Ecology of the Ráckeve-Sorokság Danube – a review

Present paper is a review on the Ráckeve-Soroksár Danube in ecological standpoint. The goal of this study is to collect and evaluate all of available publications in that conception, concerning this Danube arm. Phytoplankton, zooplankton, macroinvertebrates, vertebrates, macrophytes and also water chemistry, water management, geographical description are presented. The review comprises the main studies beginning with the earliest faunistic publications up to the recent ecological, multidisciplinary investigations. Spatial and temporal patterns likewise water quality are considered as important. Additionally checklist of aquatic invertebrate and vertebrate fauna are given based on data from literature.

Short-term changes in phosphorus storage in an oligotrophic Everglades wetland ecosystem receiving experimental nutrient enrichment

Natural, unenriched Everglades wetlands are known to be limited by phosphorus (P) and responsive to P enrichment. However, whole-ecosystem evaluations of experimental P additions are rare in Everglades or other wetlands. We tested the response of the Everglades wetland ecosystem to continuous, low-level additions of P (0, 5, 15, and 30 μg L−1 above ambient) in replicate, 100 m flow-through flumes located in unenriched Everglades National Park. After the first six months of dosing, the concentration and standing stock of phosphorus increased in the surface water, periphyton, and flocculent detrital layer, but not in the soil or macrophytes. Of the ecosystem components measured, total P concentration increased the most in the floating periphyton mat (30 μg L−1: mean = 1916 μg P g−1, control: mean = 149 μg P g−1), while the flocculent detrital layer stored most of the accumulated P (30 μg L−1: mean = 1.732 g P m−2, control: mean = 0.769 g P m−2). Significant short-term responses of P concentration and standing stock were observed primarily in the high dose (30 μg L−1 above ambient) treatment. In addition, the biomass and estimated P standing stock of aquatic consumers increased in the 30 and 5 μg L−1 treatments. Alterations in P concentration and standing stock occurred only at the upstream ends of the flumes nearest to the point source of added nutrient. The total amount of P stored by the ecosystem within the flume increased with P dosing, although the ecosystem in the flumes retained only a small proportion of the P added over the first six months. These results indicate that oligotrophic Everglades wetlands respond rapidly to short-term, low-level P enrichment, and the initial response is most noticeable in the periphyton and flocculent detrital layer.

Phosphorus enrichment effects on interactions among the ecosystem components in a long-hydroperiod oligotrophic marsh in Everglades National Park

Hydrology and a history of oligotrophy unite the massive landscape comprising freshwater marsh in Everglades National Park. With restoration of water flow to the Everglades, phosphorus (P) enrichment, both from agricultural and domestic sources, may increase nutrient load to the marsh ecosystem. Previous research of P enrichment of Everglades soil, periphyton, and macrophytes revealed each of these ecosystem components responds to increased P loads with increased production and nutrient content. Interactions among these ecosystem components and how P affects the magnitude and direction of interaction are poorly understood and are the focus of my research. Here I present results of a two-year, two-factor experiment of P enrichment and manipulation in Everglades National Park. I quantified biomass, nutrient content, and production for periphyton and macrophyes and found macrophyte removal drives change in nutrient content, biomass, and production of periphyton. Periphyton removal did not appear to control macrophyte dynamics. Soil chemical and physical characteristics were explained primarily by site differences but there was an enrichment effect of soil porewater nitrite + nitrate, nitrite, and soluble reactive phosphorus. Flocculent materials production and depth were significantly affected by macrophyte removal where depth and production were significantly greater with the no macrophyte treatment. The dominant macrophyte of the marsh, Eleocharis cellulosa, increased more in the unenriched marsh than in the enriched marsh. The combination of these findings suggests that dynamics in floc and periphyton are controlled primarily by the presence of periphyton and that this relationship is significantly affected by low-level P enrichment. These results may be valuable in their application to both managers and policy makers who are involved in the Everglades restoration process. ^

Quantitative diatom-based reconstruction of paleoenvironmental conditions in Florida Bay and Biscayne Bay, U.S.A.

The spatial and temporal distribution of modern diatom assemblages in surface sediments, on the most dominant macrophytes, and in the water column at 96 locations in Florida Bay, Biscayne Bay and adjacent regions were examined in order to develop paleoenvironmental prediction models for this region. Analyses of these distributions revealed distinct temporal and spatial differences in assemblages among the locations. The differences among diatom assemblages living on subaquatic vegetation and sediments, and in the water column were significant. Because concentrations of salts, total phosphorus (WTP), total nitrogen (WTN) and total organic carbon (WTOC) are partly controlled by water management in this region, diatom-based models were produced to assess these variables. Discriminant function analyses showed that diatoms can also be successfully used to reconstruct changes in the abundance of diatom assemblages typical for different habitats and life habits. ^ To interpret paleoenvironmental changes, changes in salinity, WTN, WTP and WTOC were inferred from diatoms preserved in sediment cores collected along environmental gradients in Florida Bay (4 cores) and from nearshore and offshore locations in Biscayne Bay (3 cores). The reconstructions showed that water quality conditions in these estuaries have been fluctuating for thousands of years due to natural processes and sea-level changes, but almost synchronized shifts in diatom assemblages occurred in the mid-1960’s at all coring locations (except Ninemile Bank and Bob Allen Bank in Florida Bay). These alterations correspond to the major construction of numerous water management structures on the mainland. Additionally, all the coring sites (except Card Sound Bank, Biscayne Bay and Trout Cove, Florida Bay) showed decreasing salinity and fluctuations in nutrient levels in the last two decades that correspond to increased rainfall in the 1990’s and increased freshwater discharge to the bays, a result of increased freshwater deliveries to the Everglades by South Florida Water Management District in the 1980’s and 1990’s. Reconstructions of the abundance of diatom assemblages typical for different habitats and life habits revealed multiple sources of diatoms to the coring locations and that epiphytic assemblages in both bays increased in abundance since the early 1990’s. ^

Seed dispersal and reproduction patterns among Everglades plants

In this study three aspects of sexual reproduction in Everglades plants were examined to more clearly understand seed dispersal and the allocation of resources to sexual reproduction—spatial dispersal process, temporal dispersal of seeds (seedbank), and germination patterns in the dominant species, sawgrass (Cladium jamaicense). Community assembly rules for fruit dispersal were deduced by analysis of functional traits associated with this process. Seedbank ecology was investigated by monitoring emergence of germinants from sawgrass soil samples held under varying water depths to determine the fate of dispersed seeds. Fine-scale study of sawgrass fruits yielded information on contributions to variation in sexually produced propagules in this species, which primarily reproduces vegetatively. It was hypothesized that Everglades plants possess a set of functional traits that enhance diaspore dispersal. To test this, 14 traits were evaluated among 51 species by factor analysis. The factorial plot of this analysis generated groups of related traits, with four suites of traits forming dispersal syndromes. Hydrochory traits were categorized by buoyancy and appendages enhancing buoyancy. Anemochory traits were categorized by diaspore size and appendages enhancing air movement. Epizoochory traits were categorized by diaspore size, buoyancy, and appendages allowing for attachment. Endozoochory traits were categorized by diaspore size, buoyancy, and appendages aiding diaspore presentation. These patterns/trends of functional trait organization also represent dispersal community assembly rules. Seeds dispersed by hydrochory were hypothesized to be caught most often in the edge of the north side of sawgrass patches. Patterns of germination and dispersal mode of all hydrochorous macrophytes with propagules in the seedbank were elucidated by germination analysis from 90 soil samples collected from 10 sawgrass patches. Mean site seed density was 486 seeds/m2 from 13 species. Most seeds collected at the north side of patches and significantly in the outer one meter of the patch edge (p = 0.013). Sawgrass seed germination was hypothesized to vary by site, among individual plants, and within different locations of a plant’s infructescence. An analysis of sawgrass fruits with nested ANOVAs found that collection site and interaction of site x individual plant significantly affect germination ability, seed viability, and fruit size (p ≤ 0.050). Fruit location within a plant’s infructescence did not significantly affect germination. As for allocation of resources to sexual reproduction, only 17.9% of sawgrass seeds germinated and only 4.8% of ungerminated seeds with fleshy endosperm were presumed viable, but dormant. Collectively, only 22% of all sawgrass seeds produced were viable.

Novel methodology for in situ carbon dioxide enrichment of benthic ecosystems

Future climate change will likely represent a major stress to shallow aquatic and coastal marine communities around the world. Most climate change research, particularly in regards to increased pCO2 and ocean acidification, relies on ex situ mesocosm experimentation, isolating target organisms from their environment. Such mesocosms allow for greater experimental control of some variables, but can often cause unrealistic changes in a variety of environmental factors, leading to “bottle effects.” Here we present an in situ technique of altering dissolved pCO2within nearshore benthic communities (e.g., macrophytes, algae, and/or corals) using submerged clear, open-top chambers. Our technique utilizes a flow-through design that replicates natural water flow conditions and minimizes caging effects. The clear, open-top design additionally ensures that adequate light reaches the benthic community. Our results show that CO2 concentrations and pH can be successfully manipulated for long durations within the open-top chambers, continuously replicating forecasts for the year 2100. Enriched chambers displayed an average 0.46 unit reduction in pH as compared with ambient chambers over a 6-month period. Additionally, CO2 and HCO3 – concentrations were all significantly higher within the enriched chambers. We discuss the advantages and disadvantages of this technique in comparison to other ex situ mesocosm designs used for climate change research.

Decadal Change in Vegetation and Soil Phosphorus Pattern across the Everglades Landscape

Wetlands respond to nutrient enrichment with characteristic increases in soil nutrients and shifts in plant community composition. These responses to eutrophication tend to be more rapid and longer lasting in oligotrophic systems. In this study, we documented changes associated with water quality from 1989 to 1999 in oligotrophic Everglades wetlands. We accomplished this by resampling soils and macrophytes along four transects in 1999 that were originally sampled in 1989. In addition to documenting soil phosphorus (P) levels and decadal changes in plant species composition at the same sites, we report macrophyte tissue nutrient and biomass data from 1999 for future temporal comparisons. Water quality improved throughout much of the Everglades in the 1990s. In spite of this improvement, though, we found that water quality impacts worsened during this time in areas of the northern Everglades (western Loxahatchee National Wildlife Refuge [NWR] and Water Conservation Area [WCA] 2A). Zones of high soil P (exceeding 700 mg P kg−1 dry wt. soil) increased to more than 1 km from the western margin canal into the Loxahatchee NWR and more than 4 km from northern boundary canal into WCA-2A. This doubling of the high soil P zones since 1989 was paralleled with an expansion of cattail (Typha spp.)-dominated marsh in both regions. Macrophyte species richness declined in both areas from 1989 to 1999 (27% in the Loxahatchee NWR and 33% in WCA-2A). In contrast, areas well south of the Everglades Agricultural Area, including WCA-3A and Everglades National Park (ENP), did not decline during this time. We found no significant decadal change in plant community patterns from 1989 and 1999 along transects in southern WCA-3A or Shark River Slough (ENP). Our 1999 sampling also included a new transect in Taylor Slough (ENP), which will allow change analysis here in the future. Regular sampling of these transects, to verify decadal-scale environmental impacts or improvements, will continue to be an important tool for long-term management and restoration of the Everglades.

Ecological effects of low-level phosphorus additions on two plant communities in a neotropical freshwater wetland ecosystem

We conducted a low-level phosphorus (P) enrichment study in two oligotrophic freshwater wetland communities (wet prairies [WP] and sawgrass marsh [SAW]) of the neotropical Florida Everglades. The experiment included three P addition levels (0, 3.33, and 33.3 mg P m−2 month−1), added over 2 years, and used in situ mesocosms located in northeastern Everglades National Park, Fla., USA. The calcareous periphyton mat in both communities degraded quickly and was replaced by green algae. In the WP community, we observed significant increases in net aboveground primary production (NAPP) and belowground biomass. Aboveground live standing crop (ALSC) did not show a treatment effect, though, because stem turnover rates of Eleocharis spp., the dominant emergent macrophyte in this community, increased significantly. Eleocharis spp. leaf tissue P content decreased with P additions, causing higher C:P and N:P ratios in enriched versus unenriched plots. In the SAW community, NAPP, ALSC, and belowground biomass all increased significantly in response to P additions. Cladium jamaicense leaf turnover rates and tissue nutrient content did not show treatment effects. The two oligotrophic communities responded differentially to P enrichment. Periphyton which was more abundant in the WP community, appeared to act as a P buffer that delayed the response of other ecosystem components until after the periphyton mat had disappeared. Periphyton played a smaller role in controlling ecosystem dynamics and community structure in the SAW community. Our data suggested a reduced reliance on internal stores of P by emergent macrophytes in the WP that were exposed to P enrichment. Eleocharis spp. rapidly recycled P through more rapid aboveground turnover. In contrast, C. jamaicense stored added P by initially investing in belowground biomass, then shifting growth allocation to aboveground tissue without increasing leaf turnover rates. Our results suggest that calcareous wetland systems throughout the Caribbean, and oligotrophic ecosystems in general, respond rapidly to low-level additions of their limiting nutrient.

Seasonal plant water uptake patterns in the saline southeast Everglades ecotone

The purpose of this study was to determine the seasonal water use patterns of dominant macrophytes coexisting in the coastal Everglades ecotone. We measured the stable isotope signatures in plant xylem water of Rhizophora mangle, Cladium jamaicense, and Sesuvium portulacastrum during the dry (DS) and wet (WS) seasons in the estuarine ecotone along Taylor River in Everglades National Park, FL, USA. Shallow soilwater and deeper groundwater salinity was also measured to extrapolate the salinity encountered by plants at their rooting zone. Average soil water oxygen isotope ratios (δ 18O) was enriched (4.8 ± 0.2‰) in the DS relative to the WS (0.0 ± 0.1‰), but groundwater δ 18O remained constant between seasons (DS: 2.2 ± 0.4‰; WS: 2.1 ± 0.1‰). There was an inversion in interstitial salinity patterns across the soil profile between seasons. In the DS, shallow water was euhaline [i.e., 43 practical salinity units (PSU)] while groundwater was less saline (18 PSU). In the WS, however, shallow water was fresh (i.e., 0 PSU) but groundwater remained brackish (14 PSU). All plants utilized 100% (shallow) freshwater during the WS, but in the DS R. mangle switched to a soil–groundwater mix (δ 55% groundwater) while C. jamaicense and S. portulacastrum continued to use euhaline shallow water. In the DS, based on δ 18O data, the roots of R. mangle roots were exposed to salinities of 25.4 ± 1.4 PSU, less saline than either C. jamaicense(39.1 ± 2.2 PSU) or S. portulacastrum (38.6 ± 2.5 PSU). Although the salinity tolerance of C. jamaicense is not known, it is unlikely that long-term exposure to high salinity is conducive to the persistence of this freshwater marsh sedge. This study increases our ecological understanding of how water uptake patterns of individual plants can contribute to ecosystem levels changes, not only in the southeast saline Everglades, but also in estuaries in general in response to global sea level rise and human-induced changes in freshwater flows.

Characteristics of surface-water flows in the ridge and slough landscape of Everglades National Park: implications for particulate transport

Over the last one hundred years, compartmentalization and water management activities have reduced water flow to the ridge and slough landscape of the Everglades. As a result, the once corrugated landscape has become topographically and vegetationally uniform. The focus of this study was to quantify variation in surface flow in the ridge and slough landscape and to relate flow conditions to particulate transport and deposition. Over the 2002–2003 and 2003–2004 wet seasons, surface velocities and particulate accumulation were measured in upper Shark River Slough in Everglades National Park. Landscape characteristics such as elevation, plant density and biomass also were examined to determine their impact on flow characteristics and material transport. The results of this study demonstrate that the release of water during the wet season not only increases water levels, but also increased flow speeds and particulate transport and availability. Further, flow speeds were positively and significantly correlated with water level thereby enhancing particulate transport in sloughs relative to ridges especially during peak flow periods. Our results also indicate that the distribution of biomass in the water column, including floating plants and periphyton, affects velocity magnitude and shape of vertical profiles, especially in the sloughs where Utricularia spp. and periphyton mats are more abundant. Plot clearing experiments suggest that the presence of surface periphyton and Utricularia exert greater control over flow characteristics than the identity (i.e., sawgrass or spike rush) or density of emergent macrophytes, two parameters frequently incorporated into models describing flow through vegetated canopies. Based on these results, we suggest that future modeling efforts must take the presence of floating biomass, such as Utricularia, and presence of periphyton into consideration when describing particulate transport.

Spatiotemporal patterns in community structure of macroinvertebrates inhabiting calcareous periphyton mats

Calcareous floating periphyton mats in the southern Everglades provide habitat for a diverse macroinvertebrate community that has not been well characterized. Our study described this community in an oligotrophic marsh, compared it with the macroinvertebrate community associated with adjacent epiphytic algae attached to macrophytes in the water column, and detected spatial patterns in density and community structure. The floating periphyton mat (floating mat) and epiphytic algae in the water column (submerged epiphyton) were sampled at 4 sites (1 km apart) in northern Shark River Slough, Everglades National Park (ENP), in the early (July) and late (November) wet season. Two perpendicular 90-m transects were established at each site and 100 samples were taken in a nested design. Sites were located in wet-prairie spikerush-dominated sloughs with similar water depths and emergent macrophyte communities. Floating mats were sampled by taking cores (6-cm diameter) that were sorted under magnification to enumerate infauna retained on a 250-μm-mesh sieve and with a maximum dimension >1 mm. Our results showed that floating mats provide habitat for a macroinvertebrate community with higher densities (no. animals/g ash-free dry mass) of Hyalella azteca, Dasyhelea spp., and Cladocera, and lower densities of Chironomidae and Planorbella spp. than communities associated with submerged epiphyton. Densities of the most common taxa increased 3× to 15× from early to late wet season, and community differences between the 2 habitat types became more pronounced. Floating-mat coverage and estimated floating-mat biomass increased 20 to 30% and 30 to 110%, respectively, at most sites in the late wet season. Some intersite variation was observed in individual taxa, but no consistent spatial pattern in any taxon was detected at any scale (from 0.2 m to 3 km). Floating mats and their resident macroinvertebrate communities are important components in the Everglades food web. This community should be included in environmental monitoring programs because degradation and eventual loss of the calcareous periphyton mat is associated with P enrichment in this ecosystem.

Sediment and soil organic matter source assessment as revealed by the molecular distribution and carbon isotopic composition of n-alkanes

The assessment of organic matter (OM) sources in sediments and soils is a key to better understand the biogeochemical cycling of carbon in aquatic environments. While traditional molecular marker-based methods have provided such information for typical two end member (allochthonous/terrestrial vs. autochthonous/microbial)-dominated systems, more detailed, biomass-specific assessments are needed for ecosystems with complex OM inputs such as tropical and sub-tropical wetlands and estuaries where aquatic macrophytes and macroalgae may play an important role as OM sources. The aim of this study was to assess the utility of a combined approach using compound specific stable carbon isotope analysis and an n-alkane based proxy (Paq) to differentiate submerged and emergent/terrestrial vegetation OM inputs to soils/sediments from a sub-tropical wetland and estuarine system, the Florida Coastal Everglades. Results show that Paq values (0.13–0.51) for the emergent/terrestrial plants were generally lower than those for freshwater/marine submerged vegetation (0.45–1.00) and that compound specific δ13C values for the n-alkanes (C23 to C31) were distinctively different for terrestrial/emergent and freshwater/marine submerged plants. While crossplots of the Paq and n-alkane stable isotope values for the C23n-alkane suggest that OM inputs are controlled by vegetation changes along the freshwater to marine transect, further resolution regarding OM input changes along this landscape was obtained through principal component analysis (PCA), successfully grouping the study sites according to the OM source strengths. The data show the potential for this n-alkane based multi-proxy approach as a means of assessing OM inputs to complex ecosystems.

Phosphorus cycling and partitioning in an oligotrophic Everglades wetland ecosystem: a radioisotope tracing study

1. Our goal was to quantify short-term phosphorus (P) partitioning and identify the ecosystem components important to P cycling in wetland ecosystems. To do this, we added P radiotracer to oligotrophic, P-limited Everglades marshes. 32PO4 was added to the water column in six 1-m2 enclosed mesocosms located in long-hydroperiod marshes of Shark River Slough, Everglades National Park. Ecosystem components were then repeatedly sampled over 18 days. 2. Water column particulates (>0.45 μm) incorporated radiotracer within the first minute after dosing and stored 95–99% of total water column 32P activity throughout the study. Soluble (<0.45 μm) 32P in the water column, in contrast, was always <5% of the 32P in surface water. Periphyton, both floating and attached to emergent macrophytes, had the highest specific activity of 32P (Bq g−131P) among the different ecosystem components. Fish and aquatic macroinvertebrates also had high affinity for P, whereas emergent macrophytes, soil and flocculent detrital organic matter (floc) had the lowest specific activities of radiotracer. 3. Within the calcareous, floating periphyton mats, 81% of the initial 32P uptake was associated with Ca, but most of this 32P entered and remained within the organic pool (Ca-associated = 14% of total) after 1 day. In the floc layer, 32P rapidly entered the microbial pool and the labile fraction was negligible for most of the study. 4. Budgeting of the radiotracer indicated that 32P moved from particulates in the water column to periphyton and floc and then to the floc and soil over the course of the 18 day incubations. Floc (35% of total) and soil (27%) dominated 32P storage after 18 days, with floating periphyton (12%) and surface water (10%) holding smaller proportions of total ecosystem 32P. 5. To summarise, oligotrophic Everglades marshes exhibited rapid uptake and retention of labile 32P. Components dominated by microbes appear to control short-term P cycling in this oligotrophic ecosystem.

Short-term changes in phosphorus storage in an oligotrophic Everglades wetland ecosystem receiving experimental nutrient enrichment

Natural, unenriched Evergladeswetlands are known to be limited by phosphorus(P) and responsive to P enrichment. However,whole-ecosystem evaluations of experimental Padditions are rare in Everglades or otherwetlands. We tested the response of theEverglades wetland ecosystem to continuous,low-level additions of P (0, 5, 15, and30 μg L−1 above ambient) in replicate,100 m flow-through flumes located in unenrichedEverglades National Park. After the first sixmonths of dosing, the concentration andstanding stock of phosphorus increased in thesurface water, periphyton, and flocculentdetrital layer, but not in the soil or macrophytes. Of the ecosystem components measured, total P concentration increased the most in the floating periphyton mat (30 μg L−1: mean = 1916 μg P g−1, control: mean =149 μg P g−1), while the flocculentdetrital layer stored most of the accumulated P(30 μg L−1: mean = 1.732 g P m−2,control: mean = 0.769 g P m−2). Significant short-term responsesof P concentration and standing stock wereobserved primarily in the high dose (30 μgL−1 above ambient) treatment. Inaddition, the biomass and estimated P standingstock of aquatic consumers increased in the 30and 5 μg L−1 treatments. Alterationsin P concentration and standing stock occurredonly at the upstream ends of the flumes nearestto the point source of added nutrient. Thetotal amount of P stored by the ecosystemwithin the flume increased with P dosing,although the ecosystem in the flumes retainedonly a small proportion of the P added over thefirst six months. These results indicate thatoligotrophic Everglades wetlands respondrapidly to short-term, low-level P enrichment,and the initial response is most noticeable inthe periphyton and flocculent detrital layer.

Phosphorus budgets in Everglades wetland ecosystems: the effects of hydrology and nutrient enrichment

The Florida Everglades is a naturally oligotrophic hydroscape that has experienced large changes in ecosystem structure and function as the result of increased anthropogenic phosphorus (P) loading and hydrologic changes. We present whole-ecosystem models of P cycling for Everglades wetlands with differing hydrology and P enrichment with the goal of synthesizing existing information into ecosystem P budgets. Budgets were developed for deeper water oligotrophic wet prairie/slough (‘Slough’), shallower water oligotrophic Cladium jamaicense (‘Cladium’), partially enriched C. jamaicense/Typha spp. mixture (‘Cladium/Typha’), and enriched Typha spp. (‘Typha’) marshes. The majority of ecosystem P was stored in the soil in all four ecosystem types, with the flocculent detrital organic matter (floc) layer at the bottom of the water column storing the next largest proportion of ecosystem P pools. However, most P cycling involved ecosystem components in the water column (periphyton, floc, and consumers) in deeper water, oligotrophic Slough marsh. Fluxes of P associated with macrophytes were more important in the shallower water, oligotrophic Cladium marsh. The two oligotrophic ecosystem types had similar total ecosystem P stocks and cycling rates, and low rates of P cycling associated with soils. Phosphorus flux rates cannot be estimated for ecosystem components residing in the water column in Cladium/Typha or Typha marshes due to insufficient data. Enrichment caused a large increase in the importance of macrophytes to P cycling in Everglades wetlands. The flux of P from soil to the water column, via roots to live aboveground tissues to macrophyte detritus, increased from 0.03 and 0.2 g P m−2 yr−1 in oligotrophic Slough and Cladium marsh, respectively, to 1.1 g P m−2 yr−1 in partially enriched Cladium/Typha, and 1.6 g P m−2 yr−1 in enriched Typha marsh. This macrophyte translocation P flux represents a large source of internal eutrophication to surface waters in P-enriched areas of the Everglades.