Translocation,remineralization, and turnover of nitrogen in the roots and rhizomes of Spartina alterniflora (Gramineae)

We used ¹⁵N to quantify rates of N translocation from aerial to belowground tissues, foliar leaching, and turnover and production of root and rhizome biomass in the plant-sediment system of short Spartina alterniflora areas of Great Sippewissett Marsh, Massachusetts. Decay of belowground tissues in litterbag incubations at 1- and 10-cm depths resulted in 80% remineralization of the original plant (¹⁵N-labeled) N and 20% burial after 3 years. Translocation of ¹⁵N from plant shoots in hydrologically controlled laboratory lysimeters maintained under field conditions was 38% of the aboveground pool while leaching of N was 10% from June to October. Most of the translocated N was not retranslocated to new aboveground growth in December but appeared to be either remineralized or buried in the sediment. Injection of ¹⁵N into field stands of grass showed initially high incorporation into plants followed by a continuous decline over the next 7 years yielding a gross tumover time of 1.5–1.6yr. Correcting the gross N turnover for recycling of label via translocation and uptake of remineralized label during this period, a net root and rhizome turnover time of 1.0–1.1 yr was obtained. Combining the turnover time with independent estimates of seasonal belowground biomass yielded an estimate of belowground production of 929–1,022 g C m⁻² yr⁻¹, similar to measurements by traditional biomass harvest, CO² based budgets and models for comparable areas of this marsh. Integration of the production and nitrogen balance estimates for short Spartina marsh yielded translocation, 1.4 g N m⁻² yr⁻¹, leaching, 0.4 g N m⁻² yr⁻¹, remineralization, 14.9–16.3 g N m⁻² yr⁻¹, and burial, 3.7–4.1 g N m⁻² yr⁻¹.     

A comparative study of above-ground productivity of dominant U.S. Gulf Coast marsh species

Above-ground productivity of dominant freshwater, brackish, and salt-marsh species from the U.S. Gulf Coast was evaluated using both gas exchange techniques and harvest methods. Both techniques showed significant differences in productivity among the study species which represent major components of their respective communities. Estimates of net aerial primary productivity using the harvest method yielded 3683 g dw (dry weight) m^?2yr^?1 for Spartina alterniflora (tall), 2008 g dw m^?2yr^?1 for S. alterniflora (short), 3677 g dw m^?yr^?1 for S. patens and 1641 g dwm^?yr^?1 for Panicum hemitomon. Carbon balance estimated from gas exchange calculation yielded values approximately equivalent to a biomass accumulation of 6024 g dw m^?2yr^?1 for S. alterniflora (tall), 3047 g dw m^?yr^?1 for S. alterniflora (short), 5702 g dw m^?yr^?1 for S. patens, and 2912 g dm^?yr^?1 for P. hemitomon. The net aerial primary production was estimated to be approximately 61% of total productivity in S. alterniflora (tall-form) and 66%o of total productivity in short-form, 64% in S. patens and 56%) in P. hemitomon. The assimilation data also indicated that Spartina alterniflora and S. patens continue carbon fixation throughout the year while assimilation in Panicum hemitomon is absent due to lack of live leaves during the winter. Various aspects of harvest and gas exchange techniques are discussed.     

Estimation of the primary productivity of Spartina alterniflora using a canopy model

A comprehensive canopy productivity model was built to study the productivity of a primary salt marsh grass, Spartina alterniflora. in Georgia, USA The canopy model was unique in employing plant demographic data to reconstruct canopy profiles and dynamics, which showed many growth processes that are otherwise difficult to discern in the field By linking canopy dynamics and leaf photosynthesis, the net total primary productivity of S alterniflora m a Georgia salt marsh was estimated to be 1421, 749, and 1441 g C m^-2 yr^-1 for the tall, short, and N-fertilized short populations respectively These estimates are reasonable in terms of the physiological capacity of S alterniflora and well below the range of 3000–4200 g C m^-2 yr^-1 as reported by some recent harvest studies Our detailed analysis suggested the net total productivity of S alterniflora might be greatly overestimated in the past This is mainly because of 1) failure to consider the translocation of photosynthate between aboveground and belowground parts, and 2) possible overestimates of belowground production We estimated the net belowground production to be 872, 397, and 762 g C m^-2 yr^-1 for the tall, short, and N-fertilized populations respectively After receiving nitrogen fertilizer, the net leaf carbon fixation in the short population increased from 1489 to 2487 g C m^-2 yr^-1, and our simulation showed the contribution of elevated leaf N to this increase was small, 21%, compared with that of increased leaf area, 79% Both tall and short populations allocated ca 48-49% of their annual gross leaf carbon fixation to belowground structures Nitrogen enrichment caused more allocation to aboveground parts in the short population, mainly for increasing leaf area The canopy model assumed that there was no leaf photosynthesis under tidal submergence, but if this assumption was relaxed, then leaf carbon fixation might increase 7–13% for different S alterniflora populations Although this research focused only on a salt marsh species, our general approaches, especially the coupling of leaf physiology with the reconstructed canopies, should be applicable to the study of production processes of many other plant populations     

Biomass and aboveground production of four angiosperms in Cantabrian (N. Spain) salt marshes

New data of aboveground biomass and production of four angiosperms over a 12 month period for the Cantabrian Sea salt marshes (Bay of Biscay, N. Spain) are presented. Based on harvest methods, maximum aboveground total biomass values for Spartina maritima (Curtis) Fernald, Spartina alterniflora Loisel, Salicornia ramosissima J. Woods and Halimione portulacoides (L.) Aellen were 628, 1109, 480 and 1267 gm^-2, respectively. We conclude that although a slight latitudinal gradient in biomass is revealed in the data compiled with reference to some of the species studied, more work is neccesary in order to assess the potential productivity of these ecosystems on the coasts of Europe and/or to make comparisons with salt marshes of the American coasts. Annual net aerial primary production estimates using Smalley's method were: 296, 1160, 486 and 952 gm^-2yr^-1, for Spartina maritima, Spartina alterniflora, Salicornia ramosissima and Halimione portulacoides, respectively. These results together with turnover rate estimates point to the lack of vigour of the native S. maritima, while the exotic S. alterniflora, which seems to be spreading along the Cantabrian estuaries, behaves like a veritable pionner throughout the low marshes in this region.     

Primary production and biomass on a Dutch salt marsh: emphasis on the below-ground component

The primary production and below-ground biomass of angiosperms were measured in four almost monospecific vegetation stands situated on a salt marsh along the Oosterschelde estuary, The Netherlands. Maximum below-ground biomass values found for Spartina anglica, Elymus pycnanthus, Halimione portulacoides and Triglochin maritima, were very high relative to values reported from other European salt marshes: 12 586, 9 717, 17 737 and 16 121 g m^-2 respectively. These relatively high values may be due to the fineness of the sieve used, compared to other studies. The actual values are likely to be even higher because the sample treatment has probably caused loss of fine root material. Below-ground production estimates, based on the difference between maximum and minimum biomass, yielded: 6 044 g m^-2 yr^-1 for Spartina, 4 421 g m^-2 yr^-1 for Elymus, 7 799 g m^-2 yr^-1 for Halimione and 3 475 g m^-2 yr^-1 for Triglochin. This high production is mainly concentrated in the deeper layers of the root environment (20–60 cm). Although these production figures are considerably higher than those generally reported for comparable species or vegetation types in Europe, statistical evidence suggests that, for the first three species, they are real values rather than figures caused by random fluctuations.     

Root turnover as determinant of the cycling of C,N, and P in a dry heathland ecosystem

Root production and turnover were studied using sequential core sampling and observations in permanent minirhizotrons in the field in three dry heathland stands dominated by the evergreen dwarfshrub Calluna vulgaris and the grasses Deschampsia flexuosa and Molinia caerulea, respectively. Root biomass production, estimated by core sampling, amounted to 160 (Calluna), 180 (Deschampsia) and 1380 (Molinia) g m^-2 yr^-1, respectively. Root biomass turnover rate in Calluna (0.64 yr^-1) was lower compared with the grasses (Deschampsia: 0.96 yr^-1; Molinia 1.68yr^-1)). Root length turnover rate was 0.75–0.77 yr^-1 (Deschampsia) and 1.17–1.49 yr^-1 (Molinia), respectively. No resorption of N and P from senescing roots was observed in either species. Input of organic N into the soil due to root turnover, estimated using the core sampling data, amounted to 1.8 g N m^-2 yr^-1(^Calluna), 1.7 g N m^-2 yr^-1 (Deschampsia) and 19.7 g N m^-2 yr^-1 (Molinia), respectively. The organic P input was 0.05, 0.07 and 0.55 g P M^-2 yr^-1, respectively. Using the minirhizotron turnover estimates these values were20–22% (Deschampsia) and 11–30% (Molinia) lower.When the biomass turnover data were used, it appeared that in the Molinia stand root turnover contributed 67% to total litter production, 87% to total litter nitrogen loss and 84% to total litter phosphorus loss. For Calluna and Deschampsia these percentages were about three and two times lower, respectively.This study shows that (1) Root turnover is a key factor in ecosystem C, N, and P cycling; and that (2) The relative importance of root turnover differs between species.     

Response of understory vegetation to variable tree mortality following a mountain pine beetle epidemic in lodgepole pine stands in northern Utah

We analyzed the long-term dynamics of aboveground biomass ofLeymus chinense steppe in relation to interannual variation of precipitation and temperature during 1980–1989 at levels of community, growth form and species in the Xilin river basin, Inner Mongolia Autonomous Region, China. Annual aboveground net primary production (ANPP) varied from 154.00 g m^-2 yr^-1 in 1980 to 318.59 g m^-2 yr^-1 in 1988, with a mean of 248.63 g m^-2 yr^-1 and the coefficient of variation of 25%. ANPP was not significantly correlated to annual precipitation and total precipitation during April–September atp0.05 level, but precipitation in May and August accounted for 69% of interannual variation of ANPP. The means of rain use efficiency and water use efficiency ofL. chinense steppe were 8.1 kg DM ha^-1 mm^-1 yr^-1 and 0.89 mg DM g^-1 H₂O respectively. Aboveground biomass of various growth forms and species had different response patterns to interannual variation of precipitation and temperature. Monthly and seasonal distribution of precipitation and temperature were the key controls of aboveground biomass of species.     

Biomass,production and woody detritus in an old coast redwood (<Emphasis Type="Italic">Sequoia sempervirens</Emphasis>) forest

We examined aboveground biomass dynamics, aboveground net primary production (ANPP), and woody detritus input in an old Sequoia sempervirens stand over a three-decade period. Our estimates of aboveground biomass ranged from 3300 to 5800 Mg ha⁻¹. Stem biomass estimates ranged from 3000 to 5200 Mg ha⁻¹. Stem biomass declined 7% over the study interval. Biomass dynamics were patchy, with marked declines in recent tree-fall patches <0.05 ha in size. Larger tree-fall patches approaching 0.2 ha in size were observed outside the study plot. Our estimates of ANPP ranged from 6 to 14 Mg ha⁻¹yr⁻¹. Estimates of 7 to 10 Mg ha⁻¹yr⁻¹ were considered to be relatively accurate. Thus, our estimates based on long-term data corroborated the findings of earlier short-term studies. ANPP of old, pure stands of Sequoia was not above average for temperate forests. Even though production was potentially high on a per stem basis, it was moderate at the stand level. We obtained values of 797 m³ ha⁻¹ and 262 Mg ha⁻¹ for coarse woody detritus volume and mass, respectively. Fine woody detritus volume and mass were estimated at 16 m³ ha⁻¹ and 5 Mg ha⁻¹, respectively. Standing dead trees (or snags) comprised 7% of the total coarse detritus volume and 8% of the total mass. Coarse detritus input averaged 5.7 to 6.9 Mg ha⁻¹yr⁻¹. Assuming steady-state input and pool of coarse detritus, we obtained a decay rate constant of 0.022 to 0.026. The old-growth stand of Sequoia studied had extremely high biomass, but ANPP was moderate and the amount of woody detritus was not exceptionally large. Biomass accretion and loss were not rapid in this stand partly because of the slow population dynamics and low canopy turnover rate of Sequoia at the old-growth stage. Nomenclature: Hickman (1993).     

Spatial heterogeneity of ecosystem carbon fluxes in a broadleaved forest in Northern Germany

Carbon fluxes were investigated in a mature deciduous forest, located in Northern Germany (53°47′N–10°36′E), by means of eddy‐covariance technique, stand survey and models. This forest has been managed following a concept of nature‐oriented forestry since the 1980s. One of the goals of the study was to test whether changed management led to increased carbon sequestration. The forest contains several broadleaved tree species. Depending on wind direction, the fetch‐area of the eddy‐covariance data was dominated by different tree species. Three subplots dominated by Oak, Beech or Alder/Ash could be distinguished from the tower data. In each of these subplots, 30 × 30 m² areas were defined to analyse leaf area index, litterfall and the increase of the wood biomass. Eddy‐covariance analysis showed that the gross primary productivity (GPP′) was higher in the Oak subplot (?1794 g C m^?2 yr^?1) in comparison with the Beech plot and the Alder/Ash plot (?1470 and ?1595 g C m^?2 yr^?1, respectively). The total ecosystem respiration (TER) was the highest in the Alder/Ash‐dominated subplot (1401 g C m^?2 yr^?1) followed by the Oak plot and the Beech plot (1235 and 1174 g C m^?2 yr^?1, respectively). The resulting net ecosystem productivity (NEP) was ?559 g C m^?2 yr^?1 for the Oak‐dominated subplot, ?295 g C m^?2 yr^?1 for the Beech plot and ?193 g C m^?2 yr^?1 for the Alder/Ash plot. From Stand survey and modelling, the net primary productivity was estimated as 1103, 702 and 671 g C m^?2 yr^?1 in the Oak, Beech and Alder/Ash plot, respectively. Also carbon flux with litterfall was the highest in the Oak plot 343 g C m^?2 yr^?1 and lowest in Alder/Ash plot (197 g m^?2 yr^?1) with the Beech plot in between (228 g m^?2 yr^?1). The observations indicate an increase of the proportion of litterfall with increasing GPP′ and a different ability of carbon sequestration of the three stands in medium temporary scale. Only in the Oak stand that comprised the oldest trees and the most structured canopy the carbon sequestration was increased compared with conventionally managed forests.     

MODELING THE ANNUAL PRODUCTION OF INTERTIDAL BENTHIC MICROALGAE IN ESTUARINE ECOSYSTEMS1

The purpose of this study was to develop and validate a habitat-specific production simulation model to quantify annual benthic microalgal production in North Inlet estuary, South Carolina. Using hourly measurements of incident irradiance during 1990–1991 as the forcing function, the simulation model was used to obtain hourly estimates of areal benthic microalgal gross primary production in five habitat types. The model, which was validated using actual measurements of production, showed good (r²= 0.63, P < 0.001) agreement between observed and predicted production in the short Spartina alterniflora Loisel zone habitats showed the highest mean hourly production (61.1 mg C m^?2 h^?1) while intertidal mudflats had the maximum hourly rate (166.9 mg C m^?2 h^?1). Daily production was highly variable, primarily due to daily fluctuations in irradiance. Annual estimates of habitat-specific production were multiplied by the mates of habitat-specific production were multiplied by the known area of each habitat type to determine total microalgal production for the estuary (3.423 × 10⁹ g C yr^?1). Short Spartina zone habitats provided 45% of total microalgal annual production, followed by intertidal mudflats (22%), tall Spartina zones (18%), shallow subtidal (13%) and microalgal production exceeds phytoplankton and microalgal production but is less than Spartina production.     

Estimates of soil respiration and net primary production of three forests at different succession stages in South China

Soil respiration (heterotropic and autotropic respiration, R_g) and aboveground litter fall carbon were measured at three forests at different succession (early, middle and advanced) stages in Dinghushan Biosphere Reserve, Southern China. It was found that the soil respiration increases exponentially with soil temperature at 5 cm depth (T_s) according to the relation R_g=a exp(bT_s), and the more advanced forest community during succession has a higher value of a because of higher litter carbon input than the forests at early or middle succession stages. It was also found that the monthly soil respiration is linearly correlated with the aboveground litter carbon input of the previous month. Using measurements of aboveground litter and soil respiration, the net primary productions (NPPs) of three forests were estimated using nonlinear inversion. They are 475, 678 and 1148 g C m^?2 yr^?1 for the Masson pine forest (MPF), coniferous and broad‐leaf mixed forest (MF) and subtropical monsoon evergreen broad‐leaf forest (MEBF), respectively, in year 2003/2004, of which 54%, 37% and 62% are belowground NPP for those three respective forests if no change in live plant biomass is assumed. After taking account of the decrease in live plant biomass, we estimated the NPP of the subtropical MEBF is 970 g C m^?2 yr^?1 in year 2003/2004. Total amount of carbon allocated below ground for plant roots is 388 g C m^?2 yr^?1 for the MPF, 504 g C m^?2 yr^?1 for the coniferous and broad‐leaf MF and 1254 g C m^?2 yr^?1 for the subtropical MEBF in 2003/2004. Our results support the hypothesis that the amount of carbon allocation belowground increases during forest succession.     

ORGANIC MATTER DYNAMICS IN FOUR SEASONALLY FLOODED FOREST COMMUNITIES OF THE DISMAL SWAMP

Budgets of organic matter dynamics for plant communities of the Great Dismal Swamp were developed to summarize an extensive data base, determine patterns of biomass allocation, transfer and accumulation, and make comparisons with other forested wetlands. Aboveground net primary production on the flooded sites (1,050–1,176 g m^-2 yr^-1) was significantly greater than on a rarely flooded site (831 g m^-2 yr^-1). Estimates of belowground net primary production were comparable to aboveground production on flooded sites (824–1,221 gm^-2 yr^-1). However, productivity was nearly three times greater belowground than aboveground on the rarely flooded site (2,256 g m^-2 yr^-1). Aboveground productivity in Dismal Swamp forests is relatively high compared to other forested wetlands. This is attributed to the timing and periodic nature of flood events. Fine root turnover is shown to be an important source of soil organic matter. Estimates indicate that roots contribute about 60% of the annual increment to soil organic matter. Leaflitter contributes 6–28% and wood debris contributes 5–15%. Comparisons with other forested wetlands suggest that detritus accounts for greater than half of the total organic matter (living + dead) in many wetland systems.     

Net primary production and net ecosystem production of a boreal black spruce wildfire chronosequence

Net primary production (NPP) was measured in seven black spruce (Picea mariana (Mill.) BSP)‐dominated sites comprising a boreal forest chronosequence near Thompson, Man., Canada. The sites burned between 1998 and 1850, and each contained separate well‐ and poorly drained stands. All components of NPP were measured, most for 3 consecutive years. Total NPP was low (50–100 g C m^?2 yr^?1) immediately after fire, highest 12–20 years after fire (332 and 521 g C m^?2 yr^?1 in the dry and wet stands, respectively) but 50% lower than this in the oldest stands. Tree NPP was highest 37 years after fire but 16–39% lower in older stands, and was dominated by deciduous seedlings in the young stands and by black spruce trees (>85%) in the older stands. The chronosequence was unreplicated but these results were consistent with 14 secondary sites sampled across the landscape. Bryophytes comprised a large percentage of aboveground NPP in the poorly drained stands, while belowground NPP was 0–40% of total NPP. Interannual NPP variability was greater in the youngest stands, the poorly drained stands, and for understory and detritus production. Net ecosystem production (NEP), calculated using heterotrophic soil and woody debris respiration data from previous studies in this chronosequence, implied that the youngest stands were moderate C sources (roughly, 100 g C m^?2 yr^?1), the middle‐aged stands relatively strong sinks (100–300 g C m^?2 yr^?1), and the oldest stands about neutral with respect to the atmosphere. The ecosystem approach employed in this study provided realistic estimates of chronosequence NPP and NEP, demonstrated the profound impact of wildfire on forest–atmosphere C exchange, and emphasized the need to account for soil drainage, bryophyte production, and species succession when modeling boreal forest C fluxes.     

A Quantitative Food Web Model for the Macroinvertebrate Community of a Northern German Lowland Stream

Trophic interactions and cycling of organic carbon within the macroinvertebrate community of a Northern German lowland stream were analyzed based on a compartment model. The network model describes the structure of the food web quantifying biomass, production, and consumption of their elements, of the entire system and between trophic levels. System primary production is 153.7 g C m⁻² yr⁻¹ and invertebrate production 53.3 g C m⁻² yr⁻¹. Invertebrate consumption amounts to 702.6 g C m⁻² yr⁻¹. Main flows are identified between trophic level 1 and 2 and are connected with highly productive compartments. ‘Anodonta and Pseudanodonta’ and Dreissena polymorpha show the highest consumption of all groups with 269.9 g C m⁻² yr⁻¹ and 114.1 g C m⁻² yr⁻¹, respectively. System consumption is highest on the import from the upstream lake with 532.5 g C m⁻² yr⁻¹, sediment detritus with 135.5 g C m⁻² yr⁻¹, and primary producers with 25.7 g C m⁻² yr⁻¹. The lowest predation pressure is observed for Bivalvia with an ecotrophic efficiency of <10% and highest for Chironomidae with 91%. Approximately 20% of organic matter entering the detritus pool are recycled to the living groups of the system. Transfer efficiencies between discrete trophic levels are generally low except for transfer of detrital material between level I and II.     

Nitrogen flows in Louisiana Gulf Coast salt marsh: Spatial considerations

Nitrogen flux data was synthesized in developing a nitrogen flow budget for a Louisiana Barataria BasinSpartina alterniflora salt marsh. Results demonstrate the importance of spatial consideration in developing a nitrogen budget for coastal marshes. Using a mass balance approach nitrogen inputs balanced nitrogen sinks or losses from a marsh soil-plant system with a specific rooting depth. However, per unit areas on a local scale, marshes serve as a large sink for nitrogen due to rapid accretion which removes 17.O g N m^–2yr^–1 through subsidence below the root zone. On a larger spatial scale (regional) it is shown that the marshes do not serve as a large nitrogen sink. The rapid marsh deterioration currently occurring in the rapidly subsiding marshes of the Mississippi River deltaic plain account for a net regional loss of 12.5 g N m^–2yr^–1. Thus, regionally the net sink is equivalent to only 5 g N m^–2yr^–1 as compared to 17.0 g N m^–2yr^–1 on a local scale.     

The soil fauna of a beech forest on limestone: trophic structure and energy budget

Summary The soil fauna of a mull beech forest on lime-stone in southern Lower Saxony (West Germany) was sampled quantitatively. Biomass estimates, trophic characteristics, and measurement and calculation of the energetic parameters of the constituent animal populations were used to construct an energy budget of the total heterotrophic subsystem of the forest. Mean annual zoomass amounted to about 15 g d wt m^–2; earthworms (about 10 g d wt m^–2) and other groups of the macrofauna were dominant. Protozoa constituted about 1.5 g d wt m^–2. Relative distribution of zoomass among the trophic categories was 50% macrosaprophages, 30% microsaprophages, 12% microphytophages, and 4% zoophages. Total annual consumption rate of the saprophagous and microphytophagous soil fauna (6328 and 4096 kJ m^–2 yr^–1, respectively) was of the same order of magnitude as annual litter fall (canopy leaves 6124 kJ m^–2 yr^–1, flowers and fruits 944 kJ m^–2 yr^–1, herbs 1839 kJ m^–2 yr^–1, fine woody material 870 kJ m^–2 yr^–1, tree roots 3404 kJ m^–2 yr^–1, without coarse woody litter). Primary decomposers (macrosaprophages) were the key group for litter comminution and translocation onto and into the soil, thus contributing to the high decomposition rate (k=0.8) for leaf litter. Consumption rates of the other trophic groups were (values as kJ m^–2 yr^–1): bacteriophages 2954, micromycophages 416, zoophages 153. Grazing pressure of macrophytophages (including rhizophages) was low. Faeces input from the canopy layer was not significant. Grazing pressure on soil microflora almost equalled microbial biomass; hence, a large fraction of microbial production is channelled into the animal component. Predator pressure on soil animals is high, as a comparison between consumption rates by zoophages and production by potential prey — mainly microsaprophages, microphytophages and zoophages — demonstrated. Soil animals contributed only about 11% to heterotrophic respiration. However, there is evidence that animals are important driving variables for matter and energy transfer: key processes are the transformation of dead organic material and grazing on the microflora. It is hypothesized that the soil macrosaprophages are donor-limited.     

Postfire carbon pools and fluxes in semiarid ponderosa pine in Central Oregon

Forest fire dramatically affects the carbon storage and underlying mechanisms that control the carbon balance of recovering ecosystems. In western North America where fire extent has increased in recent years, we measured carbon pools and fluxes in moderately and severely burned forest stands 2 years after a fire to determine the controls on net ecosystem productivity (NEP) and make comparisons with unburned stands in the same region. Total ecosystem carbon in soil and live and dead pools in the burned stands was on average 66% that of unburned stands (11.0 and 16.5 kg C m⁻², respectively, P<0.01). Soil carbon accounted for 56% and 43% of the carbon pools in burned and unburned stands. NEP was significantly lower in severely burned compared with unburned stands (P<0.01) with an increasing trend from −125±44 g C m⁻² yr⁻¹ (±1 SD) in severely burned stands (stand replacing fire), to −38±96 and +50±47 g C m⁻² yr⁻¹ in moderately burned and unburned stands, respectively. Fire of moderate severity killed 82% of trees <20 cm in diameter (diameter at 1.3 m height, DBH); however, this size class only contributed 22% of prefire estimates of bole wood production. Larger trees (> 20 cm DBH) suffered only 34% mortality under moderate severity fire and contributed to 91% of postfire bole wood production. Growth rates of trees that survived the fire were comparable with their prefire rates. Net primary production NPP (g C m⁻² yr⁻¹, ±1 SD) of severely burned stands was 47% of unburned stands (167±76, 346±148, respectively, P<0.05), with forb and grass aboveground NPP accounting for 74% and 4% of total aboveground NPP, respectively. Based on continuous seasonal measurements of soil respiration in a severely burned stand, in areas kept free of ground vegetation, soil heterotrophic respiration accounted for 56% of total soil CO₂ efflux, comparable with the values of 54% and 49% previously reported for two of the unburned forest stands. Estimates of total ecosystem heterotrophic respiration (R_h) were not significantly different between stand types 2 years after fire. The ratio NPP/R_h averaged 0.55, 0.85 and 1.21 in the severely burned, moderately burned and unburned stands, respectively. Annual soil CO₂ efflux was linearly related to aboveground net primary productivity (ANPP) with an increase in soil CO₂ efflux of 1.48 g C yr⁻¹ for every 1 g increase in ANPP (P<0.01, r²= 0.76). There was no significant difference in this relationship between the recently burned and unburned stands. Contrary to expectations that the magnitude of NEP 2 years postfire would be principally driven by the sudden increase in detrital pools and increased rates of R_h, the data suggest NPP was more important in determining postfire NEP.     

Macro-invertebrate populations and production on a salt-marsh in east England dominated by Spartina anglica

Summary The populations and production of the macroinvertebrates of a Spartina anglica salt-marsh in eastern England were studied over two years. A total of fifteen species were recorded in the sediments, of which twelve species were of regular occurrence, and the total population density recorded ranged from 3,481 m^-2 to 11,444m^-2 over the twenty-four sampling occasions.The four most abundant species were Nereis diversicolor, Tubifex costatus, Corophium volutator and Hydrobia ulvae. Thirteen further taxa were associated with the canopy of Spartina, with a total population density ranging from 0 to 1,149 m^-2. Total monthly standing crop ranged from 1.8 to 8.5 g C m^-2 with peaks in July/August in both years. Nereis diversicolor contributed 55% to 86% of total biomass in each month.Production and respiration for each species was determined and annual assimilation calculated. The total annual production was 16 g C m^-2 a^-1 in both 1979 and 1980, with a corresponding assimilation of 60 g C m^-2 a^-1 Nereis diversicolar accounted for >80% of production and assimilation in both years, and the species is clearly of considerable potential importance in the dissipation of Spartina material. The canopy dwelling species accounted for about 1% of the total annual production and assimilation.     

Secondary Production of Macrobenthic Invertebrates from Delaware Bay and Coastal Waters

Secondary production of benthic invertebrates was estimated for Delaware Bay and coastal Delaware. Production and turnover ratios were highest in Delaware Bay (P = 46,572 mg AFDW m⁻² yr⁻¹, P:B = 6,O) and progressively lower at two coastal stations (P = 7,501 to 30,124 mg AFDW m⁻² yr⁻¹, P:B = 2.3 to 5.3, and P = 4,485 to 4,492mg AFDW m⁻² yr⁻¹, P:B =2.3 to 4.8). Production was inversely related to sediment particle size. Production in Delaware Bay was relatively evenly distributed between deposit feeding polychaetes and suspension feeding molluscs with a definite shift in production dominance to suspension feeding molluscs at the coastal stations. Moreover, crustaceans and echinoderms played a larger role in production at the coastal stations than in Delaware Bay. Concerns about the health of soft-bottom communities in Delaware Bay expressed earlier were not supported here. Finally, it was concluded that P and P: B from the Delaware Bay area were very similar to those obtained from other areas in the North Atlantic which agrees with estimates for other estuaries in the northern hemisphere.     

Distribution and production ofChironomus salinarius (Diptera: Chironomidae) in a shallow coastal lagoon in the Bay of Cádiz

The abundance, generation time and production ofChironomus salinarius larvae in a lagoon fish-pond system in the Bay of Cádiz were studied by taking monthly samples at 3 sites during 1991 and 1992. Numerical abundance and biomass of larvae showed considerable spatial, seasonal and interannual variation (ANCOVAs,P<0.001). The maximum mean annual density was 7048 larvae m^–2, and corresponded to a biomass of 3.08 g dry weight (DW) m^–2. It was recorded at the site with the lowest rate of water renewal. Seasonal patterns were similar at all sites, with main annual peaks of abundance and biomass in autumn-early winter. Chironomid density was positively related to the biomass of benthic macroalgae (P<0.001). The population studied was multivoltine with a probable average of five generations per year, with overlapping cohorts and a predominance of third- and fourth-instar larvae. Estimates of annual production ranged between 72.2 g DW m^–2 yr^–1 at the site with the lowest rate of water renewal in 1991 and 0.1 g DW m^–2 yr^–1 at the site with the highest rate of water renewal in 1992. Mean annual production and the production/biomass ratio for the system was estimated to be 16.8 g DW m^–2 yr^–1 and 12.7, respectively. Possible factors leading to the observed density fluctuations are discussed, as well as possible sources of error in production estimates.