Components,drivers and temporal dynamics of ecosystem respiration in a Mediterranean pine forest

To investigate the climate impacts on the different components of ecosystem respiration, we combined soil efflux data from a tree-girdling experiment with eddy covariance CO₂ fluxes in a Mediterranean maritime pine (Pinus pinaster) forest in Central Italy. 73 trees were stem girdled to stop the flux of photosynthates from the canopy to the roots, and weekly soil respiration surveys were carried out for one year. Heterotrophic respiration (R_H) was estimated from the soil CO₂ flux measured in girdled plots, and rhizosphere respiration (R_Ab) was calculated as the difference between respiration from controls (R_S) and girdled plots (R_H).Results show that the R_S dynamics were clearly driven by R_H (average R_H/R_S ratio 0.74). R_H predictably responded to environmental variables, being predominantly controlled by soil water availability during the hot and dry growing season (May–October) and by soil temperature during the wetter and colder months (November–March). High R_S and R_H peaks were recorded after rain pulses greater than 10 mm on dry soil, indicating that large soil carbon emissions were driven by the rapid microbial oxidation of labile carbon compounds. We also observed a time-lag of one week between water pulses and R_Ab peaks, which might be due to the delay in the translocation of recently assimilated photosynthates from the canopy to the root system. At the ecosystem scale, total autotrophic respiration (R_At, i.e. the sum of carbon respired by the rhizosphere and aboveground biomass) amounted to 60% of ecosystem respiration. R_At was predominantly controlled by photosynthesis, and showed high temperature sensitivity (Q₁₀) only during the wet periods. Despite the fact that the study coincided with an anomalous dry year and results might therefore not represent a general pattern, these data highlight the complex climatic control of the respiratory processes responsible for ecosystem CO₂ emissions.     

Net ecosystem exchange, evapotranspiration and canopy conductance in a riparian forest

The ecosystem fluxes of mass and energy were quantified for a riparian cottonwood (Populus fremontii S. Watson) stand, and the daily and seasonal courses of evapotranspiration, CO₂ flux, and canopy conductance were described, using eddy covariance. The ecosystem-level evapotranspiration results are consistent with those of other riparian studies; high vapor pressure deficit and increased groundwater depth resulted in reduced canopy conductance, and the annual cumulative evapotranspiration of 1095 mm was more than double the magnitude of precipitation. In addition, the cottonwood forest was a strong sink of CO₂, absorbing 310 g C m⁻² from the atmosphere in the first 365 days of the study. On weekly to annual time scales, hydrology was strongly linked with the net atmosphere-ecosystem exchange of CO₂, with ecosystem productivity greatest when groundwater depth was ∼2 m below the ground surface. Increases in groundwater depth beyond the depth of 2 m corresponded with decreased CO₂ uptake and evapotranspiration. Saturated soils caused by flooding and shallow groundwater depths also resulted in reduced ecosystem fluxes of CO₂ and water.     

Liming and reacidification reactions of a forest lake ecosystem,lake Lysevatten,in SW Sweden

Long-term monitoring, 1973 to 1987, of reactions to liming and reacidification of a forest lake ecosystem near the Swedish west coast is reported in this study. Treatment of Lake Lysevatten with a slag product of limestone in 1974 resulted in neutralization and a positive alkalinity. Prolonged dissolution proceeded for about 7 yr whereby 86% dissolved. During 1984–86 Lake Lysevatten approached maximum reacidification with high Al concentrations and an affected biota. Asellus aquaticus L. decreased and dominance within chironomid groups approached preliming conditions. However, the most obvious biological change was the development of the filamentous algal genus Mougeotia and increased growth of Sphagnum. Populations of both plants increased notably when pH declined to about 5. Our study suggests that extensive reacidification (pH < 6.0) of limed lakes should be avoided by successive treatments to prevent development of destabilized lake ecosystems.     

Assessing the uncertainty of estimated annual totals of net ecosystem productivity: A practical approach applied to a mid latitude temperate pine forest

Values for annual NEP of micrometeorological tower sites are usually published without an estimate of associated uncertainties. Few authors quantify total uncertainty of annual NEP. Moreover, different methods to assess total uncertainty are applied, usually addressing only one aspect of the uncertainty. This paper presents a robust and easy to apply method to quantify uncertainty of annual totals of Net Ecosystem Productivity (NEP), related to multiple factors involved therein. The method was applied to NEP observations for a Scots pine forest (Loobos) in the Netherlands. Total uncertainty of annual NEP for the Loobos site was on average ±32 g C m⁻² a⁻¹ (±8% of NEP), which is a quarter of the standard deviation of annual NEP (127 g C m⁻² a⁻¹).     

A geographically-based ecosystem model and its application to the carbon balance of the luquillo forest,Puerto Rico

We have developed a geography-based computer model of the Bisley Experimental Watershed ecosystem that simulates basic forest dynamics as a function of meteorological inputs and hydrologic simulation, as influenced by topography, soils, land use, and cover. The model is parameterized based on steady slate levels of, and hurricane impacts on, biomass, necromass and rate processes of the tabonuco forest, and is stable over decades. Over a 60 yr simulation (without hurricanes) leaf biomass remains approximately constant and woody biomass of those regions not having severe sunlight, moisture, or nutrient limitations increases slowly in agreement with observations. Necromass decreases slowly. Small quantities of C leave the ecosystem in stream water, especially during large rain events. When topographically-sensitive hurricane impacts are included, leaf and woody biomass are converted to necromass. In the model the recovery of the watersheds' hydrology, leaf and woody biomass, and necromass are consistent with field observations. We used this model to simulate the C dynamics of the forest over centurics using empirical values and found that (his forest acted to pump C from the atmosphere to the ocean at a rate of about 90 kg ha^?1 yr^?1.     

Bacterial numbers in a pine forest soil in relation to environmental factors

Numbers of soil bacteria stained with acridine orange (AO) or fluorescein diacetate (FDA) were studied for 3 and 2 yr respectively. Three pine forest sites were used and both organic and mineral soil layers were included. Different patterns of fluctuation in bacterial numbers were found each year. Significant correlations were demonstrated between AO-stained bacterial numbers and soil moisture content and between FDA-stained bacteria and the accumulated precipitation during a week before the sampling date. In multiple regression analyses 60–80% of the variation in numbers of AO-stained bacteria could be accounted for by soil moisture and numbers of bacterial-feeding nematodes at the sampling dates, and 30–45% of the variation in numbers of FDA-stained bacteria was accounted for by precipitation.     

Volatile and water-soluble inhibitors of nitrogen mineralization and nitrification in a ponderosa pine ecosystem

Summary Bioassay experiments were performed to test for inhibition of the processes of nitrogen mineralization and nitrification by organics in the forest floor of a ponderosa pine ecosystem. Water-extractable organics in the forest floor were tested by applying filtered extracts to the assay soil. The extract decreased nitrate production by 17.0% and decreased net mineralization by 4.1%. Inhibition by volatile organics was tested by placing vials containing forest floor or selected terpenoids of ponderosa pine in sealed jars containing the assay soil. Nitrate production was inhibited by 87.4% and 100%, and net nitrogen mineralization was inhibited by 73.3% and 67.7% in the jars with forest floor and terpenoids, respectively. Organics which are partially water-soluble and are volatile (such as terpenoids) would be very effective inhibitors of nitrogen cycling processes.     

An analysis of food-web structure and function in a shortgrass prairie,a mountain meadow,and a lodgepole pine forest

Summary The structure of the below-ground detrital food web was similar in three different semiarid vegetation types: lodgepole pine (Pinus contorta subsp. latifolia), mountain meadow (Agropyron smithii), and shortgrass prairie (Bouteloua gracilis). The densities of component food-web functional groups and the response to removal of component groups, differed however. As measured by biomass, bacteria were dominant in the meadow and prairie, while fungi were dominant in the forest. Resourde-base dominance was reflected in consumer dominance, and both directly correlated with the form of inorganic N present. Bacterial-feeding nematodes were numerically dominant in the meadow and prairie, while microarthropods were dominant in the forest. Ammonium-N was the dominant form in the forest, while nitrate —nitrite-N was the more important form in both bacterial-dominated grasslands.Addition of a biocide solution containing carbofuran and dimethoate reduced the numbers of both microarthropods and nematodes. In the bacterial-dominated grasslands, these reductions resulted in no apparent effect on bacterial densities because one group of bacterial consumers (protozoa) increased following the decrease in bacteria-feeding nematodes, in increased fungal biomass, and in increased soil inorganic N. Conversely, in the forest, following the biocide-induced reduction in consumers, the total fungal biomass decreased, but inorganic-N levels increased. The meadow appeared to be the most resilient of the three ecosystems to biocide disturbance, as both nematode and arthropod numbers returned to control levels more rapidly in the meadow than in the prairie or the forest.     

Carbon dioxide and energy fluxes from a boreal mixedwood forest ecosystem in Ontario, Canada

A long-term flux measurement station has been established in a 74-year-old mixedwood forest ecosystem, located approximately 80 km west of Timmins in northern Ontario, as part of the Fluxnet-Canada Research Network (FCRN). Measurements of energy, water vapour, and carbon dioxide fluxes have been made continuously since August 2003 using the eddy covariance technique, along with ancillary meteorological variables. The spatial structure of the site was evaluated using a variety of sources and techniques, including remote sensing, showing that this forest is mixed but relatively homogeneous. The canopy top height is remarkably constant at between 30 and 32 m. The basal area varies from 18 to 27 m² ha⁻¹, and the aboveground biomass ranges from 82 to 122 Mg ha⁻¹. In this paper, we summarize the diurnal and seasonal patters of carbon dioxide exchange and water loss from September 1, 2003 to August 31, 2004. Net ecosystem productivity (NEP) is strongly related to temperature. Atmospheric vapour pressure deficit (VPD) in this ecosystem exerted strong biophysical control on the daily gross ecosystem productivity (GEP) and evapotranspiration. Seasonal change in shortwave albedo, as a result of the presence of mixed deciduous and coniferous species, was clearly evident. Albedo changes were comparable to the seasonal pattern of NEP. The dormant season lasts more than 6 months of the year at this station. This forest was a moderate sink of carbon over the measurement period. Annual values of GEP, ecosystem respiration (R), and NEP were 1075, 919, and 156 ± 35 g C m⁻², respectively.     

Effects of land use on soil erosion in a tropical dry forest ecosystem,Chamela watershed,Mexico

During the past few decades, Mexico has been converting tropical dry forest (TDF) into cropland and pasture, with land degradation expressed as soil erosion being the main environmental consequence. The factors and processes influencing soil erosion are related to scale. At a microscale, the stability of soil aggregates has a significant impact on soil erodibility and strongly influences other soil properties. However, at plot and watershed scales, these relationships are less well known. The relationships between the distribution of soil aggregate size, soil properties and soil erosion were examined for two soil geomorphological units (hillslopes over granite and hillslopes over tuffs) and three land uses (TDF, unburned pasture and burned pasture) within the Chamela watershed of west–central Mexico. To evaluate soil aggregation as a parameter for upscaling soil erosion, the researchers measured microtopographic features at plot scales and interpreted 1:35,000 panchromatic aerial photographs at a watershed scale. Analysis of variance indicated significant differences in soil organic carbon (P < 0.05) and soil moisture (P < 0.01) contents between the two soil geomorphological units, and field tests showed differences in soil texture and structure.     

Element Inputs and Canopy Interactions in two Pine Forest Ecosystems in Berlin,Germany

For 3 years (1986-89), element fluxes with bulk deposition and throughfall were determined in a 40 yr. old pine plantation and a 80-140 yr. old mixed pine/oak stand in the Grunewald forest in Berlin. Although SO₄-inputs are very high (60 kg S·ha^?1.a^?1) due to the urban character of the study site, acid inputs are moderate because of the buffering action of airborne alkaline dusts. This is reflected in the high Ca-inputs (12 kg·ha^?1a^?1 in bulk precipitation). Acid neutralisation was also observed in the canopies and could be attributed to 60-80% to increased Ca- and Mg-fluxes in throughfall, stemming from foliage leaching and dust interception. Heavy metal inputs were low in comparison to other Central European study sites, the decrease in Pb-inputs over the 3-year period could be attributed to the increased use of unleaded gasoline since 1987. Total inputs and canopy leaching showed clear seasonal patterns for some elements: SO₄-, H-, Ca- and Mg-fluxes were higher in winter, on the other hand, K- and Mn-enrichment in throughfall showed two peaks, in early summer and late fall.     

Spatial variability of O layer thickness and humus forms under different pine beech–forest transformation stages in NE Germany

Spatial variability of humus layer (O layer) thicknesses can have important impacts upon soil water dynamics, nutrient storage and availability, as well as plant growth. The purpose of the present study was to elucidate the impact of forest‐transformation practices on the spatial variability of O layer thicknesses. The study focused on the Kahlenberg forest area (NE Germany) with stands of Scots pine (Pinus sylvestris) and European beech (Fagus sylvatica) of different age structures that form a transformation chronosequence from pure Scots pine stands towards pure European beech stands. Topsoil profiles including both, the O layer and the uppermost humic mineral soil horizon were excavated at intervals of 0.4 m along 15–20 m long transects, and spatial variability of O layer thicknesses was quantified by variogram analysis. The correlation lengths of total O layer thickness increased in the sequence consisting of pure pine stand (3.1 m) → older mixed stand (3.7 m) → pure beech stand (4.5 m), with the exception of the younger mixed stand, for which no correlation lengths of total O layer thickness could be determined. The degree of spatial correlation, i.e., the percentage of the total variance which can be described by variograms, was highest for the two monospecies stands, whereas this percentage was distinctly lower for the two mixed stands. A similar minimum for the two mixed stands was observed for the correlation lengths of the Oh horizon. These results suggest that the spatial structures of forest‐transformation stands may be interpreted in terms of a disturbance (in the form of the underplanting of beech trees). After this disturbance, the forest ecosystem requires at least 100 y to again reach relative equilibrium. These findings are in line with the results of other soil‐related investigations at these sites.     

Dynamics and stratification of Enchytraeidae in the organic layer of a Scots pine forest

In a Dutch Scots pine forest an experiment was conducted to quantify the role of soil biota in the functioning of the soil ecosystem, and the effects of enhanced nitrogen deposition. For this, the site was sampled at 8-week intervals during 2.5 years. This paper reports on the population dynamics of enchytraeids in the field and in stratified litterbags. Mean yearly abundance of the enchytraeid community in the field was 47 600 m^–2, or 0.70 g (dry weight) m^–2. The community consisted mainly of three species: Cognettia sphagnetorum, Marionina clavata and Achaeta eiseni, of which C. sphagnetorum was dominant. The enchytraeid populations showed a marked stratification in the same sequence. Freshly fallen pine needles were colonized by C. sphagnetorum, while other species followed much later. It was found that data from the litterbags were reasonably comparable with field data, when expressed per gram of dry substrate, but less so when expressed per square metre. Multiple regression analysis of the data showed that the population dynamics in the litter layer could largely be explained by temperature and moisture fluctuations; in deeper layers other factors, such as the stage of decomposition, were probably more important. Received: 26 June 1997     

Dynamics and stratification of protozoa in the organic layer of a Scots pine forest

Fluctuations in soil biota abundance in different organic layers of a Scots pine forest in The Netherlands were studied by bimonthly counts during 2.5 years. The counts were made using litterbags which were placed in the litter (L), fragmentation (F) and humus (H) layers at the start of the experiment. Results from the L layer were also compared with results from litter which was renewed every 2 months (L′) to study colonisation. In this study the results for amoebae, flagellates and ciliates are presented. The highest numbers of soil protozoa were found in the L layer during most sampling occasions. The H layer contained the lowest numbers. The L layer also showed higher numbers than the L′ litterbags which were renewed every 2 months. Fluctuations in abundance could partly be explained by fluctuations in moisture content. Moisture content in the litterbags was rather constant throughout the experiment, although occasionally moisture contents of 10% and 80% were observed. Fluctuations in moisture content in the L layer were often larger than in the F and H layers. Flagellates were the most abundant group, reaching numbers of several hundred thousands to several millions per gram fresh weight on various occasions. Amoebae often reached numbers of between tens of thousands and several hundred thousands. Ciliates only reached numbers of up to several thousands. Received: 26 June 1997     

Watershed base-cation cycle dynamics modeled over forest regrowth in a Central Appalachian ecosystem

Watershed ecosystem analysis has been used to study aspects of nutrient cycles in many regions of the US. Here we quantify watershed input-output budgets and intrasystem cycling of the base cations Ca, K and Mg in a montane Virginia ecosystem. The intrasystem fluxes of uptake, return, canopy leaching and mineralization were simulated over the period of forest aggradation. A forest-dynamics model, based on previous models, was created to model biotically-driven fluxes at this site; biomass nutrient concentrations were parameterized with a field study. A two-year watershed mass-balance study was then conducted to estimate geologic fluxes for comparison with modeled biotic fluxes. Results show the major biotic fluxes to be much greater, highlighting the importance of considering biomass dynamics in ecosystem nutrient-cycling studies. Mineralization from forest-floor biomass compartments proved to be an increasingly important avenue for internal recycling during aggradation. Accumulation of base cations in biomass also corresponded to a production of H⁺ in soil at three times the H⁺ levels in atmospheric deposition at this location. Such high levels of base removal in soils could exceed weathering rates and may result in a depletion of bases from the soil exchange complex.     

Rapid soil organic matter loss from forest dieback in a subalpine coniferous ecosystem

Forest dieback caused by climate-change associated stresses and insect outbreaks has emerged as a global concern, and the biogeochemical consequences of this phenomenon need to be elucidated. We measured biological and chemical traits of soil beneath live trees or trees recently killed by a mountain-pine-beetle outbreak in a subalpine coniferous forest in the Front Range of Colorado. We focused on the top 5 cm of mineral soil just beneath the O horizon and measured microbial biomass, soil invertebrate abundance and composition, and soil chemical characteristics. With the termination of inputs from rhizodeposition, mycorrhizal fungal turnover and fine root turnover, soil total carbon (C) and total nitrogen (N) in the mineral soil at three sites decreased by 38–49% and 26–45%, respectively. Tree mortality was associated with reduced soil microbial biomass but soil nematode and microarthropod densities were unchanged. Nematode trophic structure was altered with an increased proportion of bacterial feeders. Soil inorganic N concentrations were inversely correlated to microbial C:N ratios. Tree death was associated with increased soil pH, a possible loss of calcium (Ca²⁺), but an accumulation of soil inorganic N, largely as NH₄⁺. Our results suggest that forest dieback results in rapid C and N loss from surface mineral soils and that the accumulation of soil inorganic N, the reduction in microbial biomass, and the more bacterial-based soil food web increase the potential of enhanced N loss from affected ecosystems.     

Landscape variation in microarthropod response to calcium addition in a northern hardwood forest ecosystem

Loss of base cations from soils of the northeastern US may adversely influence forest health. A watershed-level calcium (as wollastonite, CaSiO₃) addition experiment was initiated at the Hubbard Brook Experimental Forest, NH, USA, in fall 1999 to examine responses of forest ecosystem structure and function to restoration of exchangeable Ca that was lost over previous decades. We quantified the response of soil microarthropods to Ca addition as part of this larger study, and we also quantified microarthropod response to smaller scale (1×2 m plots) wollastonite additions within one site in an untreated reference watershed. We observed a small increase in pH in the surface organic horizon, accompanied by a decline in microarthropod abundance per gram organic matter three years following wollastonite application. However, responses varied among microarthropod groups, across years, and among study plots that were distributed along an elevation gradient in treatment and reference watersheds. Collembola declined in the treatment relative to the reference watershed in the second year but recovered in the third year following wollastonite application, whereas oribatid, prostigmatid, and mesostigmatid mites all declined in the treatment relative to the reference watershed by the third year following treatment. The largest treatment responses were observed in two base-poor hardwood stands, whereas at a slightly less poor hardwood site we found the least abundant and least variable microarthropod populations, and no response was observed to either the watershed or the plot-level treatments. Our results demonstrate that soil microarthropods are sensitive to relatively small changes in soil Ca, but that landscape position must be considered in testing responses of these soil biota to soil base status.     

Annual and long-term fluctuations of the nematode fauna in a Swedish Scots pine forest soil

Samples from an old Scots pine forest at Ivantjärnsheden in the middle of Sweden were used to study predictability and patterns of variation of soil nematode communities. There were two annual sampling series (1974–75 and 1977–78) and one long-term series sampled in September ten times over a period of 25 years. The abundance and the composition of the fauna fluctuated rather considerably in both the annual and long-term series. In the annual series abundance and species composition varied in a way which can partly be explained by changes in temperature and moisture. Total nematode abundance was influenced by soil water contents as indicated by co-variations with precipitation. Although the variations in abundance and fauna composition were large no systematic changes could be detected during 25 years. The differences in faunal structure between the two annual series were greater than between the annual and the long-term series.In all series there was a distinct vertical stratification of the fauna. In the superficial moss and litter layers species belonging to Adenophorea (Plectus) dominated. In deeper layers members of Rhabditida (Acrobeloides) contributed a greater proportion of the fauna. Variations of the annual series indicate that coexistence of different nematode species is facilitated by differences in response to temperature and moisture. The abundance of fungal and bacterial feeders changed in a regular way. During the summer the proportions of fungal and bacterial feeders were almost equal, but during the wet and cold winter the proportion of bacterial feeders increased. Rapidly growing bacterial feeding species belonging to Rhabditida were common in late summer and early autumn, whereas the more slowly growing bacterial feeders belonging to Adenophorea were most abundant during the winter. Although the community fluctuated rather much the average values indicated a rather high degree of predictability and also a high similarity with nematode faunas of other pine forest soils.     

Response of soil water chemistry and fine-roots to clean rain in a spruce forest ecosystem at Solling,FRG

In the Solling experimental forest in central Germany a ‘clean rain’ roof experiment is conducted in a 60 year old Norway Spruce (Picea abies KARST.) stand. In this experiment with application of artificially prepared pre-industrial throughfall there is now a time series of soil water chemistry data from about 2 yr of pre-experiment and 3.5 yr of manipulation treatment. The response of soil solution chemistry to reduced inputs of N and S was strong and fairly rapid. There is a clear reflection of reduced input in soil solution concentrations, particularly for the N ions. The fine-roots of the Norway spruce trees reacted strongly to these changes in soil water chemistry. Fine-root biomass increased in the clean rain plot by about 40 % compared to pre-experimental conditions. This increase was strongest in the B-horizon, indicating that acid stress has ameliorated in the mineral soil. However, low concentrations of ammonium and nitrate in the root zone may also have contributed to this effect, since more fineroots are needed to maintain the N demand of the trees. No effect was yet found for other variables (photosynthesis, respiration, transpiration). Nutrient cation concentrations in the needles remained on the same low level as in the control groups. However, these aboveground variables may react after some time lag. The results demonstrate that in spruce forests on acid soil atmospheric element input largely controls soil solution chemistry and that air pollution control measures would have a significant effect with respect to ameliorating soil water chemistry, acidity and forest health.