Influence of Earthworm Invasion on Redistribution and Retention of Soil Carbon and Nitrogen in Northern Temperate Forests

We analyzed soil organic matter distribution and soil solution chemistry in plots with and without earthworms at two sugar maple (Acer saccharum)–dominated forests in New York State, USA, with differing land-use histories to assess the influence of earthworm invasion on the retention or loss of soil carbon (C) and nitrogen (N) in northern temperate forests. Our objectives were to assess the influence of exotic earthworm invasion on (a) the amount and depth distribution of soil C and N, (b) soil ¹³C and ¹⁵N, and (c) soil solution chemistry and leaching of C and N in forests with different land-use histories. At a relatively undisturbed forest site (Arnot Forest), earthworms eliminated the thick forest floor, decreased soil C storage in the upper 12 cm by 28%, and reduced soil C:N ratios from 19.2 to 15.3. At a previously cultivated forest site with little forest floor (Tompkins Farm), earthworms did not influence the storage of soil C or N or soil C:N ratios. Earthworms altered the stable isotopic signature of soil at Arnot Forest but not at Tompkins Farm; the alteration of stable isotopes indicated that earthworms significantly increased the loss of forest floor C but not N from the soil profile at Arnot Forest. Nitrate (NO₃^–) concentrations in tension and zero-tension lysimeters were much greater at Tompkins Farm than Arnot Forest, and earthworms increased NO₃^– leaching at Tompkins Farm. The results suggest that the effect of earthworm invasion on the distribution, retention, and solution chemistry of soil C and N in northern temperate forests may depend on the initial quantity and quality of soil organic matter at invaded sites.     

Fine-root mass, growth and nitrogen content for six tropical tree species

Although fine roots might account for 50% of the annual net primary productivity in moist tropical forests, there are relatively few studies of fine-root dynamics in this biome. We examined fine-root distributions, mass, growth and tissue N and C concentrations for six tree species established in 16-year-old plantations in the Caribbean lowlands of Costa Rica in a randomized-block design (n = 4). The study included five native species (Hyeronima alchorneoides, Pentaclethra macroloba, Virola koschnyi, Vochysia ferruginea and Vochysia guatemalensis) and one exotic (Pinus patula). Under all species >60% of the total fine-root mass to 1 m deep was located in the uppermost 15 cm of the soil. Fine-root live biomass and necromass (i.e., the mass of dead fine-roots) varied significantly among species but only within the uppermost 15 cm, with biomass values ranging from 182 g m⁻² in Pinus to 433 g m⁻² in Hyeronima plots, and necromass ranging from 48 g m⁻² in Pinus to 183 g m⁻² in Virola plots. Root growth, measured using ingrowth cores, differed significantly among species, ranging from 304 g m⁻² year⁻¹ in Pinus to 1,308 g m⁻² year⁻¹ in Hyeronima. These growth rates were one to five times those reported for moist temperate areas. Turnover rates of fine-root biomass ranged from 1.6 to 3.0 year⁻¹ in Virola and Hyeronima plots, respectively. Fine-root biomass was significantly and positively correlated with fine-root growth (r = 0.79, P < 0.0001), but did not correlate with fine-root turnover (r = 0.10, P = 0.20), suggesting that fine-root accumulation is a function of growth rate rather than mortality. Fine-root longevity was not correlated (r = 0.20, P = 0.34) and growth was negatively correlated with root N concentration across species (r = −0.78, P < 0.0001), contrary to reported trends for leaves, perhaps because N was relatively abundant at this site.     

Fine root production and nutrient content in wet and moist arctic tundras as influenced by chronic fertilization

We used ingrowth cores to estimate fine root production in organic soils of wet sedge and moist tundra ecosystems near Toolik Lake on Alaska's North Slope. Root-free soil cores contained in nylon mesh tubes (5 cm diameter, 20–30 cm long) were placed in control and chronically fertilized (N plus P) plots in mid-August 1994 and were retrieved 1 year later. Estimated fine root production in control plots was 75 g m^–2 year^–1 in wet sedge and 56 g m^–2 year^–1 in moist tussock tundra. Fine root production in fertilized plots was 85 g m^–2 year^–1 in wet sedge and 67 g m^–2 year^–1 in moist tussock tundra. Although our estimates of fine root production were higher on fertilized than control plots, differences were not statistically significant within either tundra type. Comparisons between our estimates of fine root production and other estimates of aboveground (plus rhizome) production on the same (wet sedge tundra) or similar (moist tussock tundra) plots suggest that fine root production was about one-third of total net primary production (NPP) under non-fertilized conditions and about one-fifth of total NPP under chronic fertilization. Fine root N and P concentrations increased with fertilization in both tundra types, but P concentrations increased more than N concentrations in wet sedge tundra, whereas relative increases in N and P concentrations in moist tundra roots were similar. These data are consistent with other studies suggesting that NPP in wet sedge tundra is often P limited and that co-limitation by N and P is more important in moist tussock tundra.     

CO2 and N-fertilization effects on fine-root length, production, and mortality: a 4-year ponderosa pine study

We conducted a 4-year study of juvenile Pinus ponderosa fine root (≤2 mm) responses to atmospheric CO₂ and N-fertilization. Seedlings were grown in open-top chambers at three CO₂ levels (ambient, ambient+175 μmol/mol, ambient+350 μmol/mol) and three N-fertilization levels (0, 10, 20 g m⁻² year⁻¹). Length and width of individual roots were measured from minirhizotron video images bimonthly over 4 years starting when the seedlings were 1.5 years old. Neither CO₂ nor N-fertilization treatments affected the seasonal patterns of root production or mortality. Yearly values of fine-root length standing crop (m m⁻²), production (m m⁻² year⁻¹), and mortality (m m⁻² year⁻¹) were consistently higher in elevated CO₂ treatments throughout the study, except for mortality in the first year; however, the only statistically significant CO₂ effects were in the fine-root length standing crop (m m⁻²) in the second and third years, and production and mortality (m m⁻² year⁻¹) in the third year. Higher mortality (m m⁻² year⁻¹) in elevated CO₂ was due to greater standing crop rather than shorter life span, as fine roots lived longer in elevated CO₂. No significant N effects were noted for annual cumulative production, cumulative mortality, or mean standing crop. N availability did not significantly affect responses of fine-root standing crop, production, or mortality to elevated CO₂. Multi-year studies at all life stages of trees are important to characterize belowground responses to factors such as atmospheric CO₂ and N-fertilization. This study showed the potential for juvenile ponderosa pine to increase fine-root C pools and C fluxes through root mortality in response to elevated CO₂.     

Fine-root seasonal pattern,production and turnover rate of European beech (Fagus sylvatica L.) stands in Italy Prealps: Possible implications of coppice conversion to high forest

Abstract

The aim of this study was to investigate the possible effects of coppice conversion to high forest on the beech fine-root systems. We compared the seasonal pattern of live and dead fine-root mass (d < 2 mm), production and turnover in three beech stands that differed in management practices. Tree density was higher in the 40-year-old coppice stand than in the stands that were converted from coppice to high forest in 1994 and 2004, respectively. We found that a reduction in tree density reduced the total fine-root biomass (Coppice stand, 353.8 g m^?2; Conversion 1994 stand, 203.6 g m^?2; Conversion 2004 stand, 176.2 g m^?2) which continued to be characterised by a bimodal pattern with two major peaks, one in spring and one in early fall. Conversion to high forest may also affect the fine-root soil depth distribution. Both fine-root production and turnover rate were sensitive to management practices. They were lower in the Coppice stand (production 131.5 g m^?2 year^?1; turnover rate 0.41 year^?1) than in the converted stands (1994 Conversion stand: production 232 g m^?2 year^?1, turnover rate 1.06 year^?1; 2004 Conversion stand: production 164.2 g m^?2 year^?1, turnover rate 0.79 year^?1).     

Contribution of aboveground litter,belowground litter,and rhizosphere respiration to total soil CO<Subscript>2</Subscript> efflux in an old growth coniferous forest

In an old growth coniferous forest located in the central Cascade Mountains, Oregon, we added or removed aboveground litter and terminated live root activity by trenching to determine sources of soil respiration. Annual soil efflux from control plots ranged from 727 g C m⁻² year⁻¹ in 2002 to 841 g C m⁻² year⁻¹ in 2003. We used aboveground litter inputs (149.6 g C m⁻² year⁻¹) and differences in soil CO₂ effluxes among treatment plots to calculate contributions to total soil efflux by roots and associated rhizosphere organisms and by heterotrophic decomposition of organic matter derived from aboveground and belowground litter. On average, root and rhizospheric respiration (R_r) contributed 23%, aboveground litter decomposition contributed 19%, and belowground litter decomposition contributed 58% to total soil CO₂ efflux, respectively. These values fall within the range of values reported elsewhere, although our estimate of belowground litter contribution is higher than many published estimates, which we argue is a reflection of the high degree of mycorrhizal association and low nutrient status of this ecosystem. Additionally, we found that measured fluxes from plots with doubled needle litter led to an additional 186 g C m⁻² year⁻¹ beyond that expected based on the amount of additional carbon added; this represents a priming effect of 187%, or a 34% increase in the total carbon flux from the plots. This finding has strong implications for soil C storage, showing that it is inaccurate to assume that increases in net primary productivity will translate simply and directly into additional belowground storage.     

Assessing fine-root biomass and production in a Scots pine stand – comparison of soil core and root ingrowth core methods

Soil core and root ingrowth core methods for assessing fine-root (< 2 mm) biomass and production were compared in a 38-year-old Scots pine (Pinus sylvestris L) stand in eastern Finland. 140 soil cores and 114 ingrowth cores were taken from two mineral soil layers (0–10 cm and 10–30 cm) during 1985–1988. Seasonal changes in root biomass (including both Scots pine and understorey roots) and necromass were used for calculating fine-root production. The Scots pine fine-root biomass averaged annually 143 g/m² and 217 g/m² in the upper mineral soil layer, and 118 g/m² and 66 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The fine-root necromass averaged annually 601 g/m² and 311 g/m² in the upper mineral soil layer, and 196 g/m² and 159 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The annual fine-root production in a Scots pine stand in the 30 cm thick mineral soil layer, varied between 370–1630 g/m² in soil cores and between 210 – 490 g/m² in ingrowth cores during three years. The annual production calculated for Scots pine fine roots, varied between 330–950 g/m² in soil cores and between 110 – 610 g/m² in ingrowth cores. The horizontal and vertical variation in fine-root biomass was smaller in soil cores than in ingrowth cores. Roots in soil cores were in the natural dynamic state, while the roots in the ingrowth cores were still expanding both horizontally and vertically. The annual production of fine-root biomass in the Scots pine stand was less in root ingrowth cores than in soil cores. During the third year, the fine-root biomass production of Scots pine, when calculated by the ingrowth core method, was similar to that calculated by the soil core method. Both techniques have sources of error. In this research the sampling interval in the soil core method was 6–8 weeks, and thus root growth and death between sampling dates could not be accurately estimated. In the ingrowth core method, fine roots were still growing into the mesh bags. In Finnish conditions, after more than three growing seasons, roots in the ingrowth cores can be compared with those in the surrounding soil. The soil core method can be used for studying both the annual and seasonal biomass variations. For estimation of production, sampling should be done at short intervals. The ingrowth core method is more suitable for estimating the potential of annual fine-root production between different site types.     

Methane and nitrous oxide exchange in differently fertilised grassland in southern Germany

We examined the effect of fertilisation (200 kg cattle slurry-N ha^–1 year^–1) on the exchange of N₂O and CH₄ in the soil–plant system of meadow agroecosystems in southern Germany. From 1996 to 1998, we regularly determined the gas fluxes (closed chamber method) and associated environmental parameters. N₂O and CH₄ fluxes were not significantly affected by fertilisation. N₂O fluxes at the unfertilised and fertilised plots were small, generally between 50 and –20 g N m^–2 h^–1. We identified some incidents of N₂O uptake. CH₄-C fluxes ranged from 1.3 to –0.2 mg m^–2 h^–1 and were not significantly different from 0 at both plots. We budgeted an annual net emission of 15.5 and 29.6 mg m^–2 N₂O-N and an annual CH₄-C net emission of 184.2 and 122.7 mg m^–2 at the unfertilised and fertilised plots, respectively. Apparently, rapid N mineralization and uptake in the densely rooted topsoil prevents N losses and the inhibition of CH₄ oxidation.     

Earthworm secondary production and N flux in agroecosystems: a comparison of two approaches

Production was estimated for Aporrectodea spp. and Lumbricus spp. populations in corn agroecosystems with a 5-year history of manure or inorganic fertilizer applications during 1994–1995 and 1995–1996. Earthworm biomass and production were greater in manure than inorganic fertilizer plots, although biomass and production declined by about 50% between 1994–1995 and 1995–1996 due to unfavorable climatic conditions. Production was highest during the spring and autumn when soil temperatures were between 4 and 22°C. Production was higher in Lumbricus spp. than Aporrectodea spp. populations due to greater Lumbricus spp. biomass. Aporrectodea spp. production was 3.47–16.14 g ash-free dry weight (AFDW) m^–2 year^–1, while Lumbricus spp. production was 6.09–18.11 g AFDW m^–2 year^–1, depending on the fertilizer treatment and the method used to estimate production. However, production estimates from the instantaneous growth rate method were within 27% of the values calculated using the size-frequency method. Nitrogen flux through earthworms was used to estimate efficiency quotients. Net production efficiency (P/A) ranged from 0.64 to 0.76, assimilation efficiency (A/C) ranged from 0.1 to 0.3, and gross production efficiency (P/C) ranged from 0.06 to 0.22. Annual N flux through earthworm populations was higher in manure than inorganic fertilizer plots, and ranged from 2.95 to 5.47 g N m^–2 year^–1 in 1994–1995 and 1.76 to 2.92 g N m^–2 year^–1 in 1995–1996. The N flux through earthworms represented an amount equivalent to 16–30% of crop N uptake during 1994–1995 and 11–18% of crop N uptake during 1995–1996. We concluded that the effects of earthworms on N cycling in corn agroecosystems were substantial, and that N flux through earthworms was influenced significantly by fertilizer amendments. Received: 20 September 1999 / Accepted: 24 March 2000     

A historic perspective on Wadden Sea eutrophication

In this paper, a reconstruction of the pre-industrial trophic status of the Wadden Sea is presented. A conceptual model is outlined that links the organic matter and nutrient dynamics in the Wadden Sea with riverine nutrient input. Fundamental processes in this model are: a nutrient-limited offshore primary production and the subsequent import of primary produced organic matter from the North Sea into the Wadden Sea. Two approaches have been followed to estimate the production and remineralisation levels under pre-industrial conditions. The first approach is based on present-day relationships between the seasonal cycle of NH₄ and NO₂ in the western Dutch Wadden Sea and suggests, on average, sixfold lower production and remineralisation rates under pre-industrial conditions (range: four to eight times). The second approach is based on present carbon budgets extrapolated to pre-industrial budgets on the basis of present relationships between winter nutrient concentrations, annual primary production and annual organic matter turnover rates, and suggests a fivefold lower organic matter turnover under pre-industrial conditions (annual primary production: ~55 g C m^–2 year^–1, annual remineralisation: ~77 g C m^–2 year^–1). Better pre-industrial light conditions in the Wadden Sea may have allowed a more efficient use of nutrients, a higher annual primary production of about 86 g C m^–2 year^–1 and annual remineralisation rates of about 108 g C m^–2 year^–1.     

Estimation of aboveground biomass and net biomass increment in a cool temperate forest on a landscape scale

I clarified aboveground biomass (AGB), net biomass increment (NBI) and its spatial heterogeneity in a cool temperate forest on a landscape scale (>2,200 ha). The relationships among AGB, NBI, and the size frequency distribution of trees of each stand were examined by combining an analysis of vegetation using aerial photographs, and data from 146 inventory plots (28.8 ha in total). This area included natural broad-leaved stands, harvested broad-leaved stands, and artificial conifer plantations. A –3/2 power distribution density function was applied to the individual mass frequency distribution of each plot. Estimated AGB in carbon (C) equivalent was 480–5,615 g C m^–2 (3,130 g C m^–2 on average), and NBI was –98 to 436 g C m^–2 year^–1 (83.0 g C m^–2 year^–1 on average). NBI had a single significant relationship with the reciprocal of theoretical maximum individual mass, while NBI was not significantly related to AGB. My results showed that, on a landscape scale, AGB and NBI strongly depend on the size structure of forest stands.     

Anthropogenic N deposition and the fate of 15NO 3 − in a northern hardwood ecosystem

Human activity has substantially increased atmospheric NO ₃ ^– deposition in many regions of the Earth, which could lead to the N saturation of terrestrial ecosystems. Sugar maple (Acer saccharum Marsh.) dominated northern hardwood forests in the Upper Great Lakes region may be particularly sensitive to chronic NO ₃ ^– deposition, because relatively moderate experimental increases (three times ambient) have resulted in substantial N leaching over a relatively short duration (5–7 years). Although microbial immobilization is an initial sink (i.e., within 1–2 days) for anthropogenic NO ₃ ^– in this ecosystem, we have an incomplete understanding of the processes controlling the longer-term (i.e., after 1 year) retention and flow of anthropogenic N. Our objectives were to determine: (i) whether chronic NO ₃ ^– additions have altered the N content of major ecosystem pools, and (ii) the longer-term fate of ¹⁵NO ₃ ^– in plots receiving chronic NO ₃ ^– addition. We addressed these objectives using a field experiment in which three northern hardwood plots receive ambient atmospheric N deposition (ca. 0.9 g N m^–2 year^–1) and three plots which receive ambient plus experimental N deposition (3.0 g NO₃ ^–-N m^–2 year^–1). Chronic NO ₃ ^– deposition significantly increased the N concentration and content (g N/m²) of canopy leaves, which contained 72% more N than the control treatment. However, chronic NO ₃ ^– deposition did not significantly alter the biomass, N concentration or N content of any other ecosystem pool. The largest portion of ¹⁵N recovered after 1 year occurred in overstory leaves and branches (10%). In contrast, we recovered virtually none of the isotope in soil organic matter (SOM), indicating that SOM was not a sink for anthropogenic NO ₃ ^– over a 1 year duration. Our results indicate that anthropogenic NO ₃ ^– initially assimilated by the microbial community is released into soil solution where it is subsequently taken up by overstory trees and allocated to the canopy. Anthropogenic N appears to be incorporated into SOM only after it is returned to the forest floor and soil via leaf litter fall. Short- and long-term isotope tracing studies provided very different results and illustrate the need to understand the physiological processes controlling the flow of anthropogenic N in terrestrial ecosystems and the specific time steps over which they operate.     

Carbon-Degrading Enzyme Activities Stimulated by Increased Nutrient Availability in Arctic Tundra Soils

Climate-induced warming of the Arctic tundra is expected to increase nutrient availability to soil microbes, which in turn may accelerate soil organic matter (SOM) decomposition. We increased nutrient availability via fertilization to investigate the microbial response via soil enzyme activities. Specifically, we measured potential activities of seven enzymes at four temperatures in three soil profiles (organic, organic/mineral interface, and mineral) from untreated native soils and from soils which had been fertilized with nitrogen (N) and phosphorus (P) since 1989 (23 years) and 2006 (six years). Fertilized plots within the 1989 site received annual additions of 10 g N⋅m^-2⋅year^-1 and 5 g P⋅m^-2⋅year^-1. Within the 2006 site, two fertilizer regimes were established – one in which plots received 5 g N⋅m^-2⋅year^-1 and 2.5 g P⋅m^-2⋅year^-1 and one in which plots received 10 g N⋅m^-2⋅year^-1 and 5 g P⋅m^-2⋅year^-1. The fertilization treatments increased activities of enzymes hydrolyzing carbon (C)-rich compounds but decreased phosphatase activities, especially in the organic soils. Activities of two enzymes that degrade N-rich compounds were not affected by the fertilization treatments. The fertilization treatments increased ratios of enzyme activities degrading C-rich compounds to those for N-rich compounds or phosphate, which could lead to changes in SOM chemistry over the long term and to losses of soil C. Accelerated SOM decomposition caused by increased nutrient availability could significantly offset predicted increased C fixation via stimulated net primary productivity in Arctic tundra ecosystems.     

Earthworm numbers,biomass and respiratory metabolism in a beech woodland-Wytham Woods,Oxford

Summary The mean annual population density of earthworms was found to be 164.6 m^-2 during a period of detailed study between October 1971 and September 1972. In a year of less detailed study between November 1972 and October 1973 the population density was 117.5 m^-2 (139.8 m^-2 when the type of extraction method was allowed for). Mean biomass densities in the two years of investigation were 41.0 g preserved wet wt m^-2 (1971–1972) and 38.6 (possibly 39.2) g preserved wet wt m^-2 (1972–1973).Comparison of the Brogden's Belt population and biomass densities with those reported from other woodlands indicates that soil type is more important than leaf litter type in determining the numerical abundance of earthworms. Population densities are lower in beechwoods on mor soils, mor soils also support significantly fewer species. As with numbers, mean biomass density in beechwoods on mor soils was significantly lower than that occurring in beechwoods on mull soils; the latter, in turn, being lower than those found in mixed deciduous woods on mull soils. Unlike population density biomass density is influenced by both soil and litter type, this is discussed by reference to mean body weights and food quality as reflected by tannin, nitrogen and carbohydrate content.The annual respiratory metabolism of the Brogden's Belt earthworms was calculated to be between 10.7 and 13.41 O₂ m^-2 a^-1, which is equivalent to between 4.1 and 5.1% of the total soil metabolism. A production/biomass ratio of 0.49–0.58 was estimated, as was a net population efficiency of 22%.     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Denitrification and N2O emission from urine-affected grassland soil

Denitrification and N₂O emission rates were measured following two applications of artificial urine (40 g urine-N m^–2) to a perennial rye-grass sward on sandy soil. To distinguish between N₂O emission from denitrification or nitrification, urine was also applied with a nitrification inhibitor (dicyandiamide, DCD). During a 14 day period following each application, the soil was frequently sampled, and incubated with and without acetylene to measure denitrification and N₂O emission rates, respectively.Urine application significantly increased denitrification and N₂O emission rates up to 14 days after application, with rates amounting to 0.9 and 0.6 g N m^–2 day^–1 (9 and 6 kg N ha^–1 day^–1), respectively. When DCD was added to the urine, N₂O emission rates were significantly lower from 3 to 7 days after urine application onwards. Denitrification was the main source of N₂O immediately following each urine application. 14 days after the first application, when soil water contents dropped to 15% (v/v) N₂O mainly derived from nitrification.Total denitrification losses during the 14 day periods were 7 g N m^–2, or 18% of the urine-N applied. Total N₂O emission losses were 6.5 and 3 g N m^–2, or 16% and 8% of the urine-N applied for the two periods. The minimum estimations of denitrification and N₂O emission losses from urine-affected soil were 45 to 55 kg N ha^–1 year^–1, and 20 to 50 kg N ha^–1 year^–1, respectively.     

Activity of surface-casting earthworms in a calcareous grassland under elevated atmospheric CO2

Earthworms make up the dominant fraction of the biomass of soil animals in most temperate grasslands and have important effects on the structure and function of these ecosystems. We hypothesized that the effects of elevated atmospheric CO₂ on soil moisture and plant biomass production would increase earthworm activity, expressed as surface cast production. Using a screen-aided CO₂ control facility (open top and open bottom rings), eight 1.2-m² grassland plots in Switzerland have been maintained since March 1994 at ambient CO₂ concentrations (350 μl CO₂ l⁻¹) and eight at elevated CO₂ (610 μl CO₂ l⁻¹). Cumulative earthworm surface cast production measured 40 times over 1 year (April 1995–April 1996) in plots treated with elevated CO₂ (2206 g dry mass m⁻² year⁻¹) was 35% greater (P<0.05) than that measured in plant communities maintained at ambient CO₂ (1633 g dry mass m⁻² year⁻¹). At these rates of surface cast production, worms would require about 100 years to egest the equivalent of the amount of soil now found in the Ah horizon (top 15 cm) under current ambient CO₂ concentrations, and 75 years under elevated CO₂. Elevated atmospheric CO₂ had no influence on the seasonality of earthworm activity. Cumulative surface cast production measured over the 7-week period immediately following the 6-week summer dry period in 1995 (no surface casting) was positively correlated (P<0.05) with the mean soil water content calculated over this dry and subsequent wetter period, when viewed across all treatments. However, no correlations were observed with soil temperature or with annual aboveground plant biomass productivity. No CO₂-related differences were observed in total nitrogen (N_tot) and organic carbon (C_org) concentration of surface casts, although concentrations of both elements varied seasonally. The CO₂-induced increase in earthworm surface casting activity corresponded to a 30% increase of the amount of N_tot (8.9 mg N m⁻² vs. 6.9 mg N m⁻²) and C_org (126 mg C m⁻² vs. 94 mg C m⁻²) egested by the worms in one year. Thus, our results demonstrate an important indirect stimulatory effect of elevated atmospheric CO₂ on earthworm activity which may have profound effects on ecosystem function and plant community structure in the long term. Received: 3 November 1996 / Accepted: 11 January 1997     

Differences in carbon fluxes between forested and cultivated micronesian tropical peatlands

Taro is a staple crop that is often grown in wetlands throughout the Indo-Pacific, but the long-term impacts of its cultivation on wetland ecosystem functions are unknown. The objective of this study was to determine how cultivating taro affects carbon cycling by comparing key pathways in a forested peatland and an adjacent cultivated taro patch. Leaves decomposed rapidly at both sites with roughly 73% remaining after 2 weeks, 53% after 8 weeks, 38% after 17 weeks, and 17% after 36 weeks. Root decomposition proceeded much more slowly with roughly 93% remaining after 2 weeks, 80% after 8 weeks, 71% after 17 weeks, and 66% after 36 weeks. Annual litterfall was 1181 g m^–2 year^–1 and 849 g m^–2 year^–1 for the forested and cultivated sites, respectively. For the two sites combined, litterfall consisted of 78% leaves, 10% reproductive material, 3% branches, and 9% miscellaneous material. Fine root biomass was greater in the forested site than the cultivated site, averaging 205 g m^–2 and 34 g m^–2, respectively. Fine root production was much greater in the forested than the cultivated site, averaging 226 g C m^–2 year^–1 and 48 g C m^–2 year^–1, respectively. Soil respiration averaged 99 mg C m^–2 h^–1 and 55 mg C m^–2 h^–1 at the forested and cultivated sites, respectively. We found that the major change to carbon fluxes in the cultivated site was less carbon was entering the peatland, particularly less root production. Alterations to the carbon cycle caused by cultivation would probably not be permanent, because taro patches are periodically abandoned and allowed to regenerate naturally.     

Root biomass,root distribution and the fine-root growth dynamics of Quercus coccifera L. in the garrigue of southern France

Quercus coccifera L., the characteristic scrub oak of the garrigue, covers more than 100,000 ha in southern France alone. Precipitation in this area averages 900 mm/year and summer rains are not rare. A total belowground biomass of 7.2 kg/m², including rhizomes and lignotubers, was harvested. Roots were concentrated in the uppermost 50 cm of the soil. It was hypothesized that low winter temperatures inhibit active fine-root growth. This hypothesis was tested by means of fine-root extractions of soil samples from 0–50 cm depth from November 1987 to June 1988. Although the fine-root analysis could not be extended into late summer and fall, the data supported the hypothesis. Ratios of live/dead fine roots reached their minimum at 0.2–0.3 from December to April. They increased to 1.0–1.2 during late spring and early summer. Initiation of fine-root growth in early April was synchronous with bud break. Starch contents of roots, rhizomes, and lignotubers fluctuated from 4.3% in January to 8.3% in April. The starch stored in belowground organs of Q. coccifera in a closed canopy stand amounted to about 500 g/m² in April. This amount declined to 400 g with bud burst and fine-root growth initiation.     

Production,Respiration, and Overall Carbon Balance in an Old-growth <Emphasis Type="Italic">Pseudotsuga</Emphasis>-<Emphasis Type="Italic">Tsuga</Emphasis> Forest Ecosystem

Ground-based measurements of stores, growth, mortality, litterfall, respiration, and decomposition were conducted in an old-growth forest at Wind River Experimental Forest, Washington, USA. These measurements were used to estimate gross primary production (GPP) and net primary production (NPP); autotrophic respiration (R_a) and heterotrophic (R_h) respiration; and net ecosystem production (NEP). Monte Carlo methods were used to calculate uncertainty (expressed as ± 2 standard deviations of 200–400 calculations). Live carbon (C) stores were 39,800 g C m^–2 (34,800–44,800 g C m^–2). The store of C in detritus and mineral soil was 22,092 g C m^–2 (20,600–23,600 g C m^–2), and the total C stores were 61,899 g C m^–2 (56,600–67,700 g C m^–2). Total NPP was 597 g C m^–2 y^–1 (453 to 741 g C m^–2 y^–1). R_a was 1309 g C m^–2 y^–1 (845–1773 g C m^–2 y^–1), indicating a GPP of 1906 g C m^–2 y^–1 (1444–2368 g C m^–2 y^–1). R_h, including the respiration of heart rots in tree boles, was 577 g C m^–2 y^–1 (479–675 g C m^–2 y^–1). Long-term NEP was estimated to be +20 g C m^–2 y^–1 (–116 to +156 g C m^–2 y^–1), indicating this stand might be a small sink. These estimates contrast with the larger sink estimated at the same site using eddy-flux methods. Several hypotheses to explain this discrepancy were explored, including (a) undetected biomass increases, (b) underestimates of NPP, (c) unmeasured losses, and (d) a temporal mismatch between the two sets of measurements. The last hypothesis appears the most likely.