Published online 23 May 2006
Published in Soil Sci Soc Am J 70:1242-1251 (2006)
DOI: 10.2136/sssaj2003.0339
© 2006 Soil Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA
Wetland Soils
Hydrologic and Vegetation Effects on Water Column Phosphorus in Wetland Mesocosms
J. R. Whitea,*,
K. R. Reddyb and
J. Majer-Newmanc
a Wetland Biogeochemistry Institute, Dep. of Oceanography and Coastal Sciences, Louisiana State Univ., Baton Rouge, LA 70803
b Soil and Water Science Dep., Univ. of Florida, Box 110510, Gainesville, FL 32611-0510
c Everglades Division South Florida Water Management District, West Palm Beach, FL
* Corresponding author (jrwhite{at}lsu.edu)
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ABSTRACT
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Historic phosphorus (P) loading from agricultural areas has been identified as one of the major causes for ecological changes occurring in the Florida Everglades. The restoration plan for the Everglades includes construction of large stormwater treatment areas (STAs) to intercept and treat this relatively high nutrient water down to very low total P (TP) concentrations. One such STA has been in operation for approximately 10 yr and contains both emergent aquatic vegetation (EAV) and submerged aquatic vegetation (SAV) communities. The surface water TP concentrations in areas near the outflow range from 0.02 to 0.05 mg TP L1. To simulate these areas, we investigated the interaction of vegetation type; EAV or SAV; and hydrology; continuously flooded or periodic drawdown; on the P removal capacity in mesocosms packed with peat soil obtained from STA-1W. The surface water had low TP concentrations with an annual mean = 0.023 mg L1. For SRP and TP, hydrologic fluctuations alone had no discernable impact on P treatment while vegetation type showed a significant impact. Influent soluble reactive P (SRP) decreased by 49% for the SAV treatments compared with 41% for the EAV treatments, irrespective of hydrology treatment. The reduction of dissolved organic P (DOP) was also higher for the SAV treatment averaging 33% while showing a reduction of 11% for the EAV treatments. There was no significant difference in the treatment efficiency of particulate P (PP) across the treatments. For TP, SAV treatments removed 45% of TP while EAV removed significantly less at 34%. By mass calculations, the EAV required 85% more P for plant growth than was removed from the water column in 1 yr compared with only 47% for the SAV. Therefore, the EAV "mined" substantially more P from the relatively stable peat soil, translocating it into the detrital pool.
Abbreviations: DOP, dissolved organic phosphorus EAV, Emergent aquatic vegetation PP, particulate phosphorus SAV, submerged aquatic vegetation SFWMD, south Florida water management district SRP, soluble reactive P STA, stormwater treatment area TP, total P
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INTRODUCTION
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PHOSPHORUS RETENTION by constructed wetlands may include the following processes: surface adsorption on soil minerals, precipitation reactions, microbial immobilization, and plant uptake (Reddy et al., 1995). These processes may be combined into two distinct P retention pathways for wetlands: sorption and burial (Reddy et al., 1999). Phosphorus sorption includes both adsorption and precipitation reactions as mechanisms for the removal of phosphate from the soil solution to the solid phase. As plants senesce, some of the P contained in detrital tissue can be recycled within the wetland, and released into the water column. Remaining refractory detrital tissue may eventually become incorporated as organic matter in the wetland soil profile as organic matter accretes.
Accretion of organic matter has been reported as a major sink for P in wetlands (Craft and Richardson, 1993; Reddy et al., 1993). Wetlands tend to accumulate organic matter due to the production of detrital material from biota and experience relatively low rates of decomposition under flooded conditions (DeBusk and Reddy, 1998). Soil accretion rates for constructed wetlands are on the order of millimeters per year, although accretion rates in productive natural systems such as the Everglades have been reported as high as 1 cm or more per year (Craft and Richardson, 1993; Reddy et al., 1993). Over time, productive constructed wetland systems will accumulate organic matter that has different physical and biological characteristics than the original preconstruction soil. Eventually, this new material settles and compacts to form new soil, which may exhibit a different P removal capacity than the original soil.
Phosphorus accretion increases with P loading to the wetland (Reddy et al., 1993). However, an increase in accretion does not assure low surface water outflow P concentrations, especially for intermittently flooded wetland systems where decomposition of organic detritus releases available P back into the water column. Decomposition of detrital material was found to increase under high P conditions (DeBusk and Reddy, 2003, Wright and Reddy, 2001) lower water levels (White and Reddy, 2000), and higher redox conditions (White and Reddy, 2001).
Scientific investigations of P reductions in constructed wetlands generally focus on wetlands receiving much higher inflow concentrations (>0.100 mg L1) than this study. The goal of this study was to investigate the P treatment capacity of EAV and SAV communities under both continuously flooded and under periodic drawdown with a mean inflow TP concentration of 0.023 mg L1.
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STUDY AREA
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The Everglades is an internationally recognized oligotrophic ecosystem and more than half of the original 1.17 million ha has been lost to drainage and development (Davis and Ogden, 1997). Today, the Everglades wetland ecosystem is comprised of three Water Conservation Areas (WCAs) and the Everglades National Park (Fig. 1
). These areas are being negatively impacted by hydrologic changes and nutrient-rich runoff generated from urban and agricultural sources (Davis and Ogden, 1997).

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Fig. 1. View of south Florida showing the relationship of the Everglades Agricultural Area, the Water Conservation Areas (northern Everglades), the Everglades National Park and the Stormwater Treatment Areas 1-W.
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Predrainage estimates of nutrient loading, reconstructed from written records and paleoecological assessments made in nearly pristine areas of the Everglades, have indicated that the Everglades flora and fauna had adapted to a very low nutrient regime (McCormick et al., 2001). Contemporary monitoring has shown that TP concentrations in south Florida rainfall is generally <0.010 mg L1, and water column samples from the relatively unimpacted areas have TP concentrations between 0.004 and 0.010 mg L1 (McCormick et al., 2001). Over time, increased nutrient loading to a system adapted to extremely low nutrient conditions, resulted in a decrease in the native periphyton assemblage and a shift from a sawgrass dominated emergent marsh to a cattail dominated system (Davis, 1991).
In 1994, the State of Florida enacted the Everglades Forever Act (EFA) (Section 373.4592, Florida Statutes) that mandates both hydrologic modifications and nutrient reduction to protect the remaining Everglades. As part of the nutrient reduction program, the EFA requires the South Florida Water Management District (SFWMD) to construct a series of large treatment wetlands (about 17 000 ha) called STAs to reduce nutrient levels in runoff to a design target of 0.05 mg TP L1 before discharging into the Everglades. The EFA also requires the SFWMD to conduct research to optimize nutrient removal performance by the STAs in an effort to produce outflow concentrations less than the design target, and provide operational guidance to maintain and improve the long-term P retention of the STAs.
These large STAs have been designed as passive, wetland removal treatment systems, primarily dominated by emergent and submerged vegetation. The maximum and minimum standard operating water depths ranges between a low of 0.15 m to a high of 1.22 m, depending on dominant vegetation and watershed runoff volumes. However, with increased drainage and development of south Florida, the pressure for freshwater supplies during the seasonal dry periods and prolonged droughts may reduce the volume of runoff water available to the STAs, resulting in a temporary dry out of the system and potential release of P from the system. A significant increase in the mean TP outflow concentration could result from climatic and hydrologic conditions depending on the degree and extent of P flux (Olila et al., 1997; White et al., 2004).
These effects may be even more critical in areas near the outflows, resulting in export of P. Depending on the degree and extent of the P flux, a significant increase in the mean TP outflow concentration from the STA could result. We conducted a controlled mesocosms study to determine the effects of water level drawdown and rehydration and vegetation type (EAV vs. SAV) to better understand the potential effects of dry-out and vegetation type on STA P removal performance. There is a distinct paucity of information on the detailed P dynamics at very low (0.0100.10 mg L1) TP concentrations. Therefore, this study provides critical information on the detailed P dynamics at low P concentrations typical of the surface water P concentration proximal to the outflow of the STA.
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MATERIALS AND METHODS
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Experimental Design
The mesocosms were situated at the surface water outflow at the south end of STA-1W, a 2700-ha constructed wetland built on previously farmed agricultural land located 25 km west of the city of West Palm Beach in Palm Beach County. The site borders the northwest corner of WCA-1 (260 38' N and 800 25' W) situated proximal to the outflow of STA-1W (Fig. 1). Mesocosms consisted of 12 fiberglass-lined plywood tanks measuring 5.9 m long by 1.0 m wide by 1.0 m deep. Each mesocosm contained 30 cm of previously farmed peat taken from STA-1W, Cell 5, before its construction. The soil was overlaid with 40 cm of STA-1W outflow water and the mesocosms were open at the top, exposed to direct sunlight and precipitation. STA-1W outflow water was pumped from the outflow canal into a head tank located at the site and then gravity fed into a PVC distribution system. The flow-through mesocosm system operated with an average design hydraulic loading rate (HLR) of 2.61 cm d1 that resulted in a nominal hydraulic retention time of 15.4 d. The HLR was controlled through calibrated pipette tips that were replaced weekly.
There were 12 mesocosms, six of them planted with 32 Typha sp. plants and six were not planted. The mesocosms were flooded at the HLR previously described. The following treatments were evaluated in triplicate:
- Continuously Flooded with EAV: (planted with 32 mature cattail plants)
- Periodic Drawdown with EAV: (planted with 32 mature cattail plants and exposed to two 1-mo drawdown periods)
- Continuously Flooded with SAV: (emergent vegetation prevented from colonizing the tanks while the growth of SAV was permitted)
- Periodic Drawdown with SAV: (emergent vegetation prevented from colonizing the tanks while the growth of SAV was permitted and exposed to two 1-mo drawdown periods).
The mesocosms began receiving flow-through water in January 2000 and were allowed to stabilize for 2.5 mo before the initial soil sampling. The drawdown event involved draining of the surface water from six of the tanks (three EAV and three SAV) on 30 Mar. 2000, and direct precipitation was the only water received for 1 mo. The tanks were reflooded with STA-1W outflow water on 8 May 2000 and remained hydrated until 30 Aug. 2000, when they were again drained and allowed to dry-out for about 1 mo. The tanks were flooded on 2 Oct. 2000, and monitored until 31 Mar. 2001. Continuously flooded mesocosms received a constant flow of surface water during this period and were not subject to any fluctuations in water level. In both the continuously flooded and periodic drawdown treatments, the water depth was maintained at 40 cm when flooded.
Inflow and outflow grab water samples were taken twice weekly at each tank beginning in January 2000 and continued through March 2001. Soluble reactive P was determined colorimetrically within 48 h of collection (Method 365.1; USEPA, 1993). Water samples analyzed for total dissolved P (TDP) and TP were determined colorimetrically (Method 365.1; USEPA, 1993) after autoclave digestion. Particulate P was calculated by subtracting TDP from TP and DOP was calculated by subtracting SRP from TDP. Physical parameters measured weekly in the field included dissolved oxygen, temperature, and pH.
Vegetation Sampling and Analyses
A single randomly selected cattail was reserved for nutrient analysis from each of the EAV treatment mesocosms at project inception. However, no SAV were sampled at this time because these communities had not become established. After 1 yr of operation, a 0.25-m2 quadrat was used to collect the aboveground vegetation at two randomly located sites within each tank for both the EAV and SAV treatments. The plant material was dried, weighed, and ground before analyses. The roots were collected from the soil coring procedure listed below. Samples were analyzed for total C, and total N using a Carlo-Erba NA-1500 CNS Analyzer (Haak-Buchler Instruments, Saddlebrook, NJ), and TP was determined from a total Kjeldahl digestion and analyzed colorimetrically for P (Method 365.1; USEPA, 1993).
Soil Sampling and Analyses
Duplicate soil cores were collected from each mesocosm by driving a 10-cm diam. aluminum tube to the bottom of each tank for both the initial soil sampling and after 1 yr. The depth of the soil was noted; the core was sealed, and removed. Each core was sectioned into two intervals; 0 to 5 cm and 5 cm until the bottom of the mesocosm (530 cm). The core sections were extruded into a labeled plastic bag, sealed, and stored on ice until return to the laboratory the following day. All roots and rhizomes were removed, dried, weighed and digested for TP determination. The soil samples were weighed, homogenized, and transferred to 2-L polyethylene storage containers and refrigerated at 4°C until analysis. Soil redox (EH) was measured each week at the 5- and 10-cm depths using permanently installed platinum electrodes.
Gravimetric moisture content was determined on subsamples by weight percent change of the initial soil samples and samples dried at 70°C until constant weight. Bulk density was calculated for each soil core on a dry weight basis. Total C and N content of detritus and soils was determined on dried, ground samples using a Carlo-Erba NA-1500 CNS Analyzer (Haak-Buchler Instruments, Saddlebrook, NJ). Total P concentrations were determined on dried, ground subsamples by ashing at 550°C and subsequent hydrochloric acid digestion (Anderson, 1976) followed by analysis of P by an automated ascorbic acid method (Method 365.4, USEPA, 1993).
The soils were also analyzed for various inorganic P fractions following the sequential inorganic P fractionation scheme developed for Histosols (Reddy et al., 1998). The following P fractions were determined from field moist soils (0.5 g dry weight equivalent):
- 1.0 M KCL-Pi representing labile P;
- 0.1 M NaOH-Pi representing Fe and Al bound P;
- 0.1 M NaOH-Po representing fulvic and humic bound P;
- 0.5 M HCL-Pi representing Ca and Mg bound P;
- residual P representing refractory organic P.
Mass Balance of Phosphorus
The soil component was determined from the TP concentration of the soil multiplied by the mass of soil in the mesocosms. The P mass within the area of the core was normalized to a square meter. The aboveground and belowground biomass was collected in two locations within a 25 cm by 25 cm quadrat. The collective dry mass of plants collected within the total 50 cm by 50 cm area, multiplied by the TP concentration, normalized to a m2 basis and reported as g m2. The fourth component was the mass of TP in both the inflow and outflow water determined by sum and was calculated by multiplying the concentration of P by the total hydraulic loading at each time step (week) and dividing by the area of the mesocosms to obtain g P m2 yr1.
Data Analysis
Soil characteristics were statistically related using Pearson's product moment correlation and regression analysis. Analysis of variance (ANOVA) and Fisher's least significance difference (LSD) tests were used to compare the treatments. A statistical comparison was made for outflow concentrations for SRP, DOP, PP, and TP for each of the treatments. Concentration and percentage data was calculated at each time step and means and standard deviations were determined over the year long experiment.
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RESULTS AND DISCUSSION
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Soil Characteristics
Soil characteristics were determined for both the 0- to 5-cm soil intervals and the subsurface intervals (530 cm). Dry weight bulk density averaged 0.306 g cm3 in the surface soil interval and was slightly higher at 0.369 g cm3 in the subsurface soil interval (Table 1). The dry weight bulk densities are somewhat higher than expected for this organic soil and were due to significant bits of limestone incorporated in the soil. The limestone bedrock is close to the surface of the soil in the area where the soil was excavated. The higher bulk densities in the subsurface were also reflected in average moisture contents of 68 vs. 74% in the surface interval. The total C and N content of the soil averaged 260 and 13.7 g kg1 with no significant differences in the surface and subsurface soil intervals. The TP of the surface soil interval was 232 and 226 mg kg1 in the subsurface soil interval with no significant difference with depth (Table 1). There was no significant difference in any soil parameters from the initial sampling and the sampling at the end of 1 yr.
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Table 1. Soil physiochemical properties including total C, N, P, bulk density, and moisture content from the soil samples collected from the mesocosms at Year 1. Values are means ± one standard deviation (n = 3).
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Soil Phosphorus Forms
On average, the soil contained 62% organic (Table 2) vs. 38% inorganic P (Table 2). The inorganic P fractions were comprised of KCL-extractable P, which is the total of the porewater P and exchangeable P. This most available fraction comprised only 0.01% of total soil P. The Fe-Al bound P comprised 1.6% of TP and the final inorganic component was the Ca-Mg bound P 36.5% of TP. The organic P fractions include the moderately labile humic and fulvic P, which collectively made up 6.9% of TP while the residue P represented all fractions not extracted with salt, acid, or base and is generally composed of recalcitrant organic compounds comprised 55%.
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Table 2. Organic and inorganic P fractions of soils collected from the mesocosm study at Year 1. Values are means ± one standard deviation (n = 3).
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In total, the largest pools of P in this soil were the residual organic P and the Ca-Mg bound P (Table 2) in concert making up 91.5% of the TP. The Ca-Mg bound P is relatively stable in these soils due to the pH (>7.0). However, anaerobic decomposition processes and accumulation of organic acids can potentially solubilize this pool of P.
Vegetation
The mesocosms with EAV were dominated by Typha sp. while the SAV treatments were colonized by, predominately, Chara chara. and Hydrilla verticillata. Plant tissue from the SAV treatments averaged 221 and 202 g C kg1 and 6.68 and 9.24 g N kg1 for the continuously flooded and periodic drawdown treatments, respectively. Plant tissue from the EAV treatments averaged 383 and 384 g C kg1 and 5.46 and 7.08 g N kg1 for the continuously flooded and periodic drawdown treatments, respectively. The TP of the plant tissue for the SAV treatment averaged 249 mg P kg1 with a higher average for the EAV treatments at 365 mg P kg1. The C/P ratio of the EAV was 1063:1 with a lower ratio of 851:1 for the SAV. There was no statistical difference in plant variables related to hydrologic treatments.
Water Quality
The mesocosm inflow characteristics are similar to the outflow surface waters of STA 1W. The inflow water contained, on average, SRP concentrations of 0.010 mg L1, and TP concentrations of 0.024 mg L1 over the year (Table 3). The inflow water TP concentrations ranged from a low of 0.013 mg L1 to a high of 0.044 mg L1 during the experimental period. Concentrations of SRP also varied over the year ranging from 0.005 to 0.017 mg L1 (Fig. 2
).
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Table 3. Mean annual inflow and outflow concentrations of soluble reactive P (SRP), dissolved organic P (DOP), particulate P (PP), and total P (TP) and percent reductions over the inflow concentration. Data are means ± one standard deviation for replicate mesocosms (n = 3).
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Fig. 2. Soluble reactive P concentrations for surface waters entering (inflow) and discharging (outflow) the mesocosms for the continuously flooded treatments containing submerged aquatic vegetation (SAV) and emergent aquatic vegetation (EAV).
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The outflow concentration for the continuously flooded treatments had a tendency to fluctuate along with the concentration of the inflow waters (Fig. 2). This same pattern was also seen in the treatments which underwent periodic drawdown (Fig. 3
). However, these treatments also experienced increased outflow concentrations for several weeks after the reflood of the first drawdown event.

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Fig. 3. Soluble reactive P concentrations for surface waters entering (inflow) and discharging (outflow) the mesocosms for the periodic drawdown treatments containing submerged aquatic vegetation (SAV) and emergent aquatic vegetation (EAV).
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Total P concentration did not covary with changes in inflow TP concentration unlike the pattern seen with SRP (Fig. 4
and 5)
. The continuously flooded-SAV treatment had outflow concentration of 0.010 mg L1 or lower 30% of the time while the continuously flooded-EAV treatment did not reach below 0.010 mg L1 at any time during the year (Fig. 4). For either vegetative treatment (EAV vs. SAV), the outflow TP concentrations were higher than the inflow water for up to 3 wk after the reflooding for the first drawdown event, suggesting the soil and plants were acting as a source of TP to the water column (Fig. 5). There was a small effect of higher outflow concentrations from the second drawdown event on the EAV treatment, which was not apparent in the SAV treatment (Fig. 5).

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Fig. 4. Total P (TP) concentrations for surface waters entering (inflow) and discharging (outflow) the mesocosms for the periodic continuously flooded treatments containing submerged aquatic vegetation (SAV) and emergent aquatic vegetation (EAV). Shown are the Interim TP standard and the Draft (in the current rulemaking process) TP standard for discharge waters.
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Fig. 5. Total P (TP) concentrations for surface waters entering (inflow) and discharging (outflow) the mesocosms for the periodic continuously flooded treatments containing submerged aquatic vegetation (SAV) and emergent aquatic vegetation (EAV). Shown are the Interim TP standard and the Draft (in the current rulemaking process) TP standard for discharge waters.
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To investigate the mechanisms that may be responsible for P removal or release to the water column, we elucidated experimental effects on the individual P components in the water. For example, TP contains SRP, DOP, and PP fractions. The reduction of SRP in the water column is mediated by either uptake by microbes and plants and incorporated into organic forms while precipitation with calcium carbonate compounds provides for an inorganic removal mechanism.
There was no significant difference in the reduction of SRP in the SAV treatments based on concentration, whether or not periodic drawdown was imposed (Table 3). Both the drawdownEAV treatment at 42% reduction and the continuously floodedEAV treatment at 39% performed statistically worse than the SAV treatments (49%) for the outflow SRP concentrations (Table 3). The data suggests that vegetation type, more than the hydrologic fluctuations, controlled removal efficiency of SRP from the water column for these inflow P concentrations.
Decomposition processes of detrital or soil organic matter can influence the release of DOP and may have liberated the DOP from the organic detrital material or microbial pool. In the case of DOP, the continuously flooded SAV treatment had the statically best removal of DOP at 41%. Both drawdown treatments and the continuously flooded EAV treatment had lower removal efficiency for DOP ranging from 24 to 10%. While the mean percentage reduction was lower by over one-half for the EAV treatments compared with the SAV treatments, the EAV had a higher variability that likely affected statistical comparisons (Table 3). Conversion of DOP to SRP is mediated by enzymatic hydrolysis (Pant and Reddy, 2001). The DOP fraction, based on percentage removal, appears to be the most difficult fraction for treatment in terms of percentage reductions in these systems.
Treatment for PP, which is a settling process, was not significantly different for each of the four treatments, due in large part to high variability. Mean reductions ranged from a 48% down to 38% (Table 3).
Total P reductions are perhaps of most critical importance in terms of meeting the water quality standards for the Everglades. The highest average reduction was in the two SAV treatments; for the flooded at 0.013 mg L1 with a 47% reduction and for the drawdown at 0.014 mg L1 with an overall reduction of 42%. Significantly lower was the drawdown-EAV treatment averaged 0.015 mg L1 while the continuously floodedEAV averaged 0.016 mg L1 with reductions of 36 and 33%, respectively (Table 3).
The drawdown events, which appeared to have a much smaller effect on the reduction of TP of the surface water than vegetation type, did not have identical effects on the soil redox. The first drawdown period had a high EH (Fig. 6
), usually associated with drier soil conditions, which led to a release of SRP and TP for several weeks after the reflooding event, while this effect was not apparent after the second drawdown (Fig. 3 and 5). The reason for the difference in average EH values during the drawdown period could be related to rainfall patterns. The first drawdown occurred in the late winter, which is the dry period in south Florida while the second dry down period occurred in the summer when precipitation is at the highest levels. Since the mesocosms were not covered, to expose these systems to the changing climatic conditions seen over the year, the effect of the differential in precipitation likely maintained the soil moisture at a higher level in the summer than during the winter (Fig. 6). When other experimental mesocosms were maintained for 2 yr, the same pattern of higher redox in the winter drawdown was observed (White et al., 2004).

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Fig. 6. Redox reading at 5 cm in the soil for the periodic drawdown and continuously flooded treatments in the mesocosms.
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Mass Balance of Phosphorus
There were five major components of the mass balance of P calculated for this study including soil, aboveground biomass, belowground biomass, inflow TP, and outflow TP. There were only very fine root mass for the SAV treatment tanks that comprised < 0.5% of the total vegetation wet weight and were considered insignificant and not analyzed.
The largest component was the soil TP values averaging 21.7 g P m2 for the 0- to 30-cm depth (Table 4). The SAV treatment produced significantly less biomass P for both the periodic drawdown and continuously flooded treatments combined when considering only the above ground biomass at 0.121 g P m2 while the EAV treatments aboveground biomass averaged 0.214 g P m2. When you include the substantial belowground biomass for the EAV treatment, the total biomass P value rises to 0.348 g P m2. On average, all treatments removed 0.057 g P m2 from the water column (Table 4).
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Table 4. Mass balance of P in various components in the mesocosms after 1 yr. Values are means ± one standard deviation (n = 3).
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The amount of above and belowground biomass P in the emergent treatments could lead to a decline in treatment capacity over time since P is released from detrital material into the water column. The EAV produced twice as much aboveground biomass P when compared with the SAV treatments and significantly more root-associated P (Table 4). The TP of this vegetative matter was far more than was removed from the water column, therefore exploiting the more stable P in the peat for incorporation into EAV biomass. If we assume the plants were able to take up all the P that was removed from the water column, the EAV took up 85% of their P requirement from the peat soil while the SAV would have required less at 47%. The presence of significant EAV rhizosphere influences led to increased nutrient uptake from the interstitial water and likely less uptake of nutrients from the surface water (Moore et al., 1994). The SAV generally has a slower growth rate than EAV due in part, to poor light transmission and the slow diffusion rate of CO2 in water, however, SAV have shown to reduce P concentration at similar rates as emergent vegetation (Reddy et al., 1987). The SAV have also been shown in other studies to produce small (<10%) total biomass as roots and have been shown to take nutrients directly from the water column and therefore exploiting less P from the soil (Bole and Allan, 1978). Another SAV P removal mechanism is related to photosynthetically driven elevation of water column pH and the concomitant precipitation of P out of solution as Ca-P compounds are formed (Dierberg et al., 2002).
Surface water P removal was greatest in both the continuously flooded and drawdown SAV treatments compared with both hydrologic treatments containing EAV at these low surface water P concentrations suggesting that the SAV configuration might perform better than EAV for the outflow half of these large constructed wetlands (STAs) where the surface water TP concentrations are low (1430 ppb). Generally, the wetland mesocosms provided good SRP and PP removal in these systems. There was relatively poor removal of the DOP fraction. This P fraction may be of concern due to the transport of DOP carried by surface waters into the Everglades where enzymatic hydrolysis can transform the DOP into SRP, the most bioavailable form (Reddy et al., 2005).
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CONCLUSIONS
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Removal or immobilization of P at these water column concentrations was shown to be effective under both vegetative treatments with the best performance seen for the SAV communities, irrespective of hydrologic treatment. However, there was diminished effectiveness in the short-term P removal after reflooding from each drawdown event which lasted up to 3 wk before returning to pre-event removal rates. Significant reductions were seen for SAV treatments for SRP and TP when compared with the EAV treatments with no difference seen for PP. The greatest difference in treatment was seen for DOP, with an average of 11% reduction seen for the EAV treatment while the SAV treatment reduced DOP an average of 33%, irrespective of hydrology. Taken individually, the flooded SAV treatment was significantly better than all the other treatments at 41% reduction of DOP.
There was also a significant difference in total biomass P for the type of plant community with greater biomass P in the EAV compared with the SAV (Table 4). The EAV took up at least 85% of their TP requirement from the peat soil assuming they also utilized all the P that was removed from the water column. In comparison, the SAV could have met 53% of their P requirement from utilizing all the P removed from the water column while requiring less P (47%) for biomass requirements from the peat soil. Overall, the EAV community has significantly less effect on SRP and TP removal at these low water column P concentrations and also took up more P from the peat soil than the SAV. This mobilization of P from the more stable peat soil into plant biomass could potentially lead to oxidation of this organic detrital P and the concomitant release of P into the water.
Received for publication December 18, 2005.
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