Soil organic C and nutrient contents under trees with different functional characteristics in seasonally dry tropical silvopastures

Aims

The selection of tree characteristics is critical for the outcome of the tree effects on soil fertility in silvopastoral pastures. This study aims to quantify the effects of trees on soil nutrient and C stocks, as well as assessing differences on the effects between legume (Albizia saman; Enterolobium cyclocarpum) and non-legume tree species (Tabebuia rosea; Guazuma ulmifolia).

Methods

In Central Nicaragua, soil was sampled (0–10 cm deep) in paired plots, under both a canopy and in open grassland, in 12 sites per tree species and analysed for organic C, total N stocks, available P and extractable K⁺, Ca²⁺ and Mg²⁺. To assess the effects of herbaceous composition and cattle to soil proprieties, we recorded the cover of plant groups and assessed the mass of dung in each plot.

Results

Soil organic C and N, available P and extractable K⁺ and Ca²⁺ were higher under the tree canopy than under paired open grassland. The basal area of trees was positively related with the canopy effect on soil variables, thus suggesting that the age or sizes of the trees are relevant factors associated with the content of soil C and nutrients. No specific effects related to the legume species group were detected.

Conclusions

Our results indicate that in fertile seasonally dry subtropical pastures, scattered trees have an overall effect on soil fertility, and that the magnitude of the effect depends more on the tree characteristics (i.e. basal area, crown area) than on whether the species is a legume or not.     

The importance of limestone bedrock and dissolution karst features on tree root distribution in northern Yucatán, México

Background and Aims

With limited soil depth in northern Yucatán (<30 cm), roots grow deeper through rock fractures and dissolution karst features (i.e., cavities, including soil-filled ones known as soil pockets). We assessed the importance of limestone bedrock and dissolution karst features on tree root growth.

Methods

Fieldwork was conducted in a limestone quarry where the relative proportions of rock matrix, empty cavities, and soil pockets were calculated by observing recently exposed walls. Physical properties of rocks, topsoil, and soil pockets were analyzed. Root distribution was assessed and roots identified.

Results

Soil pockets represented 9% of the rock matrix. The physical properties of rock layers were different with depth. Available water capacity is higher in soil (0.11 m^-3 m^-3) than in rock layers (<0.05 m^-3 m^-3). But potential available water was much higher in subsurface features than top soil.

Conclusions

Dissolution karts features allow roots to grow deep into the bedrock, tapping water stored there. Although the limestone upper layer in northern Yucatan is highly restrictive to root growth, subsurface limestone layers and soil pockets are not restrictive and can hold important amounts of water.     

Effects of tree species composition on the CO2 and N2O efflux of a Mediterranean mountain forest soil

Background and Aims

Tree species composition shifts can alter soil CO₂ and N₂O effluxes. We quantified the soil CO₂ and N₂O efflux rates and temperature sensitivity from Pyrenean oak, Scots pine and mixed stands in Central Spain to assess the effects of a potential expansion of oak forests.

Methods

Soil CO₂ and N₂O effluxes were measured from topsoil samples by lab incubation from 5 to 25 °C. Soil microbial biomass and community composition were assessed.

Results

Pine stands showed highest soil CO₂ efflux, followed by mixed and oak forests (up to 277, 245 and 145 mg CO₂-C m^?2 h^?1, respectively). Despite contrasting soil microbial community composition (more fungi and less actinomycetes in pine plots), carbon decomposability and temperature sensitivity of the soil CO₂ efflux remain constant among tree species. Soil N₂O efflux rates and its temperature sensitivity was markedly higher in oak stands than in pine stands (70 vs. 27 μg N₂O-N m^?2 h^?1, Q₁₀, 4.5 vs. 2.5).

Conclusions

Conversion of pine to oak forests in the region will likely decrease soil CO₂ effluxes due to decreasing SOC contents on the long run and will likely enhance soil N₂O effluxes. Our results present only a seasonal snapshot and need to be confirmed in the field.     

Soil CO2 efflux in a bioenergy plantation with fast-growing Populus trees – influence of former land use, inter-row spacing and genotype

Aims

In this study we quantified the annual soil CO₂ efflux (annual SCE) of a short rotation coppice plantation in its establishment phase. We aimed to examine the effect of former (agricultural) land use type, inter-row spacing and genotype.

Methods

Annual SCE was quantified during the second growth year of the establishment rotation in a large scale poplar plantation in Flanders. Automated chambers were distributed over the two former land use types, the two different inter-row spacings and under two poplar genotypes. Additional measurements of C, N, P, K, Mg, Ca and Na concentrations of the soil, pH, bulk density, fine root biomass, microbial biomass C, soil mineralization rate, distance to trees and tree diameters were performed at the end of the second growth year.

Results

Total carbon loss from soil CO₂ efflux was valued at 589 g m^?2 yr^?1. Annual SCE was higher in former pasture as compared to cropland, higher in the narrow than in the wider inter-row spacings, but no effect of genotype was found.

Conclusions

Spatial differences in site characteristics are of great importance for understanding the effect of ecosystem management and land use change on soil respiration processes and need to be taken into account in modeling efforts of the carbon balance.     

Uptake of water from a Kandosol subsoil: I. Determination of soil water diffusivity

Aims

To determine soil water diffusivity, D(θ), on undisturbed field soil at medium to low water content (suction range from 10 to 150 m of water), for the purpose of modeling the uptake of water by plant roots.

Methods

The method is based on the analysis of one-step outflow induced by a turbulent stream of dry air over the exposed end of a soil core, with the other end of the core enclosed. The outflow is measured through time as the change in the weight of the core as it sits on a recording balance. D(θ) is calculated by deconvoluting the measured outflow function.

Results

Over the suction range of 10 to 150 m of water, D(θ) calculated on the undisturbed soil ranged from 20?×?10^?9 to 10?×?10^?9 [m²?s^?1], substantially higher than other published estimates over this range in suction.

Conclusions

These unusually large values cast doubt on the view that flow of water to roots limits uptake of water from the targeted subsoil.     

Overstorey and juvenile response to thinning and drought in a jarrah (Eucalyptus marginata Donn ex Sm.) forest of southwestern Australia

Aims

Forest thinning is expected to affect tree water use and carbon assimilation, but the related influence from climate variability is little known. Recent forest thinning in the Wungong catchment coincided with a record dry year following the thinning, which provides a rare opportunity to understand the climate influence on the thinning effect.

Methods

A field experiment was conducted to examine changes before and after thinning, especially the rainfall, soil moisture, leaf water status, tissue isotope signature (¹³?C and ¹⁵?N) and N concentration of overstorey and understorey juvenile trees of Eucalyptus marginata (Donn ex Sm.).

Results

Despite the post-thinning drought, surface soil was moister and juvenile jarrah plants were less water stressed, attributable to reduced rain interception and transpiration as a result of less canopy cover. The overstorey was under stress but mainly due to drought rather than by thinning. The concentration of N declined in both tree stems and juvenile leaves along with available N in soil, suggesting a soil N limitation. No treatment effects were detected from leaf relative water content and tissue isotope signature (¹³?C and ¹⁵?N).

Conclusions

The drought effects were superimposed over the thinning effects on overstorey growth, with stemwood δ¹³C being a major indicator of water stress. The water relations and carbon assimilation of understorey juveniles were however dependent more on topsoil moisture, and the wetter soil during the year following thinning enhanced growth activity and hence the depletion of ¹³?C (more negative δ¹³C) in juvenile leaves.     

Variability of stomatal conductance, leaf anatomy, and seasonal leaf wettability of young and adult European beech leaves along a vertical canopy gradient

This study assessed the variation of leaf anatomy, chlorophyll content index (CCI), maximal stomatal conductance (g _s ^max ) and leaf wettability within the canopy of an adult European beech tree (Fagus sylvatica L.) and for beech saplings placed along the vertical gradient in the canopy. At the top canopy level (CL_28m) of the adult beech, CCI and leaf anatomy reflected higher light stress, while g _s ^max increased with height, reflecting the importance of gas exchange in the upper canopy layer. Leaf wettability, measured as drop contact angle, decreased from 85.5°?±?1.6° (summer) to 57.5°?±?2.8° (autumn) at CL_28m of the adult tree. At CL_22m, adult beech leaves seemed to be better optimized for photosynthesis than the CL_28m leaves because of a large leaf thickness with less protective and impregnated substances, and a higher CCI. The beech saplings, in contrast, did not adapt their stomatal characteristics and leaf anatomy according to the same strategy as the adult beech leaves. Consequently, care is needed when scaling up experimental results from seedlings to adult trees.     

Effects of experimental warming and nitrogen fertilization on soil microbial communities and processes of two subalpine coniferous species in Eastern Tibetan Plateau,China

Aim

This study aimed at predicting how sub-alpine coniferous ecosystems respond to global changes in the Eastern Tibetan Plateau by understanding soil microbial communities and activities, as well as variation in the quality and quantity of soil organic matter.

Methods

An experiment was conducted to examine soil microbial communities and their related soil processes in rhizospheric soil of two coniferous species that were exposed to two levels of temperature (unwarmed and infrared heater warming) and two levels of nitrogen (unfertilized and 25 g N m^?2 a^?1) from April 2007.

Results

Four-year night warming alone slightly affected the phospholipid fatty acid contents of the microbial community. However, the combination of nitrogen addition and soil warming significantly affected soil microbial composition while reducing the biomass of major microbial groups and the activities of most enzymes, especially in Abies faxoniana plots. The combination of warming and nitrogen addition increased soil labile C and N pools in Picea asperata plots and was beneficial for soil recalcitrant C, as well as for labile and total C and N pools in A. faxoniana plots.

Conclusion

Results indicated that future warming will slightly affect soil microbial communities and their related soil processes. However, warming combined with high nitrogen deposition will significantly constrain soil microbial biomass and enzyme activities, consequently increasing soil C and N pools in sub-alpine coniferous forests of this region.     

Microbial-mediated feedbacks of leaf litter on invasive plant growth and interspecific competition

Background and Aims

In line with the Stress Gradient Hypothesis, studies of facilitation have tended to focus on plant–plant interactions (biotic nurses), while the relative role of abiotic nurses has been little studied. We assessed the role of biotic and abiotic nurses, and their interaction, on soil enhancement and the consequential performance of a native annual grass, Dactyloctenium radulans.

Methods

We used a growth chamber study with two levels of water application to compare the performance of D. radulans growing in soil from foraging pits of the Short-beaked echidna (Tachyglossus aculeatus; abiotic nurse) and non-pit soil from either under tree canopies (biotic nurse) or surrounding open areas.

Results

All measures of plant performance were more pronounced under the high than the low water treatment. The greatest differences between pit and surface Microsites occurred under the low water application, reinforcing our view that facilitatory effects are greater in resource-limited environments. Despite tree canopy soil having greater N, there was no significant effect on plant performance, nor any significant interaction with Microsite.

Conclusions

Our study provides strong evidence that foraging pits enhance soil properties and this soil, in turn, facilitates plant growth; and supports previous work documenting the positive effect of nurse-protégé interactions under greater levels of abiotic stress.     

Effects of irrigation on the soil CO2 efflux from different poplar clone plantations in arid northwest China

Aims

Soil respiration in forest plantations can be greatly affected by management practices such as irrigation. In northwest China, soil water is usually a limiting factor for the development of forest plantations. This study aims to examine the effects of irrigation intensity on soil respiration from three poplar clone plantations in this arid area.

Methods

The experiment included three poplar clones subjected to three irrigation intensities (without, low and high). Soil respiration was measured using a Li-6400-09 chamber during the growing season in 2007.

Results

Mean soil respiration rates were 2.92, 4.74 and 3.49 μmol m^?2 s^?1 for control, low and high irrigation treatments, respectively. Soil respiration decreased once soil water content was below a lower (14.8 %) or above an upper (26.2 %) threshold. When soil water content ranged from 14.8 % to 26.2 %, soil respiration increased and correlated with soil temperature. Fine root also played a role in the significant differences in soil CO₂ efflux among the three treatments. Furthermore, the three poplar hybrid clones responded differently to irrigation regarding fine root production and soil CO₂ efflux.

Conclusions

Irrigation intensity had a strong impact on soil respiration of the three poplar clone plantations, which was mainly because fine root biomass and microbial activities were greatly influenced by soil water conditions. Our results suggest that irrigation management is a main factor controlling soil carbon dynamics in forest plantation in arid regions.     

Phenotypic seedling responses of a metal-tolerant mutant line of sunflower growing on a Cu-contaminated soil series: potential uses for biomonitoring of Cu exposure and phytoremediation

Background and aims

The potential use of a metal-tolerant sunflower mutant line for both biomonitoring and phytoremediating a Cu-contaminated soil series was investigated.

Methods

The soil series (21–1,170 mg Cu kg^?1) was sampled in field plots at control and wood preservation sites. Sunflowers were cultivated 1 month in potted soils under controlled conditions.

Results

pH and dissolved organic matter influenced Cu concentration in the soil pore water. Leaf chlorophyll content and root growth decreased as Cu exposure rose. Their EC₁₀ values corresponded to 104 and 118 μg Cu L^?1 in the soil pore water, 138 and 155 mg Cu kg^?1 for total soil Cu, and 16–18 mg Cu kg^?1 DW shoot. Biomass of plant organs as well as leaf area, length and asymmetry were well correlated with Cu exposure, contrary to the maximum stem height and leaf water content.

Conclusions

Physiological parameters were more sensitive to soil Cu exposure than the morphological ones. Bioconcentration and translocation factors and distribution of mineral masses for Cu highlighted this mutant as a secondary Cu accumulator. Free Cu²⁺ concentration in soil pore water best predicted Cu phytoavailability. The usefulness of this sunflower mutant line for biomonitoring and Cu phytoextraction was discussed.     

Spatial distribution of the soil organic carbon pool in a Holm oak dehesa in Spain

Aims

Dehesas are agroforestry systems characterized by scattered trees among pastures, crops and/or fallows. A study at a Spanish dehesa has been carried out to estimate the spatial distribution of the soil organic carbon stock and to assess the influence of the tree cover.

Methods

The soil organic carbon stock was estimated from the five uppermost cm of the mineral soil with high spatial resolution at two plots with different grazing intensities. The Universal Kriging technique was used to assess the spatial distribution of the soil organic carbon stocks, using tree coverage within a buffering area as an auxiliary variable.

Results

A significant positive correlation between tree presence and soil organic carbon stocks up to distances of around 8 m from the trees was found. The tree crown cover within a buffer up to a distance similar to the crown radius around the point absorbed 30 % of the variance in the model for both grazing intensities, but residual variance showed stronger spatial autocorrelation under regular grazing conditions.

Conclusions

Tree cover increases soil organic carbon stocks, and can be satisfactorily estimated by means of crown parameters. However, other factors are involved in the spatial pattern of the soil organic carbon distribution. Livestock plays an interactive role together with tree presence in soil organic carbon distribution.     

Fine root biomass and turnover of two fast-growing poplar genotypes in a short-rotation coppice culture

Background and aims

The quantification of root dynamics remains a major challenge in ecological research because root sampling is laborious and prone to error due to unavoidable disturbance of the delicate soil-root interface. The objective of the present study was to quantify the distribution of the biomass and turnover of roots of poplars (Populus) and associated understory vegetation during the second growing season of a high-density short rotation coppice culture.

Methods

Roots were manually picked from soil samples collected with a soil core from narrow (75 cm apart) and wide rows (150 cm apart) of the double-row planting system from two genetically contrasting poplar genotypes. Several methods of estimating root production and turnover were compared.

Results

Poplar fine root biomass was higher in the narrow rows than in the wide rows. In spite of genetic differences in above-ground biomass, annual fine root productivity was similar for both genotypes (ca. 44 g DM m^?2 year^?1). Weed root biomass was equally distributed over the ground surface, and root productivity was more than two times higher compared to poplar fine roots (ca. 109 g DM m^?2 year^?1).

Conclusions

Early in SRC plantation development, weeds result in significant root competition to the crop tree poplars, but may confer certain ecosystem services such as carbon input to soil and retention of available soil N until the trees fully occupy the site.     

Soil carbon fractions in grasslands respond differently to various levels of nitrogen enrichments

Background and Aims

Soil contains many different C fractions which have diverse physical and chemical compositions. Examining these differential soil C fractions in response to N enrichment is helpful for better understanding soil C changes under the predominantly increasing N deposition. In this study, we used a field N addition experiment in a grassland to explore the effects of various N enrichment levels on soil C fractions.

Methods

We conducted a field manipulative experiment which used a Latin square design with six N addition levels of 0, 2, 4, 8, 16 and 32 g N m^?2 year^?1 since 2003 in a semiarid grassland in northern China. Soil samples were collected in August (when plants have the greatest biomass), 2011. We measured C and N concentrations in soil light fraction, microbial biomass, extractable organic matter, heavy fraction, and total soil C and N.

Results

The results showed that total soil C and N, and heavy fraction C and N were not significantly affected by N addition after 9 years of treatments. In contrast, different N enrichment levels changed soil light fraction C and N, ranging from 4.3 to 27.7 % and 3.3–30.0 %, respectively. Moreover, both light fraction C and N had a nonlinear relationship with N addition rates, and the threshold for N-induced change in light fraction C and N was near 16 g N m^?2 year^?1 in this semiarid grassland. Increases of soil light fraction C and N primarily resulted from changes in biotic (N-stimulated aboveground biomass) and abiotic (soil temperature, moisture and pH) factors under N enrichment. Soil microbial biomass exponentially declined with increasing N, but extractable organic C showed a positive linear response to N enrichment rates. Changes in microbial biomass C and extractable organic C were primarily due to the reduced soil pH under N addition.

Conclusions

Our findings suggest that various soil C fractions differentially respond to elevated N, because different sets of biotic and abiotic factors regulate those fractions under N enrichment.     

The critical soil P levels for crop yield, soil fertility and environmental safety in different soil types

Background and aims

Sufficient soil phosphorus (P) is important for achieving optimal crop production, but excessive soil P levels may create a risk of P losses and associated eutrophication of surface waters. The aim of this study was to determine critical soil P levels for achieving optimal crop yields and minimal P losses in common soil types and dominant cropping systems in China.

Methods

Four long-term experiment sites were selected in China. The critical level of soil Olsen-P for crop yield was determined using the linear-plateau model. The relationships between the soil total P, Olsen-P and CaCl₂-P were evaluated using two-segment linear model to determine the soil P fertility rate and leaching change-point.

Results

The critical levels of soil Olsen-P for optimal crop yield ranged from 10.9 mg kg^?1 to 21.4 mg kg^?1, above which crop yield response less to the increasing of soil Olsen-P. The P leaching change-points of Olsen-P ranged from 39.9 mg kg^?1 to 90.2 mg kg^?1, above which soil CaCl₂-P greatly increasing with increasing soil Olsen-P. Similar change-point was found between soil total P and Olsen-P. Overall, the change-point ranged from 4.6 mg kg^?1 to 71.8 mg kg^?1 among all the four sites. These change-points were highly affected by crop specie, soil type, pH and soil organic matter content.

Conclusions

The three response curves could be used to access the soil Olsen-P status for crop yield, soil P fertility rate and soil P leaching risk for a sustainable soil P management in field.     

Soil penetration resistance, a suitable soil property to account for variations in root elongation and branching

Background and Aims

Root system development is affected by soil conditions. The effects of bulk density, water content and penetration resistance on root development processes were investigated in peach trees.

Methods

Peach tree rootstocks were grown in various soil conditions, combining two bulk densities (1.2 and 1.5?g soil.cm^-3) and three water contents (0.14, 0.17 and 0.20?g.g^-1soil). Root parameters (tip diameter, length of apical unbranched zone, branching density and diameters of main and lateral roots) and plant growth (leaves, branches, trunk, root dry mass) were measured. Root growth processes (elongation, branching) were studied using relationships between root parameters.

Results

The proportion of biomass allocated to each plant compartment was similar whatever the soil conditions. Variations in root development were best explained by the variation in penetration resistance, rather than other soil properties. Increased soil penetration resistance reduced the root elongation rate, especially for thick roots. In addition, the branching pattern was affected. In soil with a high penetration resistance, the root system shape differs from a typical herringbone pattern.

Conclusions

These results allow quantification of the root system plasticity, and improve our understanding of the interactions between root development and soil properties.     

Root traits and soil properties in harvested perennial grassland,annual wheat,and never-tilled annual wheat

Background and aims

Root functional traits are determinants of soil carbon storage; plant productivity; and ecosystem properties. However, few studies look at both annual and perennial roots, soil properties, and productivity in the context of field scale agricultural systems.

Methods

In Long Term and Conversion studies in North Central Kansas, USA; root biomass and length, soil carbon and nitrogen, microbial biomass, nematode and micro-arthropod communities were measured to a depth of one meter in paired perennial grassland and cropland wheat sites as well as a grassland site that had been converted to cropland using no tillage five years prior.

Results

In the Long Term Study root biomass was three to seven times greater (9.4 Mg ha^?1 and 2.5 Mg ha^?1 in May), and root length two times greater (52.5 km m^?2 and 24.0 km m^?2 in May) in perennial grassland than in cropland. Soil organic carbon and microbial biomass carbon were larger, numbers of Orbatid mites greater (2084 vs 730 mites m^?2), and nematode communities more structured (Structure Index 67 vs 59) in perennial grassland versus annual cropland. Improved soil physical and biological properties in perennial grasslands were significantly correlated with larger, deeper root systems. In the Conversion Study root length and biomass, microbial biomass carbon, mite abundance and nematode community structure differed at some but not all dates and depths. Isotope analysis showed that five years after no-till conversion old perennial roots remained in soils of annual wheat fields and that all soil fractions except coarse particulate organic matter were derived from C₄ plants.

Conclusions

Significant correlation between larger, longer roots in grasslands compared to annual croplands and improved soil biological, physical and chemical properties suggest that perennial roots are an important factor allowing perennial grasslands to maintain productivity and soil quality with few inputs. Perennial roots may persist and continue to influence soil properties long after conversion to annual systems.     

Foliar chemistry and tree ring δ13C of Pinus densiflora in relation to tree growth along a soil pH gradient

Background and aims

Soil acidification is known to be one of the constraints of tree growth; however, it is unclear how it affects tree growth at photosynthesis level (i.e., through affecting stomatal conductance vs. carboxylation rate) during the growth of trees. This paper studied the effects of soil acidification on Pinus densiflora foliar chemistry and tree ring C isotope ratio (¹³C/¹²C, expressed as δ¹³C) and their relationship with tree growth.

Methods

Tree growth (diameter, annual growth ring area, and root biomass), soil chemistry (pH, mineral N, and exchangeable Ca and Al), foliage chemistry (N, Ca/Al, and δ¹³C), and tree ring δ¹³C in P. densiflora stands along a soil pH gradient (from 4.38 to 4.83, n?=?9) in southern Korea were investigated.

Results

Overall, trees with relatively poor growth under more acidic soil conditions (low pH and Ca/Al) had lower values of foliar N concentration and δ¹³C and tree ring δ¹³C, suggesting that restricted N uptake under more acidic soil conditions caused N limitation for photosynthesis, leading to poor tree growth. In addition, relationships between mean annual area increment and carbon isotope discrimination of tree rings at five-yr intervals from 1968 to 2007 revealed that the impact of soil acidification on tree growth became severer during the last 15 yrs as negative correlations between them became significant after 1993.

Conclusions

Reduced N uptake under acidic soil conditions resulted in lower radial growth of P. densiflora via non-stomatal limitation of photosynthesis.     

Minor contribution of leaf litter to N nutrition of beech (Fagus sylvatica) seedlings in a mountainous beech forest of Southern Germany

Aims

Our aims were to characterize the fate of leaf-litter-derived nitrogen in the plant-soil-microbe system of a temperate beech forest of Southern Germany and to identify its importance for N nutrition of beech seedlings.

Methods

¹⁵N-labelled leaf litter was traced in situ into abiotic and biotic N pools in mineral soil as well as into beech seedlings and mycorrhizal root tips over three growing seasons.

Results

There was a rapid transfer of ¹⁵N into the mineral soil already 21 days after tracer application with soil microbial biomass initially representing the dominant litter-N sink. However, ¹⁵N recovery in non-extractable soil N pools strongly increased over time and subsequently became the dominant ¹⁵N sink. Recovery in plant biomass accounted for only 0.025 % of ¹⁵N excess after 876 days. After three growing seasons, ¹⁵N excess recovery was characterized by the following sequence: non-extractable soil N?>>?extractable soil N including microbial biomass?>>?plant biomass?>?ectomycorrhizal root tips.

Conclusions

After quick vertical dislocation and cycling through microbial N pools, there was a rapid stabilization of leaf-litter-derived N in non-extractable N pools of the mineral soil. Very low ¹⁵N recovery in beech seedlings suggests a high importance of other N sources such as root litter for N nutrition of beech understorey.     

The age of managed heathland communities: implications for carbon storage?

Background and aims

Shrublands are ecosystems vulnerable to climate changes, with key functions such as carbon storage likely to be affected. In dwarf shrublands dominated by Calluna vulgaris, the aboveground carbon allocation is associated with community age and phase of development. As the Calluna community grows older, a shift to net biomass loss occurs and it was hypothesized this would result in carbon stock increases within the soil.

Methods

The interaction of community age with ecosystem carbon stocks was investigated through a chronosequence study on three Calluna communities, aged 11, 18 and 27 years.

Results

Aboveground Calluna carbon stock increased significantly from the 11 year community (0.73 kg C m^?2) to the 18 year community (1.11 kg C m^?2) but did not significantly change from 18 to 27 years (1.0 kg C m^?2), indicating a net carbon gain that corresponded with the growth phase of the Calluna plants. Moss was also found to be a relatively large contributor to aboveground carbon stock (e.g. 30 % in the Young community). Moss has often been excluded in aboveground assessments on Calluna heathlands which may have led to previous stock underestimation. Belowground carbon stocks to 25 cm were six to nine times greater than in the aboveground pools. For example in the Young community, 8 % of the carbon stock was located aboveground, 35 % in the organic layer and 55 % in the mineral soil.

Conclusions

Increased heathland age resulted in increased aboveground carbon stock until peak production was reached at approximately 18 years of age. However, the proportionally large belowground carbon stock eclipsed any aboveground effect when total carbon stocks were considered. The investigation emphasized both the importance of including the mineral soil in sampling programs and of consider all major species, such as bryophytes, and vegetation age in carbon stock assessments.