Primary forest succession on poor sandy soilsas related to site factors

Three main succession lines are found to occur on poor sandy soils in a blown drift sand landscape in the Netherlands. Site factors – organic matter, parent material, soil moisture regime and relief – were determined. Site types, specific combinations of these site factors, are defined. For five site types succession was studied. The primary criterion for separating succession lines and for delineating the successional stages, was the site factor organic matter and the second criterion, groundwater level. Influence of organic matter on succession was significant on site types with a buried podzol. Species composition on these site types indicated richer sites than the species composition on the site types without a buried podzol. Groundwater level of about 1m with gleyzone was critical for species requiring moist sites. Within a succession line vegetation stages are characterized and the potential forest type is presumed. The stages were related to humus form and soil development. Litter of dominant species was suggested to be the main criterion for the humus form. Initial and older vegetation stages were linked to development of soil horizons. The defined site types, their soil development and relationships with the succession lines clarify spatial and temporal structure and diversity of vegetation, and herewith is the basic information for clarifying the heterogeneity and potential biodiversity of this blown drift sand landscape.     

Partial restoration of fungal and plant species diversity by removal of litter and humus layers in stands of Scots pine in the Netherlands

A field experiment was set up in five mature Scots pine (Pinus sylvestris) stands in the Netherlands in order to study the effects of removal of litter and humus on ectomycorrhizal fungi and vegetation of higher plants, bryophytes, and lichens. Lime was also applied as a countermeasure to the removal of surplus cations. Application of lime did not have significant effects on mycorrhizal fungi. Removal of ectorganic material increased species richness of ectomycorrhizal fungi. The number of fruit bodies was also higher in the sod-cut plots, but the difference compared with the controls was not significant in all years. A major effect of removal of the ectorganic layer was a reversion of ectomycorrhizal succession to earlier stages. Liming increased the number and cover of ruderal, nitrophilous plant species. Sod-cutting caused a reversion of the green vegetation to earlier successional stages.     

Nitrogen deposition and soil carbon content affect nitrogen mineralization during primary succession in acid inland drift sand vegetation

Background and aims

Two inland dunes in the Netherlands receiving low (24) and high (41 kg N ha^?1 yr^?1) nitrogen (N) deposition were compared for N dynamics and microbial activity to investigate the potential effect of N on succession rate of the vegetation and loss of pioneer habitats.

Methods

Primary succession stages were sampled, including bare sand, and vegetation dominated by Polytrichum piliferum, Campylopus introflexus, lichens and grasses respectively, representing a series of vegetation types in undisturbed drift sand sites with succession starting on bare sand containing virtually no organic matter. Microbial characteristics and potential N mineralization were analysed in a laboratory experiment.

Results

Organic matter accumulated during succession, resulting in a lower pH and in higher microbial biomass (bacteria and fungi), respiration and net N mineralization. The increase in respiration and N mineralization was largely due to the development of an ectorganic layer in the middle stages of succession. The observed effects of N deposition were (1) decrease of microbial biomass, (2) higher net N mineralization per m², (3) higher levels of free nitrogen in the soil, and (4) a higher microbial N:P ratio.

Conclusions

Elevated N deposition leads to higher N availability which may cause accelerated succession.     

Effects of soil organic matter and nitrogen supply on competition between Festuca ovina and Deschampsia flexuosa during inland dune succession

We tested the prediction that the successional replacement of plant species during succession on inland sand dunes results from the effects of an increase in nitrogen mineralization on competitive interactions. The growth and competitive strength of Festuca ovina and Deschampsia flexuosa on soil substrates with different amounts of soil organic matter or nitrogen supply were measured. Small tillers of Festuca ovina and Deschampsia flexuosa were grown in monocultures and 1:1-mixtures on soil columns with undisturbed layers of soil organic matter from different successional age. There was (a) no visible soil organic matter, (b) a thin soil organic layer (0.5 cm) and (c) a thicker soil organic layer (6.0 cm) present on the soil columns. The species were also grown on columns with no visible soil organic matter (bare sand) with two different levels of N fertilization to mimic the increased N mineralization in the older successional stages.In monoculture, Festuca produced more biomass on the substrates with a soil organic layer compared to the unfertilized sand substrate. It also produced more biomass on sand substrates with N fertilization. Deschampsia produced more biomass in treatments with a soil organic layer compared to the bare sand treatments, but did not respond to the ammonium-nitrate addition. In competition, Festuca seemed to be the stronger competitor on the unfertilized sand substrate. Festuca was also the better competitor on the N fertilized sand treatments, while on the treatments with a soil organic layer Deschampsia was the winning species. Our results do not support the hypothesis that an increase in N supply is responsible for the replacement of Festuca by Deschampsia that concur with the accumulation of soil organic matter during succession in inland dunes.     

Carbon Quality and Stocks in Organic Horizons in Boreal Forest Soils

We investigated the mechanisms that determine the quality and quantity of organic carbon (C) stocks in boreal forest soils by analyzing both qualitative and quantitative changes in the organic fractions in the soil organic matter (OM) in a vertical gradient in the decomposition continuum of the organic horizon [litter layer (L), fermentation layer (F), and humus layer (H)] in forest soils using a sequential fractionation method at two forest types along a climatic gradient in Finland. We predicted that the concentrations of water-soluble (WSE) and non-polar (NPE) extractives should decrease and those of the acid-soluble (AS) fraction and acid-insoluble residue (AIR) should increase from the L to the F, and from the F to the H layers, but the C/N ratio of soil OM should stay constant after reaching the critical quotient. We also predicted that the AIR concentrations should be higher in the south than north boreal, and in sub-xeric than mesic forests. Consistent with our hypothesis, the concentrations of WSE and NPE fractions decreased and concentrations of AIR increased in the vertical soil gradient. The highest concentrations of the AS fraction were found in the F layer. The C/N ratio was lowest in the F layer, and the highest in the H layer, indicating that soil OM is depleted in N in relation to C along the vertical soil gradient. Concentrations of WSE and NPE were lower, and concentrations of AIR were higher in the south than in north boreal forests, which is in agreement with our hypothesis that higher soil temperatures may enhance accumulation of slowly decomposable OM in the soil. The concentrations of AIR were higher in the sub-xeric than mesic forests. Contrary to our expectations, however, the differences in the chemical quality in soil OM between the site types were amplified from the L to the H layer. The size of the C storage was significantly larger in south than north boreal sites, and larger in the mesic than in the sub-xeric sites.     

Carbon dynamics in successional and afforested spruce stands in Thuringia and the Alps

Changes in the carbon stocks of stem biomass, organic layers and the upper 50 cm of the mineral soil during succession and afforestation of spruce (Picea abies) on former grassland were examined along six chronosequences in Thuringia and the Alps. Three chronosequences were established on calcareous and three on acidic bedrocks. Stand elevation and mean annual precipitation of the chronosequences were different. Maximum stand age was 93 years on acid and 112 years on calcareous bedrocks. Stem biomass increased with stand age and reached values of 250–400 t C ha^?1 in the oldest successional stands. On acidic bedrocks, the organic layers accumulated linearly during forest succession at a rate of 0.34 t C ha^?1 yr^?1. On calcareous bedrocks, a maximum carbon stock in the humus layers was reached at an age of 60 years. Total carbon stocks in stem biomass, organic layers and the mineral soil increased during forest development from 75 t C ha^?1 in the meadows to 350 t C ha^?1 in the oldest successional forest stands (2.75 t C ha^?1 yr^?1). Carbon sequestration occurred in stem biomass and in the organic layers (0.34 t C ha^?1 yr^?1on acid bedrock), while mineral soil carbon stocks declined. Mineral soil carbon stocks were larger in areas with higher precipitation. During forest succession, mineral soil carbon stocks of the upper 50 cm decreased until they reached approximately 80% of the meadow level and increased slightly thereafter. Carbon dynamics in soil layers were examined by a process model. Results showed that sustained input of meadow fine roots is the factor, which most likely reduces carbon losses in the upper 10 cm. Carbon losses in 10–20 cm depth were lower on acidic than on calcareous bedrocks. In this depth, continuous dissolved organic carbon inputs and low soil respiration rates could promote carbon sequestration following initial carbon loss. At least 80 years are necessary to regain former stock levels in the mineral soil. Despite the comparatively larger amount of carbon stored in the regrowing vegetation, afforestation projects under the Kyoto protocol should also aim at the preservation or increase of carbon in the mineral soil regarding its greater stability of compared with stocks in biomass and humus layers. If grassland afforestation is planned, suitable management options and a sufficient rotation length should be chosen to achieve these objectives. Maintenance of grass cover reduces the initial loss.     

浙江天童植物群落演替对土壤化学性质的影响

对浙江天童植物群落演替过程的土壤腐殖质层、0～10cm层和10～20cm层化学因子的比较研究结果表明,随着进展演替,土壤全氮含量呈明显增长趋势,但硝态氮、氨态氮、速效氮、速效磷和速效钾含量除演替初期的裸地、灌丛较小外,森林群落阶段增长趋势一般不明显;土壤pH值则缺乏明显的规律性,但有机质含量呈明显增长趋势,而富里酸和胡敏酸含量,除栲树木荷林和栲树林较高,裸地最低外,其它演替阶段增长趋势也不明显;HA/FA在腐殖质层缺乏明显的规律性,但在0～10cm和10～20cm层则呈减少趋势。可见,常绿阔叶林次生演替能增加土壤有机物质含量,促进土壤有机物质的矿化和再合成作用;而土壤化学性状的改善,也为常绿阔叶林的进展演替奠定了基础。     

兴安落叶松林火干扰后土壤有机碳含量变化

在大兴安岭兴安落叶松林区,选择不同恢复年限各种火烧强度(重度、中度、轻度)的火烧迹地进行调查并采集土壤样品,对火后有机层和矿质层有机碳含量变化进行研究,以期为进一步开展森林火灾对区域碳平衡影响的定量评估提供科学依据。研究结果表明:火干扰对土壤有机碳含量变化的影响包括火烧即时影响和火后生境条件变化带来的间接影响。火干扰样地有机层的积累与转化主要是通过火后林冠郁闭度的变化影响的,其有机碳总储量低于对照样地。对于矿质层土壤,重度和中度火干扰后,如果样地发生植被序列演替,即阔叶林植被入侵,样地郁闭度增加迅速,凋落物积累量增大,土壤有机碳含量将随着过火年限的增加而增加;如果样地发生自我更新,样地郁闭度增加缓慢,凋落物分解量大于积累量,土壤有机碳含量将会在一定时间内随着过火年限的增加而减少;但是,无论样地发生植被序列演替还是自我更新,土壤有机碳含量短时间内均无法恢复到火前水平。轻度火干扰后,土壤有机碳含量短期内先增加,随着植被更新情况的发展最终趋于平衡。     

Mechanical resistance by an ectorganic soil layer on root development of seedling Pinus sylvestris

We investigated early root development of Pinus sylvestris seedlings in relation to bulk density and natural particle layering in an ectorganic soil layer from a bracken (Pteridium aquilinum) stand. Responses in root development to two levels of bulk density (0.07 and 0.15 g/cm³) in mixed bracken substrate were compared with effects in peat of similar bulk densities, and in sand of three different bulk densities (0.37, 0.52, and 0.67 g/cm³). The effect on root growth of the natural horizontal layering of the organic particles was examined by comparing intact with mixed ectorganic bracken soil profiles of similar bulk densities (resp. 0.09 and 0.07 g/cm³).Root length growth was significantly reduced in the organic and sandy substrates of high bulk density. Root diameter was not affected by bulk density in the organic substrate, but increased with higher bulk density in sand. Preservation of horizontal layering in the intact ectorganic profile significantly reduced root length compared with mixed substrate of similar bulk density.Roots growing in high bulk density, and intact, organic substrate showed increased twisting, which resulted in a smaller depth reached by the root relative to total root length produced. In sand, root twisting did not change with increased bulk density. It is suggested that roots growing through organic substrate follow a path of least resistance. This implies that organic particle size and orientation are more important in determining root development than bulk density.This study points out that the natural layering of organic particles presents another constraint on the establishment of plant species in sites with a well-developed ectorganic soil layer. Disturbance of this layer may therefore enhance establishment of seedlings by reducing the mechanical resistance of the ectorganic soil profile to developing seedling roots.     

Earthworm invasion into previously earthworm-free temperate and boreal forests

Earthworms are keystone detritivores that can influence primary producers by changing seedbed conditions, soil characteristics, flow of water, nutrients and carbon, and plant–herbivore interactions. The invasion of European earthworms into previously earthworm-free temperate and boreal forests of North America dominated by Acer, Quercus, Betula, Pinus and Populus has provided ample opportunity to observe how earthworms engineer ecosystems. Impacts vary with soil parent material, land use history, and assemblage of invading earthworm species. Earthworms reduce the thickness of organic layers, increase the bulk density of soils and incorporate litter and humus materials into deeper horizons of the soil profile, thereby affecting the whole soil food web and the above ground plant community. Mixing of organic and mineral materials turns mor into mull humus which significantly changes the distribution and community composition of the soil microflora and seedbed conditions for vascular plants. In some forests earthworm invasion leads to reduced availability and increased leaching of N and P in soil horizons where most fine roots are concentrated. Earthworms can contribute to a forest decline syndrome, and forest herbs in the genera Aralia, Botrychium, Osmorhiza, Trillium, Uvularia, and Viola are reduced in abundance during earthworm invasion. The degree of plant recovery after invasion varies greatly among sites and depends on complex interactions with soil processes and herbivores. These changes are likely to alter competitive relationships among plant species, possibly facilitating invasion of exotic plant species such as Rhamnus cathartica into North American forests, leading to as yet unknown changes in successional trajectory.     

Development of vegetation in dune slack wetlands of Cape Cod National Seashore (Massachusetts,USA)

Interdunal seasonal wetlands, known as dune slack wetlands, were mapped and their vegetation surveyed across a large region of dunes within the Cape Cod National Seashore. Wetland sizes and ages were estimated from digitized, georeferenced aerial photographs available from 1938, 1947, 1960, 1986, 1994, and 2001, and from LIDAR elevation data. A total of 346 sites were found, covering an area of ∼45.4 ha., in which 97 species of vascular plants were identified. Vegetation structure and composition exhibited a distinct sequence of development with age, following a pattern of succession from herbaceous, graminoid-dominated communities to shrub- and tree-dominated communities. Floristic variables were not related to proximity to the coast and although wetland size appeared to have some bearing on species richness, the correlation was statistically weak. Soil organic matter determined for a subset of 60 wetlands was positively correlated with age and woody cover but showed no relationship with water depth. The results suggest that vegetation development is primarily driven by the internal mechanisms of succession. Notwithstanding, any changes in the environment that alter the process of succession will collectively influence these wetlands. In addition, stabilization of the dunes resulting in a reduction in the formation of new wetlands, may translate into permanent loss of early and mid-successional dune slack communities. Maintenance of these communities will depend on succession being periodically reset by disturbance or active management.     

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.     

Specific features of soil organic matter in forest landscapes of Karelia

The organic matter reserves and the soil humus state are assessed in three types of landscapes of the middle taiga (Karelia). Peat soils are the main organic matter reservoir. Hence, the greater their area, the higher the organic matter reserves in a particular landscape. The organic matter stocks in the group of semihydromorphic and hydromorphic soils clearly correlate with the degree of their waterlogging. The distribution of organic matter reserves within these soils depends on the ratio between the areas of boggy forests and open mires. The forest supporting quality of the soil is related to the organic matter composition and parent rock (the properties of the soil mineral horizons) rather than to the organic matter stocks. The data obtained may be used for assessing the forest supporting qualities of the soils and the basis for estimating the carbon budget in the landscapes.     

Temporal and spatial variability in soil food web structure

Heterogeneity is a prominent feature of most ecosystems. As a result of environmental heterogeneity the distribution of many soil organisms shows a temporal as well as horizontal and vertical spatial patterning. In spite of this, food webs are usually portrayed as static networks with highly aggregated trophic groups over broader scales of time and space. The variability in food web structure and its consequences have seldom been examined. Using data from a Scots pine forest soil in the Netherlands, we explored (1) the temporal and spatial variability of a detrital food web and its components, (2) the effect of taxonomic resolution on the perception of variability over time and across space, and (3) the importance of organic matter quality as an explanatory factor for variability in food web composition. Compositional variability, expressed using the Bray‐Curtis similarity index, was measured over 2.5 years using a stratified litterbag design with three organic horizons per litterbag set. Variability in community composition and organic matter degradation increased over time in the litter horizon only. Seasonal variation in community composition was larger than variation between samples from the same season in different years. Horizontal spatial variability in community composition and organic matter degradation was relatively low, with no increase in variability with increasing distance between samples. Vertically, communities and organic matter degradation was more different between the non‐adjacent litter and humus horizons than between adjacent layers. These findings imply that soil food webs, at least in temperate forest plantations, are more variable than is currently appreciated in experiments and model studies, and that organic matter turnover might be an important factor explaining variability in community composition. Species composition was more variable than functional group composition, which implies that aggregated food webs will seem less sensitive to local temporal and spatial changes than they in fact are.     

Fine litter input to terrestrial humus forms in Colombian Amazonia

A comparative litter fall study was made in five rain forest stands along a gradient of humus form development and soils in the Amazon lowlands of eastern Colombia. The total fine litter fall was highest in a plot on a well drained soil of the flood plain of the Caquetá River (1.07 kg · m^-2 · y^-1), lower in three plots on well drained upland soils (0.86, 0.69, and 0.68 kg · m^-2 · y^-1), and lowest in a plot on a poorly drained, upland podzolised soil (0.62 kg · m^-2 · y^-1). In the four upland plots, leaf litter fall patterns were highly associated, which points at climatic regulation. Litter resource quality, as represented by nutrient concentrations and area/weight ratio of the leaf litter fall, was comparatively high in the flood plain plot. In the upland plots, concentrations and fluxes of Ca, Mg, K, and P were as low as in oligotrophic central Amazonian upland forests. This questions generalisations that the western peripheral region of the Amazon basin should be less oligotrophic than central Amazonia. The upland plot on the podzolised soil showed the lowest concentrations and fluxes of N. Mean residence times of organic matter and nutrients in the L horizons hardly differed between the five plots, suggesting that edaphic properties and litter resource quality are of little importance in the first step of decomposition. Mean residence time of organic matter in all ectorganic horizons combined (estimated on the basis of litter input and necromass on the forest floor, and uncorrected for dead fine root input) varied from 1.0 y in the flood plain forest, 1.1–3.3 y in the well drained upland forests, and 10.2 y in the forest on the podzolised soil.     

Testing the assumptions of chronosequences in succession

Many introductory ecology textbooks illustrate succession, at least in part, by using certain classic studies (e.g. sand dunes, ponds/bogs, glacial till, and old fields) that substituted space for time (chronosequence) in determining the sequences of the succession. Despite past criticisms of this method, there is continued, often uncritical, use of chronosequences in current research on topics besides succession, including temporal changes in biodiversity, productivity, nutrient cycling, etc. To show the problem with chronosequence-based studies in general, we review evidence from studies that used non-chronosequence methods (such as long-term study of permanent plots, palynology, and stand reconstruction) to test the space-for-time substitution in four classic succession studies. In several cases, the tests have used the same locations and, in one case, the same plots as those in the original studies. We show that empirical evidence invalidates the chronosequence-based sequences inferred in these classic studies.     

Mercury in the soil at the mouth of the Angara and Belaya interfluve (Irkutsk region)

The results of investigation of soils in the mouth of the Angara and Belaya interfluve within the zone of influence of a chemical plant in Usol’e-Sibirskoe are considered. Estimations of the degree, scale, and character of the areal distribution of mercury, as well as sulfate and chloride ions are given. The prevalence of surface run off of mercury over soil run off is given. The main fields of anomalous concentrations of mercury and anions occur in the industrial zone of a chemical plant. A correlation of mercury with organic carbon and sulfate ion for the humus horizons and with oxides of iron and aluminum, manganese, titanium, and phosphorus for the mineral horizons of the studied soils was established.     

Small-scale patterning in the recruitment of forest species during succession in tropical dry forest,Mozambique

Within a few years of field abandonment on the coastal dunes, a considerable number of species of trees and woody lianes of the climax evergreen forest enter the succession. At a relatively large-scale of analysis (a 100 m² plot), there is no evidence of soil development during the early succession, with no accumulation of soil organic matter nor nutrients being detected. At this scale there is, therefore, little support for the facilitation model of succession. However, when smaller scales of analysis are employed (e.g. a plot equivalent to the canopy area of a sapling) a very different picture emerges; most recruitment of forest woody species occurs beneath saplings found in early succession. Nutrient concentrations were higher in soils under sapling canopies than in soils beyond sapling canopies. With increasing sapling size, an increasing number of seedlings of forest woody species were found and there is progressive soil amelioration. Thus, during the early succession, there is considerable small-scale patterning in soil development and recruitment. This pattern is largely established through bird dispersal, with the intensity of recruitment being dependent on the distance to seed sources.     

Carbon stock evaluation from topsoil of forest stands in NE Italy

Gas emissions from anthropic activities, particularly CO2, are responsible for global warming. Soil is a major carbon sink on a planetary level, thereby contributing to mitigate greenhouse effect. In the present work, the objectives were: 1) to evaluate the topsoil carbon stock of different forest stands in NE Italy, and 2) to outline the relationships among humus forms, soil organic matter dynamics, and actual carbon stock under different vegetation coverage, with reference to climate change. Five forest stands and the related topsoils, were selected in the Dolomites area. The humus forms were examined in the field and samples were carried to the lab for further physical-chemical analyses. The carbon stock for each soil was calculated by means of pedotransfer functions. The less developed humus forms, as the Dysmull and the Hemimoder, presented the highest carbon storage capacity (168 t/y and 129 t/y), followed by Lithoamphimus (123 t/y) and Eu-amphimus (96 t/y), and by Oligomull (86 t/y). Organic horizons proved to recover 36% of the total carbon stocked along the soil profile, and this points to humus layers as a fundamental tool in carbon stock evaluation. Positive correlations between elevation, humus forms and soil carbon pools were found.