Relationships between the vegetation and soil conditions in beech and beech-oak forests of northern Germany

In this study we examine the relationships between the vegetation of beech and beech-oak forest communities (Hordelymo-Fagetum, Galio-Fagetum, Deschampsio-Fagetum, Betulo-Quercetum) and their soil conditions in the lowlands of northern Germany, based on 84 sample plots. In all plots the vegetation was recorded and soil parameters were analysed (thickness of the O- and the A-horizons, pH, S-value, base saturation, C/N, mean Ellenberg moisture indicator value). The vegetation classification according to the traditional Braun-Blanquet approach was compared with the result of a multivariate cluster analysis. Vegetation-site relationships were analysed by means of an indirect gradient analysis (DCA).Both traditional classification methods and the cluster analysis have produced comparable classification results. So far as the species composition is concerned, a similar grouping of sample plots was found in both approaches. Multivariate cluster analysis thus supports the classification found by the Braun-Blanquet method. The result of the DCA shows that the four forest communities mentioned above represent clearly definable ecological units. The main site factor influencing changes in the species composition is a base gradient, which is best expressed by the S-value. In addition, within the series Hordelymo-Fagetum - Galio-Fagetum - Deschampsio-Fagetum the C/N-ratios and the thickness of the organic layers (O-horizon) increase continuously. By contrast, the floristic differences between oligotrophic forest communities (i.e., Deschampsio-Fagetum and Betulo-Quercetum) cannot be explained by a base gradient and increasing C/N-ratios. It is suggested that a different forest management history in some cases (e.g., promotion of Quercus robur by silvicultural treatments) is responsible for differences in the species composition, but on the other hand the result of the DCA indicates that Fagus sylvatica is replaced by Quercus robur with increasing soil moisture (i.e., with the increasing influence of a high groundwater table). Summarizing these results, it can be concluded that the ecological importance of single site factors affecting the species composition changes within the entire site spectrum covered by the beech and beech-oak forests of northern Germany.     

Influence of forest development phase on centipede diversity in managed beech forests in Slovenia

Biodiversity in managed forests depends on management practices that determine the forest structure. The influence of stand structure on centipede diversity was investigated by comparing centipede communities from different forest development phases. Using the quadrat counts method, soil samples were taken thrice during 2000 (in spring, summer and autumn) from a deforested area and three evenly aged beech stands (juvenile, pole and timber phases) in managed beech forests in Slovenia. The characteristics of centipede communities and the similarities among them were estimated. The collection comprised 1150 centipedes representing 32 species. At a single site, 10–24 species were found. The average density was between 22 and 808 individuals per m². Differences among communities from different sites were found. The highest species number and abundance were found in the juvenile phase and the lowest in the deforested area. The occurrence of species differed among phases, but none of the species found with a high incidence in one development phase was restricted to that phase. The composition of the centipede community was more sensitive to the successional status of the forest than to the season. A comparison with differently structured forest was made, which suggests that homogenisation leads to reduced centipede diversity. To preserve biodiversity and healthy forest functioning, the maintenance of heterogeneity within a forest stand should be an important focus of management operations.     

Dynamic floristic changes of Swedish beech forest in relation to soil acidity and stand management

This study reports dynamic changes in the beech forest vegetation during one decade, using 95 permanent observation areas representing a wide variety of soils and management regimes. Current soil acidification, including decreasing pH and base cation pools, increasing solubility of toxic elements and increasing deposition of N, as well as recent changes in the beech forest management have created good conditions for the study. Most species of vascular plants increased their frequencies during the 1980's. However, there were several notable exceptions, in particularGalium odoratum, Viola riviniana/reichenbachiana, Polygonatum multiflorum, andMercurialis perennis. These species, demanding a comparatively low soil acidity for survival, are distinctly disfavoured by the long-term soil changes in the forests, which seem to have approached or exceeded their limits of existence in many sites. With most other species, differences in management regimes between the beginning and the end of the observation period were more important to the frequency changes. Sensitive to heavy thinning of the stands were, e.g.Oxalis acetosella, Lamium galeobdolon andMelica uniflora, favoured by thinning wereStellaria nemorum, Carex pilulifera, Milium effusum and an appreciable number of more ephemeral species normally occurring in clear-cut areas or otherwise open land, e.g.,Rubus idaeus, Galeopsis tetrahit, Athyrium filix-femina, Juncus effusus, Agrostic capillaris, Veronica officinalis, Urtica dioica, andMoehringia trinervia. Saplings of woody plants usually also became more frequent during the 1980's.     

Palynological investigations in sediments of ancient lake Duvensee,Schleswig-Holstein (North Germany)

The Duvensee originated before the Alleroed in the Late Glacial, and had its largest areal coverage during the Preboreal. After the lake retreat, which began in early Boreal times, the marginal shore areas and nearshore islands were repeatedly inhabited by man in the early Mesolithicum. Archaeological excavations of human settlements and pollen analyses of sediment cores show evidence of lake level fluctuations in the ensuing period. The results disclose that shallow water sediments such as lake marls, algal muds and coarse detrital gyttjas predominate in the sequence. In keeping with the shallow water conditions, strong lateral facies changes were observed in the cores. The early Holocene deposits have almost the same thickness irrespective of their position in the shallow or deep parts of the lake basin. The subaerial exposure of the nearshore and island areas sometimes resulted in fern and reed peats. The last remnants of the lake, which was drained in 1850, lay in the marginal areas over shallow water sediments.     

Effects of forest management on soil N cycling in beech forests stocking on calcareous soils

The effects of forest management (thinning) on gross and net N conversion, the balance of inorganic N production and consumption, inorganic N concentrations and on soil microbial biomass in the Ah layer were studied in situ during eight intensive field measuring campaigns in the years 2002–2004 at three beech (Fagus sylvatica L.) forest sites. At all sites adjacent thinning plots (“T”) and untreated control plots (“C”) were established. Since the sites are characterized either by cool-moist microclimate (NE site and NW site) or by warm-dry microclimate (SW site) and thinning took place in the year 1999 at the NE and SW sites and in the year 2003 at the NW site the experimental design allowed to evaluate (1) short-term effects (years 1–2) of thinning at the NW site and (2) medium-term effects (years 4–6) of thinning under different microclimate at the SW and NE site. Microbial biomass N was consistently higher at the thinning plots of all sites during most of the field campaigns and was overall significantly higher at the SWT and NWT plots as compared to the corresponding untreated control plots. The size of the microbial biomass N pool was found to correlate positively with both gross ammonification and gross nitrification as well as with extractable soil NO₃⁻ concentrations. At the SW site neither gross ammonification, gross nitrification, gross ammonium (NH₄⁺) immobilization and gross nitrate (NO₃⁻) immobilization nor net ammonification, net nitrification and extractable NH₄⁺ and NO₃⁻ contents were significantly different between control and thinning plot. At the NET plot lower gross ammonification and gross NH₄⁺ immobilization in conjunction with constant nitrification rates coincided with higher net nitrification and significantly higher extractable NO₃⁻ concentrations. Thus, the medium-term effects of thinning varied with different microclimate. The most striking thinning effects were found at the newly thinned NW site, where gross ammonification and gross NH₄⁺ immobilization were dramatically higher immediately after thinning. However, they subsequently tended to decrease in favor of gross nitrification, which was significantly higher at the NWT plot as compared to␣the␣NWC plot during all field campaigns after␣thinning except for April 2004. This increase␣in␣gross nitrification at the NWT plot (1.73 mg N kg⁻¹ sdw day⁻¹ versus 0.48 mg N kg⁻¹ sdw day⁻¹ at the NWC plot) coincided with significantly higher extractable NO₃⁻ concentrations (4.59 mg N kg⁻¹ sdw at the NWT plot versus 0.96 mg N kg⁻¹ sdw at the NWC plot). Pronounced differences in relative N retention (the ratio of gross NH₄⁺ immobilization + gross NO₃⁻ immobilization to gross ammonification + gross nitrification) were found across the six research plots investigated and could be positively correlated to the soil C/N ratio (R = 0.94; p = 0.005). In sum, the results obtained in this study show that (1) thinning can lead to a shift in the balance of microbial inorganic N production and consumption causing a clear decrease in the N retention capacity in the monitored forest soils especially in the first two years after thinning, (2)␣the resistance of the investigated forest ecosystems to disturbances of N cycling by thinning may vary with different soil C contents and C/N ratios, e. g. caused by differences in microclimate, (3) thinning effects tend to decline with the growth of understorey vegetation in the years 4–6 after thinning.     

Three-dimensional structure of a mixed broad-leaved forest in Japan

An investigation was made in a mixed broad-leaved forest in Japan to determine the three-dimensional structure of the crowns of the component species. Vertical extensions and positions of foliage above a circle with a 20-cm diamter on the forest floor were measured using a grid of 177 points in the forest. The first leaf layer above a point was defined to be the highest aggregation of the foliage above the point, and the second leaf layer to be the one below. There were generally two leaf layers above each point, the first and the second. Each of the nineteen species in the plot belonged to either the first leaf layer or the second. Because the leaf layers varied in height from place to place in the forest, the two leaf layers were not separated clearly. Mean vertical depth of each leaf layer was about 2 m irrespective of the leaf layers and component species. Since the depths were similar among the species, species mean crown volumes (volumes of the spaces occupied by foliage) per unit land area mainly depended on horizontal extensions of their crowns, or the coverage. That is, species varied in their horizontal rather than vertical crown extensions. There was an upper limit (6 m) to the sum of vertical depths of the leaf layers above a point on the forest floor. On an average, about three species occurred above a point.     

Determinants of woody species richness in Scot pine and beech forests: climate, forest patch size and forest structure

We analysed patterns of woody species richness in Pinus sylvestris and Fagus sylvatica forests in Catalonia (NE Spain) from forestry inventory databank in relation to climate and landscape structure. Both types of forests are found within the same climatic range, although they have been managed following somewhat different goals. Overall, woody species richness significantly increased when conditions get closer to the Mediterranean ones, with milder temperatures. Differences between the two types of forests arose when comparing the relationship between richness and forest patch size. Woody species richness increased in pine forests with patch size, while the opposite trend was observed in beech forests. This pattern is explained by the different behaviour of structural canopy properties, since leaf area index and canopy cover showed a steeper increase with increasing forest patch size in Fagus forests than in Pinus ones. Accordingly, richness decreased with canopy cover in Fagus plots, but not in Pinus ones. We suggest that these differences would be related to management history, which may have enhanced the preservation of beech stands in larger forest landscape units.     

Respiration activity of ectomycorrhizas from Cenococcum geophilum and Lactarius sp. in relation to soil water potential in five beech forests

Forest trees are involved in root symbioses with hundreds of species of ectomycorrhizal fungi which constitute functional guilds able to improve the water and mineral nutrition of host trees. In temperate ecosystems, water shortage is a main factor limiting tree vitality. To assess how soil water conditions affected the physiological state of beech (Fagus silvatica L.) ectomycorrhizal roots, we monitored glucose respiration of two ectomycorrhizal types (Lactarius sp. and Cenococcum geophilum) during two complete growing seasons. Five stands of contrasting soil conditions were chosen in north-eastern France. The top soil horizons were equipped with micropsychrometers for measuring water potential and temperature. Glucose respiration on individual ectomycorrhizas was measured in vitro by trapping [¹⁴C]-CO₂ from radiolabelled glucose. For soil water potential <-0.2 MPa, the potential respiration activity of C. geophilumectomycorrhizas was significantly less altered than that of Lactariussp. ectomycorrhizas, indicating that C. geophilumis more likely than Lactariussp. to maintain the physiological integrity of beech roots facing drought stress.     

Soil microbial community structure in European forests in relation to forest type and atmospheric nitrogen deposition

The objective of the present study was to evaluate the combined effect of vegetation and N deposition on microbial community composition in forest soils. For this, microbial biomass and community structure were assessed by ester linked fatty acid methyl ester (EL-FAME) analyses for 12 European forest sites representing different forest types (coniferous/deciduous) and differing in annual N loads (2?C40 kg?N?ha^?1). Microbial community composition was affected by vegetation as indicated by a higher proportion of the marker for arbuscular mycorrhiza (AM) fungi??16:1 11???in deciduous forest soils (1.2%?C5.7% of total EL-FAMEs) compared to acidic coniferous forest soils (0.5%?C1.6%). The two pine forest sites investigated showed the highest proportion of fungi (up to 28% of total EL-FAMEs) and the lowest proportions of Gram-negative and Gram-positive bacteria of all study sites. Nitrogen deposition rates were highly correlated with the ratios of cyclopropyl fatty acids to their precursors (r?=?0.82; P?<?0.01) and of bacteria to fungi (r?=?0.71; P?<?0.05). The two sites with the highest N deposition (??32.3 kg?N?ha^?1a^?1) were depleted in the marker fatty acids for AM fungi and other fungi. Our findings suggest that vegetation has a pronounced effect on microbial community structure, but this effect is masked by high N inputs (>30 kg?N?ha^?1a^?1).     

Differences in beech (Fagus crenata) regeneration between two types of Japanese beech forest and along a snow gradient

Differences in beech (Fagus crenata) regeneration were quantitatively investigated using power function analysis for the size–class (diameter at breast height, DBH) distribution and juvenile-to-canopy tree (J/C) ratio along a snow gradient throughout Japan. In snowy areas, all species combined, as well as F. crenata alone, showed constant regeneration, with parameter b≈−1.6 for the power function y=ax ^b (x=DBH, y=density), which is related to the DBH–class distribution. The good fit of the data to the function suggests that beech regenerates constantly with self-thinning patch dynamics. Parameter a, which indicates the abundance of small trunks, was high. Furthermore, the mean J/C ratio was ≈8, i.e., each parent beech tree produced eight juveniles. These results suggest that beech regenerates constantly with gap dynamics in snowy beech forests on the Japan Sea side of Japan (snowy). However, the fit of F. crenata was lower and nonsignificant in some forests in less snowy areas, despite the high fit of all species combined. In these areas, the mean of a was low, and b was often near zero for F. crenata regressions. These results suggest that the abundance of beech was low, and self-thinning was not evident because of the initial low abundance. Moreover, the mean J/C ratio was <1.0, suggesting that juvenile density was lower than that of canopy trees. Thus, the regeneration of F. crenata on the Pacific Ocean side of Japan (less snowy) is rather sporadic. Less snowy conditions may promote seed desiccation, predation of beechnuts and seedlings, and water stress. Lower F. crenata density may also reduce predator satiation and wind pollination.     

Variation in the aboveground stand structure and fine-root biomass of Bornean heath (kerangas) forests in relation to altitude and soil nitrogen availability

Key message

Borneo’s tropical heath (kerangas) forest has limited soil nutrient availability, and high variation in aboveground structure and fine-root biomass. This variation depends on altitude and soil nitrogen availability.

Abstract

To elucidate the biotic and abiotic factors affecting the variation in fine-root biomass (FRB, <2 mm diameter) of trees growing under nutrient-poor environments in Sabah, North Borneo, we investigated FRB in different forests with varying soil nitrogen (N) availability. We selected two study sites at different altitudes: the Maliau Basin (ca. 1000 m asl) and Nabawan (ca. 500 m asl). Both sites included tropical heath (kerangas) forest, on infertile soils (podzols) with a surface organic horizon overlying a bleached (eluviated) mineral horizon, and taller forests on more fertile non-podzolic soils. FRB was obtained from each plot by soil coring (to a depth of 15 cm). FRB increased with decreasing soil inorganic N content (NH₄–N and NO₃–N), tree height, and aboveground biomass. Thus, higher proportions of carbon resources were allocated to fine-roots in stands with lower N availability. FRB was significantly greater at the Maliau Basin than at Nabawan, reflecting lower soil N availability at higher altitude. Our results demonstrate high variation in FRB among the heath forests, and suggest that fine-root development is more prominent under a cooler climate where N availability limits tree growth owing to slower decomposition. The variation in N availability under the same climate (i.e., at the same altitude) appears to be related to the extent of soil podzolization.

     

Abundance and trophic structure of pine forest nematodes in relation to soil layers and ground cover

In a 15-20 yr old forest of Scots pine, the total number of nematodes under the dominating cover grass Deschampsia flexuosa , reached 4.0–4.7 × 10⁶ ind. m⁻² stone free soil, corresponding to densities of 8770–9090 and 3500–3790 ind. 100 ml⁻¹ in FH and mineral soil, respectively. Root/fungal feeding nematodes formed 53–59% of the total nematode fauna. The majority in this group was potential mycophages. Obligate root feeders formed only 11–17% of the total number of nematodes. Soil layers and ground cover affected the abundance of trophic groups. Bacterial feeders, potential mycophages and miscellaneous feeders reached their maximal abundance in the FH-layer. Obligate root feeders tended to reach higher densities in the mineral soil. Higher abundance values of most trophic groups were found under forest grass. The species Calamagrostis arundinacea had a striking positive effect on the relative and absolute abundance of obligate root feeders. This group may be most numerous in early stages of forest development.     

Chemical soil conditions in pristineNothofagus forests of New Zealand as compared to German forests

In many German forest soils low base saturation of CEC in deeper soil layers was reported and acidic deposition is seen as the major cause of these findings. To test this hypothesis we sampled 5 New Zealand forest soils from pristine beech (Nothofagus fusca, N. menziesii, N. solandri) sites under climatic and geological conditions comparable to higher elevations in Germany. The soils developed from granite and greywacke. Soil samples were analyzed for pH and the exchangeable cations were extracted with 1M NH₄Cl. The base saturation of all soil profiles was very low, even in deeper layers and was thus similar to the patterns found in many German forest soils. The pH was generally higher in the New Zealand soils as compared to Germany. The reason for the depletion of base cations in deeper soil layers of New Zealand forest soils is most likely the leaching of base cations with HCO₃ ^- resulting from the dissociation of carbonic acid in connection with high amounts of seepage. Thus, under high rainfall conditions, the low base saturation found in deeper layers of forest soils cannot exclusively be attributed to the effects of acidic depositions and land use. ei]Section editor: R F Huettl     

Representative estimates of soil and ecosystem respiration in an old beech forest

Respiration has been proposed to be the main determinant of the carbon balance in European forests and is thus essential for our understanding of the carbon cycle. However, the choice of experimental design strongly affects estimates of annual respiration and of the contribution of soil respiration to total ecosystem respiration. In a detailed study of ecosystem and soil respiration fluxes in an old unmanaged deciduous forest in Central Germany over 3 years (2000–2002), we combined soil chamber and eddy covariance measurements to obtain a comprehensive picture of respiration in this forest. The closed portable chambers offered to investigate spatial variability of soil respiration and its controls while the eddy covariance system offered continuous measurements of ecosystem respiration. Over the year, both fluxes were mainly correlated with temperature. However, when soil moisture sank below 23 vol.% in the upper 6 cm, water limitations also became apparent. The temporal resolution of the eddy covariance system revealed that relatively high respiration rates occurred during budbreak due to increased metabolic activity and after leaf fall because of increased decomposition. Spatial variability in soil respiration rates was large and correlated with fine root biomass (r ² = 0.56) resulting in estimates of annual efflux varying across plots from 730 to 1,258 (mean 898) g C m⁻² year⁻¹. Power function calculations showed that achieving a precision in the soil respiration estimate of 20% of the full population mean at a confidence level of 95%, requires about eight sampling locations. Our results can be used as guidelines to improve the representativeness of soil respiration measurements by nested sampling designs, being applied in long-term and large-scale carbon sequestration projects such as FLUXNET and CarboEurope.     

Variation of soil and biomass carbon pools in beech forests across a precipitation gradient

Temperate forests have recently been identified as being continuing sinks for carbon even in their mature and senescent stages. However, modeling exercises indicate that a warmer and drier climate as predicted for parts of Central Europe may substantially alter the source/sink function of these economically important ecosystems. In a transect study with 14 mature European beech (Fagus sylvatica L.) forests growing on uniform geological substrate, we analyzed the influence of a large reduction of annual precipitation (970–520 mm yr^?1) on the carbon stocks in fast and slow pools, independent of the well‐known aging effect. We investigated the C storage in the organic L, F, H layers, the mineral soil to 100 cm, and in the biomass (stem, leaves, fine roots), and analyzed the dependence of these pools on precipitation. Soil organic carbon decreased by about 25% from stands with > 900 mm yr^?1 to those with < 600 mm yr^?1; while the carbon storage in beech stems slightly increased. Reduced precipitation affected the biomass C pool in particular in the fine root fraction but much less in the leaf biomass and stem fractions. Fine root turnover increased with a precipitation reduction, even though stand fine root biomass and SOC in the organic L, F, and H layers decreased. According to regression analyses, the C storage in the organic layers was mainly controlled by the size of the fine root C pool suggesting an important role of fine root turnover for the C transfer from tree biomass to the SOC pool. We conclude that the long‐term consequence of a substantial precipitation decrease would be a reduction of the mineral soil and organic layer SOC pools, mainly due to higher decomposition rates. This could turn temperate beech forests into significant carbon sources instead of sinks under global warming.     

Prediction of macrofungal occurrence in Swedish beech forests from soil and litter variable models

The spatial distribution of 49 macrofungal species in Swedish beech forests was related to the statistical variation in 31 edaphic variables. In order to reduce the multicollinearity problem, the variables were transformed into eight principal components, PCs, which are used in two-group discriminant analysis (on absence/presence patterns) and multiple regression analysis (on number of fruit-bodies). The results suggested that base saturation and organic matter content are of outstanding importance. However, significant relationships were also found with other variables, i.e. Cd or Zn in soil and litter, soil nitrogen mineralization rate, and Na or S in litter. One interesting interpretation of the results is that fungi do not only respond to the main variables of a gradient (soil pH, organic mater, base saturation) but also to other variables. Attempts were made to interpret the PCs to characterize fungal occurrence from the models they formed.     

Significant and persistent impact of timber harvesting on soil microbial communities in Northern coniferous forests

Forest ecosystems have integral roles in climate stability, biodiversity and economic development. Soil stewardship is essential for sustainable forest management. Organic matter (OM) removal and soil compaction are key disturbances associated with forest harvesting, but their impacts on forest ecosystems are not well understood. Because microbiological processes regulate soil ecology and biogeochemistry, microbial community structure might serve as indicator of forest ecosystem status, revealing changes in nutrient and energy flow patterns before they have irreversible effects on long-term soil productivity. We applied massively parallel pyrosequencing of over 4.6 million ribosomal marker sequences to assess the impact of OM removal and soil compaction on bacterial and fungal communities in a field experiment replicated at six forest sites in British Columbia, Canada. More than a decade after harvesting, diversity and structure of soil bacterial and fungal communities remained significantly altered by harvesting disturbances, with individual taxonomic groups responding differentially to varied levels of the disturbances. Plant symbionts, like ectomycorrhizal fungi, and saprobic taxa, such as ascomycetes and actinomycetes, were among the most sensitive to harvesting disturbances. Given their significant ecological roles in forest development, the fate of these taxa might be critical for sustainability of forest ecosystems. Although abundant bacterial populations were ubiquitous, abundant fungal populations often revealed a patchy distribution, consistent with their higher sensitivity to the examined soil disturbances. These results establish a comprehensive inventory of bacterial and fungal community composition in northern coniferous forests and demonstrate the long-term response of their structure to key disturbances associated with forest harvesting.     

The significance of life history strategies in the developmental history of mixed beech (Nothofagus) forests,New Zealand

Size and age structure analysis, dated past disturbances, treefall replacement patterns, and spatial pattern analysis were used to reconstruct the developmental history of two old-growth Nothofagus fusca/N. menziesii stands, South Island, New Zealand. Diameter and height class distributions suggested that N. menziesii was replacing N. fusca, however, stand history reconstruction analysis showed that both species had regenerated intermittently after small-scale disturbances. Although large-scale disturbances such as blowdowns may occasionally generate even-aged stands, gap-phase regeneration maintains the forests in compositional equilibrium. In the absence of other competing tree species and understorey plants the two species appear to coexist by way of different life history strategies, where one species (N. menziesii) has low juvenile mortality and the other (N. fusca) has faster height growth rates and greater longevity and adult survivorship.     

Secondary coordination of dichlorodioxomolybdenum(VI) to crown ethers

The reactions of MoO₂Cl₂ with 15-crown-5 and 18-crown-6 in ether solution produced crystals of [MoO₂Cl₂(H₂O)₂] · (15-crown-5) and [MoO₂Cl₂(H₂O)₂] · (H₂O)₂ · (18-crown-6) due to adventitious water. Further hydrolysis lead to crystals of [(H₃O) · (crown)]₂[Mo₆O₁₉]. The structures of the three complexes are briefly discussed.     

Difference of pine ectomycorrhizal biomass in relation to forest conditions

In this study, two plots in a secondary and another two in planted Pinus densiflora stands were exposed to different forest treatments, and the ectomycorrhizal (EM) biomass and its ergosterol content was measured for a year. The unmanaged plot in the secondary stand had greater EM biomass than those in any other plots. Whereas understory cutting had less effect on EM biomass, litter and humus removal decreased pine EM biomass and its ergosterol content, suggesting that such forest treatment alters EM biomass and its structure.