Changes in diversity and storage function of ectomycorrhiza and soil organoprofile dynamics after introduction of beech into Scots pine forests

Diversity and storage function of mycorrhiza as well as soil organoprofile formation were investigated in a chronosequence of a pure Scots pine (Pinus sylvestris L.) stand, of Scots pine stands that were underplanted with beech (Fagus sylvatica L.) and in three pure beech stands of different age. Mycorrhiza diversity was higher in the pure beech stands compared to the pure pine stand. Beech and pine trees in the mixed stands had similar dominant mycorrhiza morphotypes. However, trees in two of the three pure beech stands were mycorrhized with other types. Mycorrhizal abundance and nutrient amounts of mycorrhizae associated with beech trees were higher in the mixed and in the pure beech stands compared to pine mycorrhizae indicating that nutrient uptake was higher in older beech than in older pine trees. Humus quality varied from pine to beech stands. Plant litter storage in the humus layer was highest in the youngest mixed stand and lowest in the oldest beech stand. Humus forms changed from moder grass-type in the pure Scots pine stand to mor-like moder and moder rich in fine humus with increasing age of beeches in the mixed stands. The older beech stands were characterised by oligomull and mull-like moder as the dominating humus forms. The ecologically favourable humus forms, i.e., nutrient rich humus forms in the older beech stands correlate well with the higher mycorrhizal diversity and abundance as well as the higher nutrient storage of their mycorrhizae in these stands. The results are also discussed with regard to the 'base-pump effect' of beech trees.     

Restoration of Aboveground Ectomycorrhizal Flora in Stands of Pinus sylvestris (Scots Pine) in The Netherlands by Removal of Litter and Humus

Species richness and sporocarp density of ectomycorrhizal fungi in stands of Pinus sylvestris (Scots pine) in The Netherlands have decreased during the last decades. The lowest species diversity was found in P. sylvestris stands situated in areas with high atmospheric deposition of nitrogen originating from intensive livestock industry. In these stands, litter and humus have accumulated into thick layers, and the herbaceous understory vegetation is dominated by the grass Deschampsia flexuosa (wavy hair grass). Earlier investigations showed negative correlations between the number of species of ectomycorrhizal fungi above ground and the depth of humus layers. Our aim was to investigate whether removal of litter, humus layers, and herbaceous vegetation (sods)—so-called "sod cutting"—increased species diversity above ground and sporocarp density of ectomycorrhizal fungi in P. sylvestris stands of different age. Therefore, three P. sylvestris stands of different ages (planted in 1987, 1963, and 1924) on Haplic Arenosol were selected. In 1990, litter, humus layers, and herbaceous vegetation were removed to create nutrient-poor sandy soils without overlying litter and humus layers. Untreated plots served as controls. Surveys conducted in 1991, 1992, and 1993 indicated that sod cutting enhanced the species diversity and sporocarp density of ectomycorrhizal fungi. These results suggest that sod cutting is a way to restore ectomycorrhizal flora in medium-aged and old stands of P. sylvestris where litter and humus have accumulated.     

Ectomycorrhizal root growth in scots pine stands in response to manipulation of litter and humus layers

 The effect on ectomycorrhizal root growth in a nitrogen-enriched planted stand of Scots pine (Pinus sylvestris L.) on podzolic sandy soil to manipulation of litter and humus layers (removal, doubling and control treatments) was examined, and compared to ectomycorrhizal root growth in an untreated naturally established Scots pine stand on nutrient-poor non-podzolic sandy soil. Half a year after manipulation of litter and humus layers in the planted stand, ingrowth-cores to a depth of 60 cm were installed in both stands. Scots pine roots were sampled four times during two growing seasons. Ectomycorrhizal roots were found at all sampled soil depths to 60 cm in all plots. Root growth and ectomycorrhizal development were greater in the naturally established stand than in all plots in the planted stand. Numbers of ectomycorrhizal root tips in the litter and humus removal plots were generally higher than in the control plots in the planted stand until May 1992. Doubling litter and humus did not significantly affect root length or the numbers of ectomycorrhizal root tips. The N _dissolved , NH₄ ⁺ and NO₃ ^– concentrations and the organic matter content in the upper 5 cm of the mineral soil in the planted stand on podzolic sandy soil were generally higher and the pH significantly lower than in the naturally established stand on non-podzolic sandy soil. Root growth and ectomycorrhizal development in the secondary stand may have been negatively affected by the chemical composition of the podzolic sandy soil. Accepted: 19 March 1997     

Degradability of dissolved soil organic carbon and nitrogen in relation to tree species

The degradability and chemical characteristics of water-extractable dissolved organic carbon (DOC) and nitrogen (DON) from the humus layer of silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) stands were compared in short-term incubation of soil solutions. For all extracts the degradation of DOC and DON was low (12-17% loss) and increased in the order: birch, spruce and pine. In the humus layer under pine a relatively larger pool of rapidly degrading dissolved soil organic matter (DOM) was indicated by the [3H]thymidine incorporation technique, which measures the availability of DOM to bacteria. The degradation of DOC was explained by a decrease in the hydrophilic fraction. For DON, however, both the hydrophilic and hydrophobic fractions tended to decrease during incubation. No major differences in concentrations of hydrophilic and hydrophobic fractions were detected between tree species. Molecular size distribution of DOC and DON, however, revealed slight initial differences between birch and conifers as well as a change in birch extract during incubation. The depletion of very rapidly degrading fractions (e.g., root exudates and compounds from the litter) may explain the low degradability of DOM in the humus layer under birch.     

Ectomycorrhizal development on Scots pine (Pinus sylvestris L.) seedlings in different soils

In the present study ectomycorrhizal development of Laccaria bicolor, Rhizopogon luteolus and Suillus bovinus associated with Scots pine (Pinus sylvestris) seedings was studied as affected by primary stand humus, secondary stand humus, podsolic sandy soil or peat in perspex growth chambers. After 9 weeks, ectomycorrhizal development with S. bovinus was significantly greater in peat and primary stand humus than in secondary stand humus or podsolic sandy soil. Ectomycorrhizal development with R. luteolus in secondary stand humus was higher than in primary stand humus. Degree of ectomycorrhizal development of L. bicolor, R. lutuelus and S. bovinus on Scots pine was related to potassium concentration, organic matter content and pH of the soils suggesting that chemical composition of the soils affects ectomycorrhizal development.     

Why does land-use history facilitate non-native plant invasion? A field experiment with Celastrus orbiculatus in the southern Appalachians

Although historic land use is often implicated in non-native plant invasion of forests, little is known about how land-use legacies might actually facilitate invasion. We conducted a 2-year field seeding experiment in western North Carolina, USA, to compare germination and first-year seedling survival of Celastrus orbiculatus Thunb. in stands that had been cultivated and abandoned a century earlier and were dominated by tulip poplar (Liriodendron tulipifera L.), and in paired stands that had never been cultivated and were dominated by oaks (Quercus spp.). Experiments were conducted at five sites with paired tulip poplar and oak stands by varying litter mass (none, low, or high) and litter type (tulip poplar or oak). We also performed reciprocal soil translocations using pots seeded with C. orbiculatus. Soil moisture and temperature were measured throughout the growing season. Germination and survival were highest in the tulip poplar stands. Germination was also higher in plots with low litter mass. Seedling survival was highest in plots with low litter mass or no litter. Soil moisture was higher in tulip poplar stands and under low-mass litter. Differences in germination and survival among the potted plants were minimal, suggesting that soil type and ambient site conditions were less important than litter conditions for C. orbiculatus establishment. Our results suggest that the low litter mass and mesic soil conditions that are characteristic of tulip poplar stands may confer higher invasibility and explain the higher abundance of C. orbiculatus in areas with successional overstory communities associated with historically cultivated forests.     

Post-logging organic matter recovery in forest ecosystems of eastern Baikal region

The dynamics of organic matter accumulated in the soil and main vegetation elements was analyzed for post-logging forest ecosystem succession series in eastern Baikal region. The phytomass was found to allocate up 63 and 50% of carbon in undisturbed Scots pine and fir stands, respectively. The post-logging phytomass contribution to the total carbon pool appeared to decrease down to 16% in Scots pine and 6% in fir stands. In Scots pine stands, carbon storage was determined to account for almost 70% of the initial carbon 60 years after logging. In 50- to 55-year-old fir stands, carbon recovered its initial pool only by 10%. Soil carbon recorded in recently logged Scots pine and fir sites appeared to be 5 and 16 times that accumulated in the phytomass, respectively. The ratio between phytomass carbon and soil organic matter recovered back to the prelogging level in Scots pine stands by the age of 50–60 years. While phytomass carbon also increased in fir stand of the same age, it did not reach the level of the control stand.     

Tree Species Traits Influence Soil Physical,Chemical, and Biological Properties in High Elevation Forests

Background

Previous studies have shown that plants often have species-specific effects on soil properties. In high elevation forests in the Southern Rocky Mountains, North America, areas that are dominated by a single tree species are often adjacent to areas dominated by another tree species. Here, we assessed soil properties beneath adjacent stands of trembling aspen, lodgepole pine, and Engelmann spruce, which are dominant tree species in this region and are distributed widely in North America. We hypothesized that soil properties would differ among stands dominated by different tree species and expected that aspen stands would have higher soil temperatures due to their open structure, which, combined with higher quality litter, would result in increased soil respiration rates, nitrogen availability, and microbial biomass, and differences in soil faunal community composition.

Methodology/Principal Findings

We assessed soil physical, chemical, and biological properties at four sites where stands of aspen, pine, and spruce occurred in close proximity to one-another in the San Juan Mountains, Colorado. Leaf litter quality differed among the tree species, with the highest nitrogen (N) concentration and lowest lignin∶N in aspen litter. Nitrogen concentration was similar in pine and spruce litter, but lignin∶N was highest in pine litter. Soil temperature and moisture were highest in aspen stands, which, in combination with higher litter quality, probably contributed to faster soil respiration rates from stands of aspen. Soil carbon and N content, ammonium concentration, and microbial biomass did not differ among tree species, but nitrate concentration was highest in aspen soil and lowest in spruce soil. In addition, soil fungal, bacterial, and nematode community composition and rotifer, collembolan, and mesostigmatid mite abundance differed among the tree species, while the total abundance of nematodes, tardigrades, oribatid mites, and prostigmatid mites did not.

Conclusions/Significance

Although some soil characteristics were unaffected by tree species identity, our results clearly demonstrate that these dominant tree species are associated with soils that differ in several physical, chemical, and biotic properties. Ongoing environmental changes in this region, e.g. changes in fire regime, frequency of insect outbreaks, changes in precipitation patterns and snowpack, and land-use change, may alter the relative abundance of these tree species over coming decades, which in turn will likely alter the soils.     

Effects of manipulation of litter and humus layers on ectomycorrhizal colonization potential in Scots pine stands of different age

Effects of manipulation of litter and humus layers (removal, doubling and control treatments) on the colonization potential of ectomycorrhizal fungi were studied in two secondary stands of Pinus sylvestris (5 and 18 years old) in The Netherlands. Five-mont-hold, sterile-grown Scots pine seedlings, inoculated with Laccaria bicolor, Paxillus involutus or Rhizopogon luteolus and noninoculated seedlings were used as baits. The seedlings were harvested after one growing season. For comparison, sporocarps of ectomycorrhizal fungi were also investigated. Genus composition on the seedlings was independent of initial inoculum, but determined by both treatment and age of the stands. In both stands, removal of litter and humus layers increased, and addition of organic material decreased the number of ectomycorrhizal types on the seedlings. Not all indigenous genera were observed by either outplanting seedlings or sporocarp surveys.     

Thiosulfate sulfurtransferase (rhodanese) in forest soils

Concentrations of thiosulfate sulfurtransferase (rhodanese; EC 2.8.1.1) in soils of five types of forest stands (spruce and pine stands without ground vegetation, and mountain ash, birch and pine stands with grass cover) were followed. Soils from grassed stands contained much higher concentrations of the enzyme than soils without ground vegetation. On grassed locations higher enzyme concentrations were found in fermentation horizons, whereas on locations without the ground vegetation they were detected in humus horizons. Urea fertilization and SO₂ pollution did not exhibit any significant effect.     

Sensitivity of French temperate coniferous forests to climate variability and extreme events (Abies alba,Picea abies and Pinus sylvestris)

Questions: (1) How do extreme climatic events and climate variability influence radial growth of conifers (silver fir, Norway spruce, Scots pine)? (2) How do elevation and soil water capacity (SWC) modulate sensitivity to climate? Location: The sampled conifer stands are in France, in western lowland and mountain forests, at elevations from 400 to 1700 m, and an SWC from 50 to 190 mm. Methods: We established stand chronologies for total ring width, earlywood and latewood width for the 33 studied stands (985 trees in total). Responses to climate were analysed using pointer years and bootstrapped response functions. Principal component analysis was applied to pointer years and response function coefficients in order to elucidate the ecological structure of the studied stands. Results: Extreme winter frosts are responsible for greater growth reductions in silver fir than in Norway spruce, especially at the upper elevation, while Scots pine was the least sensitive species. Exceptional spring droughts caused a notable growth decrease, especially when local conditions were dry (altitude<1000 m and SWC<100 mm for silver fir, western lowlands for Scots pine). Earlywood of silver fir depended on previous September and November and current‐year February temperature, after which current June and July water supply influenced latewood. Earlywood of Norway spruce was influenced by previous September temperature, after which current spring and summer droughts influenced both ring components. In Scots pine, earlywood and latewood depended on the current summer water balance. Local conditions mainly modulated latewood formation. Conclusions: If the climate becomes drier, low‐elevation dry stands or trees growing in western lowlands may face problems, as their growth is highly dependent on soil moisture availability.     

Phytoremediation of subarctic soil contaminated with diesel fuel

The effects of several plant species, native to northern latitudes, and different soil amendments, on diesel fuel removal from soil were studied. Plant treatments included Scots Pine (Pinus sylvestris), Poplar (Populus deltoides x Wettsteinii), a grass mixture (Red fescue, Fesuca rubra; Smooth meadowgrass, Poa pratensis and Perennial ryegrass, Lolium perenne) and a legume mixture (White clover, Trifolium repens and Pea, Pisum sativum). Soil amendments included NPK fertiliser, a compost extract and a microbial enrichment culture. Diesel fuel disappeared more rapidly in the legume treatment than in other plant treatments. The presence of poplar and pine enhanced removal of diesel fuel, but removal under grass was similar to that with no vegetation. Soil amendments did not enhance diesel fuel removal significantly. Grass roots accumulated diesel-range compounds. This study showed that utilisation of selected plants accelerates removal of diesel fuel in soil and may serve as a viable, low-cost remedial technology for diesel-contaminated soils in subarctic regions.     

Vegetation Composition Determines Microbial Activities in a Boreal Forest Soil

Abstract Forest soil ecology was studied in Fennoscandinavian dry Scots pine forests grazed by reindeer to varying extents (ungrazed, lichen-dominated-sites; grazed sites; and bryophyte-dominated sites). We hypothesized that the productivity parameters of the site (i.e., tree growth and soil nutrient concentrations), the vegetation composition, and the microbial activities are directly correlated. Since the productivity of the lichen-dominated ecosystem is low, microbial activities are assumed to be naturally low. Grazing was expected to decrease both the amount of Scots pine fine roots and the soil microbial activities. Several variables on the characteristics of the soil microbial community, Scots pine fine roots, soil nutrients, and tree growth were studied in relation to vegetation composition by using non-metric multidimensional scaling (NMDS). Basal respiration (Bas), metabolic quotient of the microbial community (qCO2), and pine fine root parameters increased toward the ungrazed, nutrient-poor, lichen-dominated sites, which were grouped at one end of the first axis in the NMDS ordination. Soil nutrient and tree growth parameters and thickness of the humus layer increased toward bryophyte-dominated sites, which were grouped at the other end of the first axis in the ordination. The grazed sites fell between them. These were characterized by lower Bas and qCO2 values and longer lag, compared to ungrazed lichen- or bryophyte-dominated sites, probably due to decreased carbon input and microclimatic change (the soil without lichen carpet is exposed to direct sunlight and wind). Microbial biomass (Cmic), fungal biomass (ergosterol concentration), and the specific growth rate (μCO2) were not related to vegetation ordination. The high fine root production is the most plausible explanation for the high microbial activities at nutrient-poor, lichen-dominated sites, which produce qualitatively poor and slowly decomposing litter, as fine roots secrete considerable amounts of organic substances. At bryophyte-dominated sites, the higher soil nutrient concentrations and the higher production of easily decomposable substrates are likely to maintain the microbial activities.     

Soil CO2 efflux in a mixed pine-oak forest in Valsaín (central Spain)

Soil-surface CO2 efflux and its spatial and temporal variation were investigated in a southern Mediterranean, mixed pine-oak forest ecosystem on the northern slopes of the Sierra de Guadarrama in Spain from February 2006 to July 2006. Measurements of soil CO2 efflux, soil temperatures, and moisture were conducted in nine 1963-m2 sampling plots distributed in a gradient around the ecotone between Pinus sylvestris L. and Quercus pyrenaica Lam. forest stands. Total soil organic matter, Walkey-Black C, particulate organic matter, organic matter fraction below 53 microm, total soil nitrogen content, total soil organic carbon content, and pH were also measured under three representative mature oak, pine, and mixed pine-oak forest stands. Soil respiration showed a typical seasonal pattern with minimums in winter and summer, and maximums in spring, more pronounced in oak and oak-pine stands. Soil respiration values were highest in pine stands during winter and in oak stands during spring and summer. Soil respiration was highly correlated with soil temperatures in oak and pine-oak stands when soil moisture was above a drought threshold of 15%. Below this threshold value, soil moisture was a good predictor of soil respiration in pine stands. Greater soil organic matter, particulate organic matter, Walkey-Black C, total organic C, and total N content in pine compared to oak sites potentially contributed to the greater total soil CO2 efflux in these stands during the winter. Furthermore, opposing trends in the organic matter fraction below 53 microm and soil respiration between plots suggest that in oak stands, the C forms are less affected by possible changes in use. The effects of soil properties on soil respiration were masked by differences in soil temperature and moisture during the rest of the year. Understanding the spatial and temporal variation even within small geographic areas is essential to assess C budgets at ecosystem level accurately. Thus, this study bears important implications for the study of large-scale ecosystem dynamics, particularly in response to climatic change.     

Greater accumulation of litter in spruce (Picea abies) compared to beech (Fagus sylvatica) stands is not a consequence of the inherent recalcitrance of needles

Background and aims

Replacement of beech by spruce is associated with changes in soil acidity, soil structure and humus form, which are commonly ascribed to the recalcitrance of spruce needles. It is of practical relevance to know how much beech must be admixed to pure spruce stands in order to increase litter decomposition and associated nutrient cycling. We addressed the impact of tree species mixture within forest stands and within litter on mass loss and nutritional release from litter.

Methods

Litter decomposition was measured in three adjacent stands of pure spruce (Picea abies), mixed beech-spruce and pure beech (Fagus sylvatica) on three nutrient-rich sites and three nutrient-poor sites over a three-year period using the litterbag method (single species and mixed species bags).

Results

Mass loss of beech litter was not higher than mass loss of spruce litter. Mass loss and nutrient release were not affected by litter mixing. Litter decay indicated non-additive patterns, since similar remaining masses under pure beech (47%) and mixed beech-spruce (48%) were significantly lower than under pure spruce stands (67%). Release of the main components of the organic substance (C_org, N_tot, P, S, lignin) and associated K were related to mass loss, while release of other nutrients was not related to mass loss.

Conclusions

In contradiction to the widely held assumption of slow decomposition of spruce needles, we conclude that accumulation of litter in spruce stands is not caused by recalcitrance of spruce needles to decay; rather adverse environmental conditions in spruce stands retard decomposition. Mixed beech-spruce stands appear to be as effective as pure beech stands in counteracting these adverse conditions.     

Total, organic and extractable-P in humus and soil beneath Sitka spruce planted in pure stands and in mixture with Scots pine

Total, organic and extractable P were measured in the humus and underlying soil to 10 cm depth beneath Sitka spruce (SS) and mixed Sitka spruce and Scots pine (SS+SP) stands planted on upland heath. The humus beneath SS+SP contained significantly (p<0.01) greater amounts of total and organic-P than that in SS and the mixed stands had more effectively retained approximately 87 per cent of previously applied fertilizer-P, totalling 100 kg P ha^–1, compared with 70 per cent in SS. Despite the larger amounts of total-P in the mixed plots 0.01 M CaCl₂ extractable molybdate reactive phosphorus (MRP) was significantly (p<0.05) greater in SS+SP humus only during March and April. Greater concentrations of MRP were released from the humus and soil during July and August at a mean rate of 58 g P ha^–1 day^–1. This coincided with drying of the soil during the summer and the rate of release, attributed to death of fine roots and microorganisms, was 4 to 30 times greater than reported values for rates of net mineralization of P from forest soils.     

A comparison of pinewood and moorland soils in the Abernethy Forest Reserve, Scotland

Despite considerable published literature on the above-ground ecology of the pinewoods of Scotland, little research has considered the way in which pinewood soils differ from those under other vegetation types. Soil properties were compared between ancient, semi-natural Scots pine forest and moorland on three soil types in the Abernethy Forest Reserve in the Cairngorm Mountains of Scotland. Soil morphology differed considerably between the vegetation types on each soil type, principally in the thickness of organic layers. Forest soils had thicker organic layers and this was particularly true of the F horizon. Forest soils were slightly less acid than equivalent moorland soils and had accumulated significantly more carbon. Forest soils in this environment therefore have the capacity to sequester larger amounts of carbon than moorland, and therefore represent a significant potential carbon sink. The quantity of nitrogen and phosphorus was also consistently larger in the organic layers under pine forest and since little difference existed in these properties in the mineral horizons, it was concluded that this accumulation was real and represented a net addition to the tree-soil system.     

Long term effect of liming and fertilization on ectomycorrhizal colonization and tree growth in old Scots pine (Pinus sylvestris L.) stands

In this study, we surveyed the long term effects of liming and fertilizing in old Scots pine stands on the ectomycorrhiza (ECM) colonization, tree growth and needle nutrient concentration 35 years later. Four mature stands of Scots pine on low productive mineral soil were limed in 1959 and 1964 with total doses of limestone ranging from 3 to 15 Mg ha^?1 and fertilized with nitrogen (N) in 1970. Thirty-five years after the first liming treatment, all stands were analysed for tree growth and needle nutrient concentrations and two of the stands were also analysed for ECM colonization. ECM colonization increased significantly with liming from 61.5% in the control plots to 88% in the plot with the highest limestone dose. ECM colonization increased with increasing pH in the humus layer from 62% colonization at pH?=?3.5 to 90% at pH?=?6.5 and decreased with increasing amount of extractable phosphorus (P) in the humus. Liming did not affect the frequencies of different ECM morphotypes or dead short root tips, the fine root biomass or necromass. ECM colonization was uncorrelated with needle nutrient concentrations or tree increment. Liming did not significantly affect tree growth. However, nutrient concentrations of current-year needles were affected by prior liming. Ca concentrations in current-year needles increased from approximately 15 mg g^?1 in control treatments to more than 30 mg g^?1 in limed plots, whereas concentrations of Mn, Al, Fe, and in two stands, B, decreased due to liming. In conclusion, liming with doses up to 15 Mg ha^?1 was detectable in stands 35 years after treatment. The liming significantly increased the ECM colonization of Scots pine fine roots, increased the needle nutrient concentration of Ca and decreased the needle concentrations of Mn, Al, and Fe.     

In situ gross nitrogen transformations differ between temperate deciduous and coniferous forest soils

Despite long-term enhanced nitrogen (N) inputs, forests can retain considerable amounts of N. While rates of N inputs via throughfall and N leaching are increased in coniferous stands relative to deciduous stands at comparable sites, N leaching below coniferous stands is disproportionally enhanced relative to the N input. A better understanding of factors affecting N retention is needed to assess the impact of changing N deposition on N cycling and N loss of forests. Therefore, gross N transformation pathways were quantified in undisturbed well-drained sandy soils of adjacent equal-aged deciduous (pedunculate oak (Quercus robur L.)) and coniferous (Scots pine (Pinus sylvestris L.)) planted forest stands located in a region with high N deposition (north Belgium). In situ inorganic ¹⁵N labelling of the mineral topsoil (0–10?cm) combined with numerical data analysis demonstrated that (i) all gross N transformations differed significantly (p?<?0.05) between the two forest soils, (ii) gross N mineralization in the pine soil was less than half the rate in the oak soil, (iii) meaningful N immobilization was only observed for ammonium, (iv) nitrate production via oxidation of organic N occurred three times faster in the pine soil while ammonium oxidation was similar in both soils, and (v) dissimilatory nitrate reduction to ammonium was detected in both soils but was higher in the oak soil. We conclude that the higher gross nitrification (including oxidation of organic N) in the pine soil compared to the oak soil, combined with negligible nitrate immobilization, is in line with the observed higher nitrate leaching under the pine forest.     

The ectomycorrhizal flora of primary and secondary stands of Pinus sylvestris in relation to soil conditions and ectomycorrhizal succession

Abstract. Ectomycorrhizal species composition and sporocarp abundance in two 15–20-yr old primary stands of Pinus sylvestris in the central part of the Netherlands was compared with those in two 16- and 27-yr old secondary stands of P. sylvestris in the Northeast of the country. The trees of the primary stands were spontaneously seeded in a drifting sand area. Only thin litter and humus layers were present. The trees of the 16-yr old secondary stand were planted on podzolic sandy soil and those of the 27-yr old stand on non-podzolic sandy soil. In both secondary stands thick litter and humus layers had developed. The litter and humus partly originated from former stands at the same sites and partly from the present stands. In the secondary stands the thick litter and humus layers and herb vegetation were removed (‘sod-cutting’) in order to simulate the thin litter and humus layers in the primary stands. Control treatments were present. Surveys in 1991, 1992 and 1993 showed that sod-cutting enhanced both abundance and diversity of ectomycorrhizal fungi. However, species richness and diversity were higher in the primary stands than in the secondary ones, also even in the sod-cut plots. High species richness and diversity were associated with low concentrations of nitrogen and relatively high pH in the litter and humus layers, and in the mineral soil.