Ecuador in terms of species diversity (richness, abundance), species composition and life forms of terrestrial bryophytes.

Methods: We used Non-metric Multidimensional Scaling (NMDS) to group bryophyte relevés by study location, habitat type and exposure class. Species indicator values were calculated and compared for different habitats.

Results: In total, 140 species were recorded, the majority being liverworts. NMDS analyses and Mantel correlations clearly separated between slope and ridge relevés, and between sunny and shaded microhabitats on ridges. Bryophyte life forms also showed different distribution patterns in slope and in ridge habitats. Mosses were more prominent in sunny than in shaded microhabitats.

Conclusions: Environmental differentiation between ridges and slopes, and small-scale variation in microclimatic conditions caused by differences in exposure, were stronger predictors of species richness and composition than geographical distance between study sites.     

Diversity,variability and habitat characteristics of the communities dominated by Sesleria species (Poaceae) in the Western Carpathians

This study provides a view of vegetation types dominated by Sesleria species in the Western Carpathians (Sesleria caerulea, S. heufleriana, S. tatrae, and S. uliginosa). We also took into account characteristics/traits such as plant life forms, autochthonous status, endemism, and ploidy level occurring within each relevé in our data set. Altitude and Ellenberg indicator values derived for each relevé were considered as well. Eight vegetation types/formations/habitats were recognized in a data set of 942 phytosociological relevés: (1) synanthropic vegetation, (2) rock fissures and screes, (3) alpine grasslands and heaths, (4) mesic grasslands and pastures, (5) springs and fens, (6) forests, (7) xerophilous shrubland, and (8) high-mountain scrubs (krummholz). Results corroborated and clearly emphasized that Sesleria caerulea has the widest ecological amplitude of all studied species because the species occurred among all studied vegetation types. Sesleria tatrae was present only in several vegetation types occurring from montane to alpine vegetation belts in the highest mountains. Sesleria heufleriana and S. uliginosa were recorded only in low-altitude areas. The difference between them lies mainly in the dampness of each locality. Sesleria heufleriana was frequently found in xerophilous communities, whereas S. uliginosa preferred humid habitats of springs and fens.     

Picea abies andAbies alba forests of the austrian alps: Numerical classification and ordination

A TWINSPAN classification of a representative set of 3026 relevés of spruce and fir forests from the Eastern Alps (Austria) is presented. Ecological features of relevé clusters and species groups are described by means of Ellenberg indicator values, site factors and stand characteristics. The most important floristic discontinuity in the data set separates acidophilous communities on mostly silicate substrates from basiphilous communities on mostly carbonate substrates. Further divisions reflect a combined gradient of temperature, nutrient regime and shading. This is supported by the correlation of average Ellenberg values of sample plots with DCA axes. A qualitative comparison between TWINSPAN clusters and a syntaxonomic system widely used in the Austrian Alps is drawn. The two ordersPiceetalia excelsae andAthyrio-Piceetalia largely coincide with the clusters of the first level of divisions. Alliances are partly reproduced by TWINSPAN. Clusters on the fifth and fourth level of division mostly correspond to associations. However, a considerable portion of the lower level clusters is of a transitional type. Out of thirteen fir and spruce associations described for the Austrian Alps, five associations are not reproduced by TWINSPAN, i.e.Bazzanio-Piceetum, Veronico-Piceetum, Adenostylo alliariae-Abietetum, Asplenio-Piceetum andCarici-Piceetum. Three associations are split on the second level of division, i.e.Larici-Piceetum, Luzulo nemorosae-Piceetum andCalamagrostio variae-Piceetum.     

Geographical and ecological differentiation in Greek Fagus forest vegetation

Question: Which are the gradients of floristic differentiation in Greek beech (Fagus sylvatica) forests? Which is the role of geographical and ecological factors in this differentiation? Location: Beech forests of the plant geographical regions Northeast, North Central and East Central Greece. Methods: A total of 1404 published and unpublished phytoso‐ciological relevés were used in the analyses. TWINSPAN and DCA were applied to classify and ordinate the relevés. Altitude, Indicator Values of relevés and their X and Y coordinates were used in a posteriori interpretation of the ordination axes. Kendall's correlation coefficients were calculated between DCA relevé scores and explanatory variables. Multiple linear regression was used to partition the variation explained by the first two DCA axes, between the geographical and the ecological variables. Results: Classification resulted in 14 vegetation units defined by species composition. Two types of gradients, ecological and geographical, were revealed by the DCA of all releves. The partition of the variation accounted for by the first and second DCA axis was attributed mainly to ecological and geographical variables, respectively. Conclusions: Beech forests of northeast and Central Greece show phytogeographical differences, while ecologically similar vegetation units occur in both regions. A west‐east gradient is revealed in Greek beech forest vegetation. The extent of the study area, its position along regional gradients and the comprehensiveness of the data set that is analysed determine the types of the gradients which can be revealed in a vegetation study.     

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.     

Trends in species diversity and composition of urban vegetation over three decades

Question: What was the change in diversity of urban synantropic vegetation in a medium‐sized Central European city during the period of increasing urbanization (1960s‐1990s)? Location: The city of Plzeň, an industrial centre of the western part of the Czech Republic. Methods: Sampling of various types of synanthropic vegetation, conducted in the 1960s, was repeated by using the same methods in the 1990s. This yielded 959 relevés, of which 623 were made in the 1960s and 336 in the 1990s. The relevés were assigned to the following phytosociological classes: Chenopodietea, Artemisietea vulgaris, Galio‐Urticetea, Agropyretea repentis and Plantaginetea majoris. Total number of vascular plant species, evenness index J, number of alien species (classified into archaeophytes and neophytes), and mean Ellenberg indicator values for light, temperature, continentality, moisture, soil reaction, and nutrients were obtained for each relevé. Results: From 1960s to 1990s, there was a significant decrease of species richness and diversity in synanthropic vegetation. The proportion of archaeophytes decreased in most vegetation types, indicating the contribution of this group of species, often confined to specific rural‐like habitats, to the observed impoverishment of ruderal vegetation. The proportion of neophytes did not change between the two periods. Comparison between 1960s and 1990s indicated a decrease in light, temperature, moisture, soil reaction and nutrient indicator values in some vegetation types. In both periods, Artemisieta, Galio‐Urticetea and Chenopodietea formed a distinct group harbouring more species than Agropyretea and Plantaginetea. Neophytes, i.e. recently introduced species, were most represented in the early successional annual vegetation of Chenopodietea, rather than in perennial vegetation of the other classes. Conclusions: Synanthropic vegetation of Plzeň exhibited a general trend of decrease in species diversity.     

Indicator species of broad-leaved oak forests in the eastern Iberian Peninsula

The degraded state, scattered distribution and substrate type of Quercus pyrenaica forests in eastern Spain make them a subject of particular interest, as well as a priority for conservation efforts. An extension of Dufrêne and Legendre's indicator value (IndVal) analysis was applied to one hundred relevés. This extension of the IndVal method allowed us, by considering the preference of species for combinations of relevé groups, to obtain those species that represent local, but also regional, characteristics (as a new practical use). Our objective was to identify the species that best indicate the main ecological features of these broad-leaved oak forests. The usefulness of our analysis was evidenced at two levels: local indicator species, showing the characteristic species of individual sites and including the presence of endemics in the area; and indicator species of combinations of two or more sites, reflecting the affinities or relationships between these scattered forests.Most singular forests showed a higher number of local indicator species. The indicator species of each site combination represent the main ecological traits shared by these groups – such as proximity to the coast, land uses or postglacial migration pathways – thereby defining their relationships. Indicator value analysis with site group combinations (Multipatt function) proved to be a useful tool for the identification of different species that could serve both to determine the local forest conservation status and aid in its preservation, as well as contributing to an understanding of the life history of larger territories.     

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.     

Vegetational changes in a wet meadow complex,South Bohemia,Czech Republic

Directional changes of vegetation in a wet meadow complex are described, comparing phytosociological relevés taken in 1956, 1963, 1984 and 1989, and vegetation maps made in 1956 and 1984. The relevés were located at various distances from the pond shore predominantly reflecting differences in water regime and intensity of farming. The data were elaborated using direct and indirect gradient analyses. Alterations of environmental factors (moisture, nitrogen) were expressed using Ellenberg's indicator values. Over most of the area, the former distinct vegetation mosaic has been replaced by uniform meadows due to intensive farming. The area was eutrophicated and partly drained, clearly shown in Ellenberg's indicator values, in species composition and vegetation pattern. Rapid processes of ruderalization took place. A scheme of successional pathways which finally emerged can be applied to analogous wet meadows in comparable geographical regions and used as a tool for prognoses of successional processes in meadows under human impact. The paper demonstrates ways to exploit earlier phytosociological records.     

A simple method for estimating homotoneity of sets of phytosociological relevés

The homogeneity of sets of phytosociological relevés or syntaxa—the homotoneity—is tested by means of a homotoneity-coefficient. This coefficient consists of the basic homotoneity-coefficient expressing the proportion of highly constant species (61–100%) in the species composition of the average relevé and the correcting factor which is proportional to the difference of species number between the richest and poorest relevé. Successive analysis of heterotoneity of sets of relevés for detecting sources of heterotoneity is proposed.     

Variability of the <Emphasis Type="Italic">Molinion</Emphasis> meadows in Slovakia

Nine vegetation types were distinguished using cluster analysis within Molinion meadows in Slovakia. Vegetation of cluster 1 occurs on most acidic soils and is characterized by the occurrence of species of the Caricion fuscae alliance and of the Nardus grasslands. Vegetation of cluster 2 is also found on rather acidic soils but in contrast to cluster 1 vegetation it contains species of base-rich sites, such as Betonica officinalis, Galium boreale or Serratula tinctoria. Vegetation of cluster 3 occurs in wet base-rich habitats and often contains species of the Caricion davallianae alliance. Species of dry and Nardus grasslands are typical for vegetation of cluster 4, which is found at the driest sites and is confined to oligotrophic sandy soils. Vegetation of clusters 5 and 6 occurs on moist mesotrophic soils. Their species composition is quite similar, the main difference being that the former includes species-poor relevés and the latter includes species-rich relevés. Relevés of cluster 7 include species of dry grasslands and some ruderal species and represent degraded types of inundated floodplain meadows of the Deschampsion alliance. Vegetation of clusters 8 is characterized by species of the Phragmito-Magnocaricetea class and of the Deschampsion alliance, and occurs in wet nutrient-rich habitats. Vegetation of cluster 9, which usually develops from vegetation of cluster 8 due to decrease in the ground-water table, often contains species of dry grasslands and mesic meadows. Except for relevés of clusters 1 and 7, all others can be assigned to the Molinietum caeruleae Koch 1926 association. Cluster 1 corresponds to the Junco effusi-Molinietum caeruleae Tüxen 1954 association. Average Ellenberg indicator values for relevés, which were passively projected on the ordination biplot of detrended correspondence analysis, showed that the first ordination axis correlates with nutrients, soil base status and temperature, and second axis with moisture.     

The influence of earthworms (Lumbricidae) on the nitrogen dynamics in the soil litter system of a deciduous forest

Summary The influence of earthworms (Aporrectodea caliginosa (Savigny) and Lumbricus castaneus (Savigny)) on the rate of nitrogen net mineralization of the soil was studied in the laboratory and in the field. The additional mineralization of nitrogen cause by the burrowing activity of the substrat feeding earthworm A. caliginosa (N _L )was directly correlated to the biomass of the lumbricids independently of their number. A rise in temperature caused an exponential increase in N _L values. The Q ₁₀ value of this process (2.18) was found to be much higher than that of the nitrogen mineralization without earthworms (Q ₁₀=1.22). At 15°C the N _L value caused by A. caliginosa was calculated to be about 250 g N g^-1 fresh body wt d^-1. Using the experimentally determined exponential relationship between temperature and N _L values, the additional nitrogen mineralization caused by a population of A. caliginosa in a beechwood on limestone was calculated to be 4.23 kg ha^-1 a^-1.In contrast to A. caliginosa the litter dwelling species L. castaneus lost considerable amounts of biomass (56%) during the 4 week incubation period. Only 1/3 of the nitrogen equivalent to the weight loss of the animals was recovered in the mineral nitrogen pool.The addition of litter (old beech leaf litter, freshly fallen beech and ash leaf litter) had a pronounced effect on both nitrogen net mineralization and N _L values of the soil. Presence of old beech leaves caused an increase in both values, wheres the other litter types effected a decrease in nitrogen net mineralization. Fragmented ash litter was found to have the most distinct effect on N _L values (-69%) and nitrogen net mineralization (-74%).     

常绿阔叶林8种乔灌木叶片氮含量及其分配与光合能力的关系

以浙江天童国家森林公园常绿阔叶林为研究对象,采用空间代替时间的方法,研究了5个不同演替阶段常见的4种乔木以及4种灌木叶片的光饱和速率(Pmax)、光合氮素利用效率(PNUE)及其与叶片氮含量(NL)、叶片氮素在细胞壁的分配比例(细胞壁N/叶片总N,NCW/NL)、氮素在光合酶中的分配比例(NR/NL)、单位面积叶干重(LMA)的相互关系。结果表明:(1)演替系列4种乔木和4种灌木各种间指标除NL外均表现出显著差异,前期种较后期种具有更高的NR/NL、PNUE、Pmax,而后期种LMA、NCW/NL、MCW/ML(细胞壁干重/叶片总干重)更大,NL在乔木各种间差异不明显,在灌木种间则差异显著;乔木种较灌木种具有更大的LMA、NCW/NL、MCW/ML,而NR/NL则较灌木小;8种植物的Pmax与NL以杨梅为最高,连蕊茶最低;苦槠具有最高的PNUE,而栲树最低。(2)随着演替的进行,前期种的NR/NL、PNUE、Pmax有减小趋势,而LMA、NCW/NL、MCW/ML逐渐增大,后期种则表现出相反的趋势。(3)NR/NL与Pmax、PNUE之间呈显著正相关关系,而LMA、NCW/NL、MCW/ML则与Pmax、PNUE、NR/NL显著负相关。研究认为,NR/NL与NCW/NL之间的负相关性及其对PNUE的影响可以在一定程度上解释树木在光合与维持两方面的权衡关系以及演替的生理机制。