Comparison of giant cutgrass productivity in tidal and impounded marshes with special reference to tidal subsidy and waste assimilation

Aboveground live standing crop of giant cutgrass (Zizaniopsis miliacea) populations in similar freshwater tidal and impounded nontidal marshes were almost identical (peaking at 1,039 g per m² in each). The mortality, however, was greater in the tidal marsh resulting in significantly (95% level) greater annual production of aboveground cutgrass in the tidal (1,530±103 g per m² per yr) than the impounded (1,172±88 g per m² per yr) marsh, a 31% difference which we consider to be a measure of tidal subsidy. Belowground production also was found to average higher in the tidal marsh, but estimates were not as satisfactory as the aboveground results due to sampling difficulties. Combined annual above and belowground net production comes to an estimated 2,048 ±101 g per m² per yr for the tidal and 1,481±219 for the impounded cutgrass marsh. The potential of freshwater tidal marshes for tertiary treatment of wastes is briefly discussed.     

Productivity,detritus flux,and nutrient cycling in a Carex lyngbyei tidal marsh

Net annual primary production of a sedge Carex lyngbyei dominated tidal marsh in the Fraser River estuary, British Columbia, Canada was 634 g ash-free dry weight (AFDW) per m² per yr (687 g dry weight per m² per yr). Mean maximum shoot elongation during the short (May to August) growing season was 1.88 cm per day from overwintering shoots. The maximum aboveground standing crop of 690 g AFDW per m² represented only 25% of the total below-ground biomass, which appears to be controlling most of the critical life history processes of the sedge marsh. An estimate of 14 percent of the aboveground standing crop was lost through leaching of dissolved organic carbon from the growting plant. Aboveground tissue losses, which were negligible during the growing season, occurred primarily via translocation in autumn and tidal export during the winter. In situ measurements showed that of the original maximum standing crop, approximately 38%, 37%, and 25% were lost by downward translocation, tidal export, and sediment burial, respectively. Based on changes in above and belowground nutrient pools, rapid spring (May to late June) uptake rates of 109 mg N per m² per day and 23.0 mg P per m² per day by shoots were followed by downward translocation rates of 44.8 mg N per m² per day and 12.2 mg P per m² per day during late June to the end of August. Aboveground leaching rates were estimated as 23.9 mg N per m² per day and 7.8 mg P m² per day and belowground uptake rates as 100 mg N per m² per day and 26 mg P per m² per day; root uptake occurred primarily after late June. Nutrient levels in decomposing litter more than doubled over the winter period showing a pattern of nutrient enrichment characteristic of marsh ecosystems. *** DIRECT SUPPORT *** A01BY023 00004     

The role of tides in redistributing macrodetrital aggregates within the Swartvlei estuary

The Swartvlei estuary possesses a prolific growth of both intertidal and subtidal eelgrass,Zostera capensis. During 1984 less than 12% of the eelgrass beds were located in the upper half of the estuary, yet deposition ofZostera/macroalgal wrack in this region, when the estuary was linked to the sea (open phase), was similar to that in the lower half. Over a period of 20 semidiurnal tidal cycles there was a net gain of 2.5 tonnes dry mass of plant material into the upper reaches. Export of aquatic macrophytes and filamentous algae from the lower reaches toward the sea over 20 tidal cycles amounted to 1.6 tonnes dry mass. The amount of plant material transported during spring tides was 2 to 3 times greater than that carried during neap tides. Shallowing of the estuary mouth due to sand deposition resulted in a decline in the tidal prism and a decrease in macrodetrital flux. Total export ofZostera and associated algae amounted to 0.87 g ash-free dry mass m^?2d^?1 and represented a monthly export of 18% ofZostera bed biomass. Deposition of plant wrack during the 1984–1985 closed phase amounted to 63 g dry mass per meter of shore per day at the lower reaches site but only 10 g m^?1d^?1 was recorded at the upper reaches site. The relatively low latter value was attributed to the absence of tidal action which transports macrodetritus from the lower and middle reaches into the upper part of the system. During the 1984 open phase 70 g m^?1d^?1 was deposited at the lower reaches site and 68 g m^?1d^?1 at the upper reaches site. The role of tides in the redistribution of aquatic macrophyte primary production in the Swartvlei estuary was therefore clearly underscored.     

Productivity of cordgrass,Spartina alterniflora,estimated from live standing crops,mortality, and leaf shedding in a Virginia salt marsh

Net annual productivity of tall and medium form cordgrass,Spartina alterniflora, was estimated by a new clip sampling method in a sloping foreshore salt marsh at Wallops Island, Virginia. This method measured live standing crops only, to avoid problems of measuring dead biomass inherent in other methods. Losses from live standing crops by shoot mortality and by leaf shedding were estimated from these measurements and added separately to production of live tillers and of live culms. This allowed quantification of various components of production.Spartina tillering in different zones of the marsh produced 62 to 211 g dry weight per m² per yr. Tiller mortality removed 37 to 106 g per m² per yr from live standing crops. Culms produced 348 to 1,132 g per m² before flowering and die-back. Culm mortality removed 28 to 246 g per m² before flowering. Leaf shedding removed an additional 83 g per m² in tall formSpartina. Altogether, net annual productivity These estimates are much higher than previous estimates of productivity and standing crops inSpartina marshes nearby.     

Growth,survival and production in young-of-year populations ofLeiostomus xanthurus Lacépéde residing in tidal creeks

Growth, mortality, and production rates for spot,Leiostomus xanthurus Lacépede, were studied from January 1977 through August 1978 in tidal creeks of the Cape Fear estuary, North Carolina. Winter growth rates were low, averaging 0.14 and 0.16 mm per day in 1977 and 1978, respectively. Mortality rates were significantly higher in polyhaline marshes during 1977, however, similar rates were observed in oligohaline marshes in 1978. The difference is believed due to higher freshwater flows experienced in 1978, which apparently increased mortality rates upriver. Pooled monthly production estimates for all sampling sites combined yielded a value of 0.17 g per m² (257 cal per m²). This figure is similar to those reported for spot collected in seagrass meadows in the vicinity of the Cape Fear estuary (Adams 1976). Because spot migrate offshore in the fall, this production also represents potential export of energy to the marine environment in the form of living biomass.     

Natural Cycles and Transfer of Mercury Through Pacific Coastal Marsh Vegetation Dominated by Spartina foliosa and Salicornia virginica

The potential for marsh plants to be vectors in the transport of mercury species was studied in the natural, mature, tidal China Camp salt marsh on San Pablo Bay. The fluxes of organic matter, mercury (THg), and monomethylmercury (MeHg) were studied in natural stands of Spartina foliosa and Salicornia virginica. Seasonal fluxes from the sediment into aboveground biomass of live plants and subsequent transfer into the dead plant community by mortality were measured. Loss of THg and MeHg from the dead plant community through fragmentation, leaching, and excretion were calculated and were similar to net uptake. Seasonal data were added up to calculate annual mass balances. In S. foliosa, annual net production was 1,757 g DW m^?2, and the annual net uptakes in the aboveground biomass were 305 μg THg m^?2 and 5.720 μg MeHg m^?2. In S. virginica, annual net production was 2,117 g DW m^?2, and the annual net uptakes in aboveground biomass were 99.120 μg THg m^?2 and 1.990 μg MeHg m^?2. Of both plant species studied, S. foliosa had a slightly lower production rate but greater mercury species uptake and loss rates than S. virginica, and, consequently, it is to be expected that S. foliosa matter may affect the local and possibly the regional food web relatively more than S. virginica. However, the actual effects of the input of mercury-species-containing plant-derived particulate matter into the food webs would depend on trophic level, food preference, seasonal cycle of the consumer, total sediment surface area vegetated, location of the vegetation in the marsh landscape, and estuary bay landscape. Since the levels of mercury species in dead plant material greatly exceed those in live plant material (on a dry weight basis), detritivores would ingest greater mercury species concentrations than herbivores, and consumers of S. foliosa would ingest more than consumers of S. virginica. The greatest THg and MeHg losses of both plant species due to mortality and to fragmentation–leaching–excretion occurred in late spring and early autumn, which corresponds to peak MeHg levels observed in sediments of coastal systems of previous studies, suggesting enhanced THg–MeHg export from the marsh to the nearshore sediment.     

Standing crops of marsh vegetation of two tributaries of Chesapeake Bay

A comparative study of the standing crop of marsh vegetation was made of the Patuxent River and Parker Creek, two tributaries of Chesapeake Bay. The biomass of marsh vegetation in the tidal freshwater and brackish regions of the Patuxent was relatively uniform with regard to salinity, seasonally high concentrations of dissolved nitrogen, and phosphorus and nutrient gradient. Maximum values of biomass occurred in the tidal freshwater and slightly brackish water region of Parker Creek, a system whose nutrient concentrations approximated 20% of those of Patuxent River. Biomass values for the Patuxent River and Parker Creek averaged about 1417 and 895 g m^?2 dry weight, respectively. Estimates of total annual marsh production based on the maximum standing crop was 27×10³ and 519 metric tons, respectively, for the Patuxent River and Parker Creek.     

The autecology and production dynamics of eelgrass (Zostera marina L.) in Netarts Bay,Oregon

Changes in biomass, growth form and shoot net primary production in an eelgrass, Zostera marina L., bed were monitored along transects at three tidal heights in Netarts Bay, Oregon, from May 1979 through June 1981. During the growing season, April through October, the mean plastochrone interval was 16.5 d along the low intertidal transect and 11.6 d along the high intertidal transect. The mean export interval was 13.3 d along the low intertidal transect and 11.6 d along the high intertidal transect. The life span of a leaf averaged 48 d along the low intertidal transect and 36 d along the high intertidal transect. Shoot density was positively correlated with mean leaf area index (LAI) until the LAI reached 3.8 to 5.5, above which LAI was negatively correlated with density. The maximum Zostera biomass ranged from 143 (high intertidal transect) to 463 (low intertidal transect) g dry wt m^?2. Maximum values of shoot net production ranged from 4.7 (high intertidal transect) to 13.6 (low intertidal transect) g dry wt m^?2d^?1. Zostera shoot net production was related to light and to the physical damage to the shoots associated with a rapid accumulation of Enteromorpha biomass in the bay. In addition, patterns of biomass accumulation were related to the duration of water coverage, as determined by both tidal height and local impoundments of water. At all transects, biomass sloughed was equal to at least 50% of the shoot net primary production in that area during that time period; sloughed leaves accounted for 25 to 97% of these losses. An estimate of the total annual net primary production of aboveground Zostera in the bed was 17,500 kg, dry wt (SE=3,080 kg dry wt), which was equivalent to a mean annual rate of 383 g C m^?2 (SE=67 g C m^?2)     

Interactive effects of animal disturbance and elevation on vegetation of a tidal freshwater marsh

We studied interactions between animal disturbance (geese, carp, and muskrat) and elevation in a field experiment in tidal freshwater marshes of the Patuxent River, Maryland, United States. Vegetation changes were recorded in fenced and unfenced plots in high and low marsh community types for 2 yr using measurements of areal cover and within-plot frequency (which were averaged to create a dominance index), Leaf Area Index (LAI), and aboveground biomass. We related light environment to differences in vegetation using below-canopy measurements of Photosynthetically Active Radiation (PAR). In the low marsh, total cover of all species, cover of annual species, biomass, and LAI were significantly higher in plots fenced to exclude animals (exclosures) than in unfenced plots (fenced/unfenced total cover=76/40%, annual cover=45/10%, biomass=936/352 g m^?2, LAI=3.3/1.4). PAR was significantly lower in fenced than unfenced plots (fenced/unfenced=115/442 μmol s-1 m^?2). Despite the strong effect of fencing on biomass, species richness per plot (i.e., the number of species per plot, or species density) was not affected significantly by fencing in the low marsh. Most of the observed differences in cover, biomass, LAI, and PAR were due to variation in the abundance of the herbaceous annual speciesBidens laevis (dominance index fenced/unfenced=45/10%) andZizania aquatica (30/12%). In the high marsh community, fencing had only minor effects on plant community composition and did not significantly affect species richness, cover, biomass, PAR, or LAI. Our results show that animals can dramatically affect low marsh vegetation, primarily via physical disturbance or herbivory of shallowly rooted seedlings of annual species.     

Nitrogen cycling and ecosystem exchanges in a Virginia tidal freshwater marsh

Tidal freshwater marshes are diverse habitats that differ both within and between marshes in terms of plant community composition, sediment type, marsh elevation, and nutrient status. Because our knowledge of the nitrogen (N) biogeochemistry of tidal freshwater systems is limited, it is difficult to assess how these marshes will respond to long-term progressive nutrient loading due to watershed development and urbanization. We present a process-based mass balance model of N cycling in Sweet Hall marsh, a pristine (i.e., low nutrient)Peltandra virginica-Pontederia cordata dominated tidal freshwater marsh in the York River estuary, Virginia. The model, which was based on a combination of field and literature data, revealed that N cycling in the system was largely conservative. The mineralization of organic N to NH₄ ⁺ provided almost twice as much inorganic N as was needed to support marsh macrophyte and benthic microalgal primary production. Efficient utilization of porewater NH₄ ⁺ by nitrifiers and other microbes resulted in low rates of tidal NH₄ ⁺ export from the marsh and little accumulation of NH₄ ⁺ in marsh porewaters. Inputs of N from the estuary and atmosphere were not critical in supporting marsh primary production, and served to balance N losses due to denitrification and burial. A comparison of these results with the literature suggests that the relative importance of tidal freshwater marsh N cycling processes, including plant productivity, organic matter mineralization, microbial immobilization, and coupled nitrification-denitrification, are largely independent of small changes in water column N loading. Although very high (millimolar) concentrations of dissolved inorganic N can affect processes including denitrification and plant productivity, the factors that cause the switch from efficient N recycling to a more open N cycle have not yet been identified.     

Comparison of production computation methods in a Southeastern North CarolinaSpartina alterniflora salt marsh

Tagging studies ofSpartina alterniflora Loisel showed no significant differences in stem longevity of short, medium, and tall height forms. Mean stem longevity was 7.9 months, and the experimental turnover rate was 1.5 crops per yr. Five methods to measure productivity (peak standing crop, Milner and Hughes, Smalley, Wiegert and Evans, and Lomnicki, et al.) yielded annual net aerial primary production (NAPP) estimates ranging from 214 to 1,038 g dry wt per m² per yr in a stand of shortSpartina. Turnover rates were computed for each of the methods by dividing the respective production value by the peak standing crop (242 g dry wt per m² per yr). Each computed turnover rate was compared with the experimental value of 1.5 crops per yr to ultimately determine that the methods of peak standing crop, Milner and Hughes, and Smalley were underestimates and that the Wiegert and Evans method was an overestimate of NAPP in tidal marsh systems. Based on its calculated turnover rate of 1.9 crops per yr, a modified Lomnicki, et al. method provided the best NAPP estimate (454 g dry wt per m² per yr).     

Secondary production within a seagrass bed (Zostera marina andRuppia maritima) in lower Chesapeake Bay

Monthly sampling of a 140-ha seagrass bed in the lower Chesapeake Bay, Virginia, revealed that 13 numerically and trophically important species, representing about 20% of the total community densities over the year-long study period, accounted for the production of ≈42 g dry wt m^?2 yr^?1. This estimate is likely conservative due to our assumptions on voltinism and fixed size at maturity regardless of season for the species studied. The isopodErichsonella attenuata accounted for 17.6 g dry wt m^?2 yr^?1 or 42% of the calculated total production, while the portunid decapodCallinectes sapidus and the amphipodGammarus mucronatus each accounted for 7.7 g dry wt m^?2 yr^?1. The 10 remaining species (4 peracarids, 4 molluscs, and 2 decapods) each produced less than 2 g dry wt m^?2 yr^?1. Total seagrass-associated secondary production was estimated to equal or exceed 200 g dry wt m^?2yr^?1. By applying this estimate to the entire 140-ha grassbed, we projected that, on average, 4.8 metric tons dry wt of invertebrate standing stock and 55.9 metric tons of invertebrate production occur over the year.     

The effect of chronic oil pollution on salt-marsh nitrogen fixation (acetylene reduction)

Annual acetylene reduction rates associated with interidal communities in a chronically oil polluted Virginia salt marsh were compared to rates measured in an undisturbed marsh. Chronic oil treatment resulted in visible damage to the higher plants of theSpartina alterniflora zones; however, vegetation-associated acetylene, reduction was not different from the untreated control. Sediment rates generally were affected little by oil application, except during the summer when rates in the median tidal elevation zones were considerably higher than those of the control. Acetylene reduction occurred in all transects, each of which extended from upper mudflat to theSpartina patens zone. Intertidal sediment acetylene reduction was patchy, both spatially and seasonally. Estimated rates were greatest near the surface; free-living bacterial N₂ fixation activity averaged 2.23 mg N per m² per d (range=undetectable to 365 mg N per m² per d) in the untreated and 3.17 mg N per m² per d (range=undetectable to 564 mg N per m² per d) in the oil-treated marsh during the year. Vegetation-associated N₂ fixation activity yielded highest overall mean rates (156 mg N per m² per d). The seasonal pattern of sediment and vegetation-associated fixation may be controlled by temperature and availability of oxidizable substrates.     

Loading, Transformation, and Retention of Nitrogen and Phosphorus in the Tidal Freshwater James River (Virginia)

A nutrient mass balance for the tidal freshwater segment of the James River was used to assess sources of nutrients supporting phytoplankton production and the importance of the tidal freshwater zone in mitigating nutrient transport to marine waters. Monthly mass balances for 2007–2010 were based on riverine inputs, local point sources (including sewer overflow events), ungauged inputs, riverine outputs, and tidal exchange. The tidal freshwater James River received exceptionally high areal loads (446 mg TN m^?2 day^?1 and 55 mg TP m^?2 day^?1) compared to other estuaries in the region and elsewhere. P inputs were principally from riverine sources (84 %) whereas point sources contributed appreciably (54 %) to high N loads. Despite high loading rates and short water residence time, areal mass retention was high (143 mg TN m^?2 day^?1 and 33 mg TP m^?2 day^?1). Retention of particulate fractions occurred during high discharge, whereas dissolved inorganic fractions were retained during low discharge when chlorophyll-a concentrations were high. On an annualized basis, P was retained more effectively (59 %) than N (32 %). P was retained by abiotic mechanisms via trapping of particulate forms, whereas N was retained through biological assimilation of dissolved inorganic forms. Results from a limited suite of stable isotope determinations suggest that DIN from point sources was preferentially retained. Combined inputs from diffuse and point sources accounted for only 20 % and 36 % (respectively) of estimated algal N and P demand, indicating that internal nutrient recycling was important to sustaining high rates of phytoplankton production in the tidal freshwater zone.     

Algal mat productivity: Comparisons in a salt marsh

This paper documents the role of salt marsh algal mats in the productivity of a southern California tidal wetland. The productivity of the mats, which are composed of filamentous bluegreen and green algae and diatoms, varies both temporally and spatially in relation to tidal inundation and overstory vegetation. The estimates of net primary productivity (NPP) were highest under the canopy ofJaumea carnosa (Less.) Gray (341 g C m^?2 yr^?1) at low elevation. Elsewhere, NPP appeared to be limited by low light (276 g C m^?2 yr^?1 underSpartina foliosa Trin.) and desiccation (185 g C m^?2 yr^?1 underBatis martima L. and 253 g C m^?2 yr^?1 underMonanthochloe littoralis Engelm). Algal NPP was from 0.8 to 1.4 times that of the vascular plant overstory NPP. It is hypothesized that the arid environment of southern California and resulting hypersaline soils reduce vascular plant cover, which leads to high algal productivity.     

Organic matter fluxes and marsh stability in a rapidly submerging estuarine marsh

We studied organic matter cycling in two Gulf Coast tidal, nonsaline marsh sites where subsidence causes marine intrusion and rapid submergence, which mimics increased sea-level rise. The sites experienced equally rapid submergence but different degrees of marine intrusion. Vegetation was hummocked and much of the marsh lacked rooted vegetation. Aboveground standing crop and production, as measured by sequential harvesting, were low relative to other Gulf CoastSpartina patens marshes. Soil bulk density was lower than reported for healthyS. alterniflora growth but that may be unimportant at the current, moderate sulfate levels. Belowground production, as measured by sequential harvesting, was extremely fast within hummocks, but much of the marsh received little or no belowground inputs. Aboveground production was slower at the more saline site (681 g m^?2 yr^?1) than at the less saline site (1,252 g m^?2 yr^?1). Belowground production over the entire marsh surface averaged 1,401 g m^?2 yr^?1 at the less saline site and 585 g m^?2 yr^?1 at the more saline site. Respiration, as measured by CO₂ emissions in the field and corrected for CH₄ emissions, was slower at the less saline site (956 g m^?2 yr^?1) than at the more saline site (1,438 g m^?2 yr^?1), reflecting greater contributions byS. alterniflora at the more saline site which is known to decompose more rapidly thanS. patens. Burial of organic matter was faster at the less saline site (796 g m^?2 yr^?1) than at the more saline site (434 g m^?2, yr^?1), likely in response to faster production and slower decomposition at the less saline site. Thus vertical accretion was faster at the less saline site (1.3 cm yr^?1) than at the more saline site (0.85 cm yr^?1); slower vertical accretion increased flooding at the more saline site. More organic matter was available for export at the less saline site (1,377 g m^?2 yr^?1) than at the more saline site (98 g m^?2 yr^?1). These data indicated that organic matter production decreased and burial increased in response to greenhouse-like conditions brought on by subsidence. *** DIRECT SUPPORT *** A01BY069 00016     

Growth dynamics of cordgrass,Spartina alterniflora loisel., on control and sewage sludge fertilized plots in a Georgia salt marsh

Seasonal plant growth dynamics were followed for a year in undisturbed plots of tall and short formSpartina alterniflora Loisel. and in plots of short formS. alterniflora which were enriched with sewage sludge at a rate of 100 g dry sludge m^?2wk^?1, corresponding to a nitrogen enrichment of 2 g N m^?2wk^?1. Monthly determinations of aboveground live and dead biomass, density of live stems, the ratio of number of young shoots to total number of shoots, and belowground mass of macro-organic matter to a depth of 30 cm were made for each area. Sludge fertilization increased the live biomass of the short formS. alterniflora by up to 150% of the control live biomass, but had little effect on the dead biomass, stem density, or proportion of young shoots. There was a trend of increased amount of belowground macro-organic matter in fertilized compared to control plots during the last 6 months of the study. In all areas, there was a marked decrease in the proportion of young shoots from winter to early summer, followed by a rapid increase in the percent of young shoots from late summer to fall. Sampling of plots 7 and 20 months after termination of sludge enrichment showed higher plant biomass and % N content in surface soils, but no difference in N content of live plant tissue, in fertilized compared to unfertilized marsh. After 20 months, about half of the sludge nitrogen remaining in the soils of the fertilized plots had disappeared.     

Evapotranspiration in tidal marshes

Field experiments were completed to determine patterns of evapotranspirative water loss from salt and tidal freshwater marshes in Virginia. Water losses from “Mariotte systems” attached to open-water lysimeters and lysimeters vegetated by dominant marsh macrophytes were used to calculate hourly rates of open-water evaporation (Eo) and evapotranspiration (ET), respectively, during low tide. In the tidal freshwater marsh, ET was significantly greater than Eo (p=0.002, n=6); in the salt marsh, there were no differences between mean rates of ET and Eo (p=0.200, n=3). The ratio ET:Eo was highly correlated with leaf area index (LAI) (r²=0.82). In the tidal freshwater marsh, the amount of water loss due to plant transpiration was partitioned from total evapotranspiration by covering the water surface of the lysimeters with styrofoam beads. Measured transpiration rates in the tidal freshwater marsh were strongly correlated with leaf area index according to the following linear regression equation: T=0.355(LAI)?0.084 (r²=0.797, n=10). Because LAI was shown to be a good predictor of the relative increase in ET over Eo, it is likely that in vegetated tidal freshwater marshes with high leaf densities most atmospheric water loss comes from plants, not from the surface of the marsh. In salt marshes, low plant densities do not contribute substantially to atmospheric water loss, suggesting that paths of water transport and patterns of solute concentration in the subsurface environment are different compard to the tidal freshwater marsh.     

The Cananéia Lagoon estuarine system,São Paulo,Brazil

The Cananéia Lagoon estuarine system lies at 25°S, near the latitudinal limit for mangroves. It is 110 km long, consisting of 1–3 km wide channels behind a barrier island, with narrow inlets at the southern and northern ends. Average and maximum depths are 6 m and 12 m. The system is microtidal and subtropical. Mean annual temperature is 21.4°C (annual amplitude=7.0°C). When the area receives sporadic frosts, temperatures close to 2°C occur in the estuary. Annual precipitation (2,270 mm) exceeds annual potential evapotranspiration (1,656 mm). The water budget of the 1,339 km² watershed is controlled primarily by local rainfall. Before 1978, a large river discharged a significant portion of its flow into the lagoon, but closure of the diversionary channel has since caused changes in salinity, phytoplankton populations, and mangrove coverage. About 90 km² of intertidal habitat is occupied by mangroves and tidal marsh; mangroves are dominant. Fringe and riverine forests (dominated byRhizophora) are more structurally developed than the basins dominated byLaguncularia and have higher litterfall rates (2.08 g m^?2 d^?1, fringes; 1.04 g m^?2d^?1, basins). Primary production exhibits pronounced seasonal pulses; heterotrophic processes lag photosynthetic production and are partially driven by particulate matter inputs. Synthetic models must consider the spatial and temporal heterogeneity of this region.     

Bivalve contribution to benthic metabolism in a California Lagoon

Populations of suspension feeding bivalves constitute a metabolically important component of the benthos in Colorado Lagoon, Long Beach, California. Oxygen and nutrient flux were measuredin situ at monthly intervals over a two-year period. Estimates of bivalve metabolism were based on regressions of oxygen and ammonium flux on a measure of clam biomass adjusted allometrically. The introduced bivalve,Mercenaria mercenaria, occurs at maximum densities exceeding 400 per m². Based on mean densities (143 per m²), bivalves contributed more than 50% of the 77 mg O₂ per m² per hr mean annual oxygen uptake and the 191 μM per m² per hr mean ammonium release. Although bivalve biomass was not correlated with other inorganic nutrient flux, on an annual basis the sediments were a source of phosphate (26 μM per m² per hr annual average, range 5 to 50) and a small sink for nitrate and nitrite. Net primary production, ammonium flux, and phosphate flux showed great annual variability while respiration was relatively constant. Total community oxygen uptake was not correlated with temperature. Long term incubations revealed no obvious rhythms having a period between four hours and four days. The benthic flux of ammonium and phosphate was calculated to supply more than the annual requirement of Lagoon phytoplankton.