Nutrient additions by waterfowl to lakes and reservoirs: predicting their effects on productivity and water quality

Lakes and reservoirs provide water for human needs and habitat for aquatic birds. Managers of such waters may ask whether nutrients added by waterfowl degrade water quality. For lakes and reservoirs where primary productivity is limited by phosphorus (P), we developed a procedure that integrates annual P loads from waterfowl and other external sources, applies a nutrient load-response model, and determines whether waterfowl that used the lake or reservoir degraded water quality. Annual P loading by waterfowl can be derived from a figure in this report, using the days per year that each kind spent on any lake or reservoir. In our example, over 6500 Canada geese (Branta canadensis) and 4200 ducks (mostly mallards, Anas platyrhynchos) added 4462 kg of carbon (C), 280 kg of nitrogen (N), and 88 kg of P y^–1 to Wintergreen Lake in southwestern Michigan, mostly during their migration. These amounts were 69% of all C, 27% of all N, and 70% of all P that entered the lake from external sources. Loads from all external sources totaled 840 mg P m^–2 y^–1. Application of a nutrient load-response model to this concentration, the hydraulic load (0.25 m y^–1), and the water residence time (9.7 y) of Wintergreen Lake yielded an average annual concentration of total P in the lake of 818 mg m^–3 that classified the lake as hypertrophic. This trophic classification agreed with independent measures of primary productivity, chlorophyll-a, total P, total N, and Secchi disk transparency made in Wintergreen Lake. Our procedure showed that waterfowl caused low water quality in Wintergreen Lake.Contribution 824 of the National Fisheries Research Center-Great Lakes, 1451 Green Road, Ann Arbor, Michigan 48105, U.S.A. and 722 of the Kellogg Biological Station, Michigan State University.     

The influence of impoundments on nutrient budgets in two catchments of Southeastern Michigan

N and P budgets quantify inputs and outputs of nutrients at the catchment scale to allow evaluation of inputs and outputs as well as inferences about transport and processing based on unaccounted-for nutrients. N and P budgets were constructed for two catchments in southeastern Michigan with markedly different numbers of impoundments, over two years, to evaluate the influence of impoundments on nutrient fluxes from each catchment. The Huron, with 88 impoundments >10 ha, stored 156 kg P km⁻² y⁻¹, while the Raisin (with 14 impoundments) had a net export of 102 kg P km⁻² y⁻¹. The Huron catchment also stored and denitrified more N than the Raisin catchment – 2,418 kg N km⁻² y⁻¹ compared to 1,538 kg N km⁻² y⁻¹. Riverine export of N and P also varied markedly between the catchments, with the Huron River exporting 288 kg N and 7 kg P km⁻² y⁻¹ and the Raisin River exporting 1,268 kg N and 34 kg P km⁻² y⁻¹. We then re-calculated budget results from previous studies using the approach of the present study, altering input and outputs fluxes as well as system boundaries to obtain comparable budgets. For these comparable budgets, annual P outputs on average accounted for 77% of inputs whereas N outputs accounted for only 39% of N inputs. Across catchments, the percent of inputs exported by the river averaged 16% for N and 5% for P, indicating more effective retention of P than N.     

Migratory Atlantic salmon as vectors for the transfer of energy and nutrients between freshwater and marine environments

SUMMARY 1. Annual energy, carbon, nitrogen and phosphorus fluxes across the river mouth by Atlantic salmon were estimated for 18 years (1976–94) in the Norwegian River Imsa. The total energy content of the emigrating smolts in each year varied considerably with a mean value of 237 × 10³ kJ. That of returning adults also varied between years with a mean value of 141 × 10⁴ kJ. One‐sea‐winter salmon (grilse) made up 65% of the total energy content of the spawners in the river. Dead carcasses remaining in the river after spawning were estimated to have a mean annual energy content of 175 × 10³ kJ. 2. The net annual energy flux from the sea to the river varied between 48 × 10³ kJ (1987) and 152 × 10⁴ kJ (1989) with a mean of 616 × 10³ kJ, and a coefficient of variation of 67%. Average net marine import of the returning adults was 83 × 10⁴ kJ year^?1 with a coefficient of variation of 52%. Mean annual export of C, N and P to sea by the smolts was 595, 131 and 22 kg, and by kelts 1535, 352 and 70 kg, respectively, whereas gross import via the adults was 3176 kg C, 735 kg N and 132 kg P. The annual flux across the river mouth was 1046 kg C, 253 kg N and 39 kg P. The net marine import were 1585 kg C, 371 kg N and 60 kg P. The net flux was estimated at 0.2% for nitrogen and 5% for phosphorus of the total river load. 3. The energy flux caused by Atlantic salmon spawning in the River Imsa was relatively high because the general nutrient load in the river is low. Thus, even though most Atlantic salmon survive spawning, their contribution to the nutrient flux in the river is significant.     

Particulate inputs to Dublin Bay and to the South Lagoon,Bull Island

There are substantial riverine and sewage particulate inputs into Dublin Bay. The main river, the R. Liffey, accounts for almost 85% of the riverine input, which amounts to 37.1 t d^–1 of SPM. The sewage input, which is discharged into the estuary, is slightly less at 35.3 t d^–1. The combined inputs deliver 17.4 t d^–1, 2.9 t d^–1and 1.2 t d^–1of particulate C, N and P, respectively, to the Bay as a whole. Although the particulate N load accounts for just 20% of the total N input, the retention of particulates within the system, and the subsequent remobilisation of N from the sediments is heavily implicated in the macroalgal blooms. A particulate mass balance of SPM, and of C, N and P for the South Lagoon showed no consistent pattern of import/export except that the fluxes of the individual elements reflected SPM load and varied with quantity rather than quality. The molar C:N ratio was around 6:1 although on occasion values approaching 3:1 were obtained, indicating very high quality and potentially labile SPM. The controlling factors in the budget appeared to be the water velocities and to a lesser extent, the SPM load, which surprisingly was no greater on a spring tide than on a neap. In this shallow, and primarily intertidal, system, wind- or wave-driven resuspension may exert more influence than predictable factors such as tidal range.     

Seasonal oxygen,nitrogen and phosphorus benthic cycling along an impacted Baltic Sea estuary: regulation and spatial patterns

The regulatory roles of temperature, eutrophication and oxygen availability on benthic nitrogen (N) cycling and the stoichiometry of regenerated nitrogen and phosphorus (P) were explored along a Baltic Sea estuary affected by treated sewage discharge. Rates of sediment denitrification, anammox, dissimilatory nitrate reduction to ammonium (DNRA), nutrient exchange, oxygen (O₂) uptake and penetration were measured seasonally. Sediments not affected by the nutrient plume released by the sewage treatment plant (STP) showed a strong seasonality in rates of O₂ uptake and coupled nitrification–denitrification, with anammox never accounting for more than 20 % of the total dinitrogen (N₂) production. N cycling in sediments close to the STP was highly dependent on oxygen availability, which masked temperature-related effects. These sediments switched from low N loss and high ammonium (NH₄ ⁺) efflux under hypoxic conditions in the fall, to a major N loss system in the winter when the sediment surface was oxidized. In the fall DNRA outcompeted denitrification as the main nitrate (NO₃ ^?) reduction pathway, resulting in N recycling and potential spreading of eutrophication. A comparison with historical records of nutrient discharge and denitrification indicated that the total N loss in the estuary has been tightly coupled to the total amount of nutrient discharge from the STP. Changes in dissolved inorganic nitrogen (DIN) released from the STP agreed well with variations in sedimentary N₂ removal. This indicates that denitrification and anammox efficiently counterbalance N loading in the estuary across the range of historical and present-day anthropogenic nutrient discharge. Overall low N/P ratios of the regenerated nutrient fluxes impose strong N limitation for the pelagic system and generate a high potential for nuisance cyanobacterial blooms.     

The fate of nitrogen and phosphorus at the land-sea margin of the North Atlantic Ocean

Five large rivers that discharge on the western North Atlantic continental shelf carry about 45% of the nitrogen (N) and 70% of the phosphorus (P) that others estimate to be the total flux of these elements from the entire North Atlantic watershed, including North, Central and South America, Europe, and Northwest Africa. We estimate that 61 · 109 moles y^–1 of N and 20 · 10⁹ moles y^–1 of P from the large rivers are buried with sediments in their deltas, and that an equal amount of N and P from the large rivers is lost to the shelf through burial of river sediments that are deposited directly on the continental slope. The effective transport of active N and P from land to the shelf through the very large rivers is thus reduced to 292 · 10⁹ moles y^–1 of N and 13 · 10⁹ moles y^–1 of P.The remaining riverine fluxes from land must pass through estuaries. An analysis of annual total N and total P budgets for various estuaries around the North Atlantic revealed that the net fractional transport of these nutrients through estuaries to the continental shelf is inversely correlated with the log mean residence time of water in the system. This is consistent with numerous observations of nutrient retention and loss in temperate lakes. Denitrification is the major process responsible for removing N in most estuaries, and the fraction of total N input that is denitrified appears to be directly proportional to the log mean water residence time. In general, we estimate that estuarine processes retain and remove 30–65% of the total N and 10–55% of the total P that would otherwise pass into the coastal ocean. The resulting transport through estuaries to the shelf amounts to 172–335 · 10⁹ moles y^–1 of N and 11–19 · 10⁹ moles y^–1 of P. These values are similar to the effective contribution from the large rivers that discharge directly on the shelf.For the North Atlantic shelf as a whole, N fluxes from major rivers and estuaries exceed atmospheric deposition by a factor of 3.5–4.7, but this varies widely among regions of the shelf. For example, on the U.S. Atlantic shelf and on the northwest European shelf, atmospheric deposition of N may exceed estuarine exports. Denitrification in shelf sediments exceeds the combined N input from land and atmosphere by a factor of 1.4–2.2. This deficit must be met by a flux of N from the deeper ocean. Burial of organic matter fixed on the shelf removes only a small fraction of the total N and P input (2–12% of N from land and atmosphere; 1–17% of P), but it may be a significant loss for P in the North Sea and some other regions. The removal of N and P in fisheries landings is very small. The gross exchange of N and P between the shelf and the open ocean is much larger than inputs from land and, for the North Atlantic shelf as a whole, it may be much larger than the N and P removed through denitrification, burial, and fisheries. Overall, the North Atlantic continental shelf appears to remove some 700–950· 10⁹ moles of N each year from the deep ocean and to transport somewhere between 18 and 30 · 10⁹ moles of P to the open sea. If the N and P associated with riverine sediments deposited on the continental slope are included in the total balance, the net flux of N to the shelf is reduced by 60 · 10⁹ moles y^–1 and the P flux to the ocean is increased by 20 · 10⁹ moles y^–1. These conclusions are quite tentative, however, because of large uncertainties in our estimates of some important terms in the shelf mass balance.     

Retention of Sediments and Nutrients in the Iron Gate I Reservoir on the Danube River

This work addresses an intensively debated question in biogeochemical research: “Are large dams affecting global nutrient cycles?” It has been postulated that the largest impoundments on the Lower Danube River, the Iron Gates Reservoirs, act as a major sink for silica (Si) in the form of settling diatoms, for phosphorus (P) and to a lesser extent for nitrogen (N). This retention of P and N in the reservoir would represent a positive contribution to the nutrient reduction in the Danube River. Based on a 9-month monitoring scheme in 2001, we quantified the nutrient and the sediment retention capacity of the Iron Gate I Reservoir. The sediment accumulation corresponded to 5% TN (total nitrogen), 12% TP (total phosphorus) and 55% TSS (total suspended solids) of the incoming loading. A mass balance revealed that more N and P are leaving the reservoir than entering via the inflow. Based on these current results, the reservoir was temporarily acting as a small nutrient source. The nutrient accumulation in the sediments of the Iron Gate I Reservoir represents only 1% of the “missing” load of 10⁶ t N and 1.3 × 10⁵ t P defined as the difference between the estimated nutrient export from the Danube Basin and the measured flux entering the Black Sea. This result disproves the hypothesis that the largest impoundment on the Danube River, the Iron Gates Reservoir, plays a major role in N and P elimination.     

Effects of Sewage Dumping on Macrobenthic Invertebrates in the Jordan Gulf of Aqaba,Red Sea

There are two sewage outfalls along the Jordanian coastline in the Gulf of Aqaba. During 1982 and 1983 a total of 328 core samples (0.01 m² to a depth of 15 cm) were collected from the two outfalls, two control stations which resemble the outfalls in depth and sediment texture, and from two stations 100 m on both sides of each outfall. Faunal analysis revealed that the total number of individuals, number of species, species richness, and faunal similarities of macrobenthic invertebrates were lower at the sewage outfall near the phosphate loading port than the control station during both collections. At the 100 m stations, the numbers of individuals were generally higher than the sewage and control stations. The number of species, however, was highest at the control station and lowest at the sewage outfall. At the other sewage outfall, where the sewage effluent is discharged sporadically, no measurable effects on macrobenthic invertebrates were found.     

The influence of Procambarus clarkii (Cambaridae,Decapoda) on water quality and sediment characteristics in a Spanish floodplain wetland

This study describes the effects of the American red swamp crayfish, Procambarus clarkii Girard, on water quality and sediment characteristics in the Spanish floodplain wetland, Las Tablas de Daimiel National Park. Our short term enclosure study during a summer drawdown revealed that crayfish acted as a nutrient pump that transformed and translocated sediment bound nutrients to the water column. Water quality impoverishment was mainly due to the increase of dissolved inorganic nutrients (soluble reactive phosphorus and ammonia), and a significant increase of total suspended solids occurred likely as a result of crayfish associated bioturbation. At the same time, crayfish reduced the content of organic matter in the sediment and we observed a slight increase of total sediment phosphorus and nitrogen content as a result of crayfish benthic activity. P. clarkii effects, in terms of internal nutrient loading (229.91 mg TP m^–2 d^–1), were shown to be important on a local scale, indicating the significance of internal nutrient supply to water column primary producers particularly under low external supply (summer). Extrapolations to the whole ecosystem, however, revealed a negligible crayfish contribution (0.06%) to total internal nutrient loading (0.035 mg TP m^–2 d^–1). Hence, crayfish spatial heterogeneity patterns are important in global and local matter fluxes and nutrient cycles in wetlands.     

Macrophyte-related shifts in the nitrogen and phosphorus contents of the different trophic levels in a biomanipulated shallow lake

Lake Zwemlust, a small highly eutrophic lake, was biomanipulated without reducing the external nutrient loading, and the effects were studied for four years. In this paper we pay special attention to the shifts in relative distribution of nitrogen and phosphorus in the different trophic levels and to the changes in growth limitation of the autotrophs.Despite of the high external nutrient loads to the lake (ca 2.4 g P m^–2 y^–1 and 9.6 g N m^–2 y^–1), the effects of biomanipulation on the lake ecosystem were pronounced. Before biomanipulation no submerged vegetation was present in the lake and P and N were stored in the phytoplankton (44% N, 47% P), fish (33% N, 9% P) and in dissolved forms (23% N, 44% P). P and N contents in sediments were not determined. In the spring and summer following the biomanipulation (1987), zooplankton grazing controlled the phytoplankton biomass and about 90% of N and P were present in dissolved form in the water. From 1988 onwards submerged macrophyte stands continue to thrive, reducing the ammonium and nitrate concentrations in the water below detection levels. In July 1989 storage of N and P in the macrophytes reached 86% and 80%, respectively. Elodea nuttallii (Planchon) St.John, the dominant species in 1988 and 1989, acted as sink both for N and P during spring and early summer, withdrawing up to ca 60% of its N and P content from the sediment. At the end of the year only part of the N and P from the decayed macrophytes (ca 30% of N and 60% of P) was recovered in the water phase of the ecosystem (chiefly in dissolved forms). The rest remained in the sediment, although some N may have been released from the lake by denitrification.In summer 1990 only 30% of the N and P was found in the macrophytes (dominant species Ceratophyllum demersum L.), while ca 30% of N and P was again stored in phytoplankton and fish.     

Retention of nitrogen and phosphorus in a Danish lowland river system: implications for the export from the watershed

In a Danish lowland river system intensive measurements were made, in four 80 m reaches, of the nitrogen (N) and phosphorus (P) stored in the stream sediment. The results were used for calculation of the total retention in the river system during two summers (June to August). In addition, the mobilization of nutrients from the stream bottom in autumn 1987 was compared with the export from the watershed.During the study period (June 1987 to September 1988) the amounts of N and P stored in stream reaches were determined fortnightly using a core-sample technique. In reaches dominated by submersed macrophytes, 25–40 g N m^–2 and 20–30 g P m^–2 were stored during two summers, against only 10–15g N and P m^–2 for sandy and gravely reaches. In riparian zones with emergent macrophytes the retention was even higher than in the submersed macrophytes. Gross retention exceeded net retention by a factor of two to three.Net retention of P in the river system during the summer of 1987 was equal to the summer export from the watershed. On an annual basis, retention in the summer constituted 20% of the P export. In contrast, retention in the summer of 1988 amounted to 60% of the total P export during the same period (38% reduction) and 22% in comparison with the annual export. The corresponding figures for N were lower, showing reductions of 16% and 12% of the export of total N in the two summer periods, and about 1% of the annual exports.In September 1987 6.4% of the total N export and 65% of the total P export from the watershed consisted of resuspended material. In 1987 the N and P retained during the summer was almost completely resuspended during storm events during September to November.     

Long-term phytoplankton changes in a shallow hypertrophic lake,Albufera of Valencia (Spain)

A long-term phytoplankton study was carried out in the Albufera of Valencia, a shallow hypertrophic lake (surface area 21 km², mean depth 1 m, total inorganic nitrogen load 155 g m^-2 y^-1, total inorganic phosphate load 15 g m^-2 y^-1) from 1980 to 1988. The lake functions as a reservoir for the surrounding rice cultivation. From 1940's to 1988, its phytoplankton assemblage has been altered from a mesotrophic to a hypertrophic character, as consequence of the increasing pollution. For 1980–88, annual variations in the phytoplankton were less pronounced than seasonal changes. The hypertrophic and morphometric features of the lake favoured the stability of the phytoplankton assemblage and chlorophyll a levels during the study period. Seasonal and horizontal distribution of the total phytoplankton abundance and biomass were highly influenced by the hydrological cycle of the lagoon. Compared with other shallow nutrient rich lakes, the Albufera of Valencia is similar to the shallow hypertrophic lakes of the Netherlands.     

Rainfall input, throughfall and stemflow of nutrients in a central African rain forest dominated by ectomycorrhizal trees

Incident rainfall is a major source of nutrient input to a forest ecosystem and the consequent throughfall and stemflow contribute to nutrient cycling. These rain-based fluxes were measured over 12 mo in two forest types in Korup National Park, Cameroon, one with low (LEM) and one with high (HEM) ectomycorrhizal abundances of trees. Throughfall was 96.6 and 92.4% of the incident annual rainfall (5370 mm) in LEM and HEM forests respectively; stemflow was correspondingly 1.5 and 2.2%. Architectural analysis showed that ln(funneling ratio) declined linearly with increasing ln(basal area) of trees. Mean annual inputs of N, P, K, Mg and Ca in incident rainfall were 1.50, 1.07, 7.77, 5.25 and 9.27 kg ha^–1, and total rain-based inputs to the forest floor were 5.0, 3.2, 123.4, 14.4 and 37.7 kg ha^–1 respectively. The value for K is high for tropical forests and that for N is low. Nitrogen showed a significantly lower loading of throughfall and stemflow in HEM than in LEM forest, this being associated in the HEM forest with a greater abundance of epiphytic bryophytes which may absorb more N. Incident rainfall provided c. 35% of the gross input of P to the forest floor (i.e., rain-based plus small litter inputs), a surprisingly high contribution given the sandy P-poor soils. At the start of the wet season leaching of K from the canopy was particularly high. Calcium in the rain was also highest at this time, most likely due to washing off of dry-deposited Harmattan dusts. It is proposed that throughfall has an important 'priming' function in the rapid decomposition of litter and mineralization of P at the start of the wet season. The contribution of P inputted from the atmosphere appears to be significant when compared to the rates of P mineralization from leaf litter.     

Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

Effect of nutrient loading on biogeochemical processes in tropical tidal creeks

The effect of increased nutrient loads on biogeochemical processes in macrotidal, mangrove-lined creeks was studied in tropical Darwin Harbour, Australia. This study uses an integrative approach involving multiple benthic and pelagic processes as measures of ecosystem function, and provides a comparison of these processes in three tidal creeks receiving different loads of treated sewage effluent. There were significant differences in process rates between Buffalo Creek (BC) (hypereutrophic), which receives the largest sewage loads; Myrmidon Creek (MC) (oligotrophic–mesotrophic) which receives smaller sewage inputs; and Reference Creek (RC) (oligotrophic) which is comparatively pristine. Benthic nutrient fluxes and denitrification were more than an order of magnitude higher and lower, respectively, in BC and denitrification efficiency (DE) was <10%. Pelagic primary production rates were also much higher in BC but respiration exceeded primary production resulting in severe drawdown of O₂ concentrations at night. Hypoxic conditions released oxide-bound phosphorus and inhibited coupled nitrification–denitrification, enhancing benthic nitrogen and phosphorus fluxes, leading to a build-up of excess nutrients in the water column. Poor water quality in BC was exacerbated by limited tidal flushing imposed by a narrow meandering channel and sandbar across the mouth. In contrast to BC, the effect of the sewage load in MC was confined to the water column, and the impact was temporary and highly localized. This is attributed to the effective flushing of the sewage plume with each tidal cycle. Denitrification rates in MC and RC were high (up to 6.83 mmol N m^?2 day^?1) and DE was approximately 90%. This study has identified denitrification, benthic nutrient fluxes and pelagic primary production as the biogeochemical processes most affected by nutrient loading in these tidal creek systems. Physical process play a key role and the combined influence of nutrient loading and poor tidal flushing can have serious consequences for ecosystem functioning.     

Spatial and temporal variability of internal and external phosphorus loads in Mona Lake,Michigan

A study was conducted in Mona Lake, a small eutrophic lake located in western Michigan (USA) to address the temporal and spatial variability of external and internal phosphorus loading. External P load varied among subbasins, which was mostly related to discharge, but also to land use. Black Creek, which drains lands with natural cover and agriculture, accounted for the majority of flow, and total phosphorus (TP) and soluble reactive phosphorus (SRP) load, to Mona Lake. However, the relative contribution of SRP load was greater in Little Black Creek, which flows through a mostly urbanized subbasin, than in Black Creek. The relative importance of internal loading was strongly related to season, as internal TP loads contributed only ∼9% of the overall P load in April 2005, but ∼68–82% of the overall P load in the summer and early fall seasons. Internal TP and SRP loading was greater under anaerobic than aerobic conditions. Mean anaerobic TP release rates ranged from 0.80 to 15.56 mg P m⁻² d⁻¹, varying with site and season. Spatial variability in both internal phosphorus loading and sediment P concentration was also evident. By taking into account the spatial and temporal variability of different loading sources, management practices can be targeted to optimize nutrient source control strategies.     

The influence of cyanobacteria blooms on the attenuation of nitrogen throughputs in a Baltic coastal lagoon

We combined a mass balance approach with measurements of air–water and sediment–water nitrogen (N) exchange to better understand the mechanisms attenuating N throughputs in a eutrophic coastal lagoon. We were particularly interested in how seasonal shifts in external versus internal N fluxes and the transition from diatom- to cyanobacteria- dominated phytoplankton communities influence N storage and loss to the atmosphere. We found that on an annual basis almost all of the N removed by the lagoon was due to sediment storage following the spring diatom bloom. This period was characterized by high riverine inputs of dissolved inorganic nitrogen, high rates of assimilatory conversion to particulate nitrogen (PN), and net accrual of N in sediments. By contrast, the larger summer bloom was associated with low sediment N storage, which we attribute in part to the presence of positively-buoyant cyanobacteria. Low settling rates during cyanobacteria blooms favored export of PN to the Baltic Sea over sediment accrual in the lagoon. In addition, summer dinitrogen (N₂) fixation by cyanobacteria largely offset annual N₂ losses via denitrification. These findings show that cyanobacteria blooms diminish N attenuation within the lagoon by altering the balance of N exchange with the atmosphere and by promoting export of particulate N over sediment burial.     

Groundwater contribution to the nutrient budget of Tomales Bay, California

Tomales Bay, a graben structure along the San Andreas Fault, was selected for modeling ecosystem nutrient dynamics because of its linear, one-dimensional morphology and relatively pristine state. Groundwater is a potentially important term in the nutrient budget. The geologic complexities created by the San Anreas Fault, however, complicate the hydrogeology and require the area to be subdivided into three regions: granite to the west, Franciscan Formation to the east, and alluvial fill in the trough. Nutrient concentrations in the groundwater were determined through extensive well sampling; groundwater discharge was estimated using both Darcy's Law calculations and a soil moisture budget. Results indicate that groundwater discharge is of the same order of magnitude as summer streamflow into the Bay, while being significantly less than other freshwater inputs in winter. Dissolved nutrient (phosphate, nitrate + nitrite, ammonium, silica and DIC) concentrations in groundwater were consistently higher (by as much as an order of magnitude) than in surface water discharges. During the summer months, groundwater flow contributes about as much nutrient load to the bay as does streamflow. During the winter, the groundwater contribution to nutrient loading is about 20% of the streamflow contribution. Our findings indicate that groundwater is a significant component of terrestrial nutrient and freshwater loading to Tomales Bay, particularly so during the summer months. However, neither groundwater nor streamflow nutrient fluxes are large in comparison to the mixing flux at the bay mouth or the flux of N₂ gas across the air-water interface.     

Modeling the hydrochemistry of the Choptank River basin using GWLF and Arc/Info: 2. Model Validation and Application

We used the hydrochemical model GWLF to estimate terrestrial diffuse fluxes from ungauged areas of a coastal plain catchment, the Choptank River basin. The gauged area of the basin is 17% of the land surface, and we divided the remaining ungauged area into 21 subbasins. Three comparative approaches were used: (1) application of area yield coefficients based on 11 years of observations from the gauged area to extrapolate over ungauged subbasins without modeling, (2) application of GWLF to estimate export from all subbasins using model parameters calibrated in the gauged subbasin, and (3) application of GWLF with parameter adjustments based on the local characteristics in each subbasin. Comparison of the predicted export from 6 selected subbasins with observed export data showed that application of GWLF with local adjustments reduced model errors of N export from 43% to 27%. With only one adjustment for sediment P, application of GWLF alone reduced errors of P export from 92%to 40–45%, with or without local adjustments for flow and sediment retention. The data supported the hypothesis that significant spatial variations in N and P yields introduce large errors when extrapolating from gauged to ungauged subbasins, and estimated TN and TP yield coefficients varied over 1–21 kg N and 0.1–0.5 kg P ha^–1 y^–1 in ungauged areas due to varying human population densities, soil drainage characteristics, and amounts of agriculture. The most accurate estimates of terrestrial diffuse sources were combined with point source discharges and wet atmospheric inputs to estimate annual average inputs of 2.5 × 10⁶ kg N and 5.8 × 10⁴ kg P y^–1 to the Choptank estuary during 1980–1996. These results illustrate the problems of spatial extrapolation from gauged to ungauged areas and emphasize the need for application of local characteristics for accurate assessment of watershed export.