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.

Spatial and temporal patterns of aboveground net primary productivity (ANPP) along two freshwater-estuarine transects in the Florida Coastal Everglades

We present here a 4-year dataset (2001–2004) on the spatial and temporal patterns of aboveground net primary production (ANPP) by dominant primary producers (sawgrass, periphyton, mangroves, and seagrasses) along two transects in the oligotrophic Florida Everglades coastal landscape. The 17 sites of the Florida Coastal Everglades Long Term Ecological Research (FCE LTER) program are located along fresh-estuarine gradients in Shark River Slough (SRS) and Taylor River/C-111/Florida Bay (TS/Ph) basins that drain the western and southern Everglades, respectively. Within the SRS basin, sawgrass and periphyton ANPP did not differ significantly among sites but mangrove ANPP was highest at the site nearest the Gulf of Mexico. In the southern Everglades transect, there was a productivity peak in sawgrass and periphyton at the upper estuarine ecotone within Taylor River but no trends were observed in the C-111 Basin for either primary producer. Over the 4 years, average sawgrass ANPP in both basins ranged from 255 to 606 g m−2 year−1. Average periphyton productivity at SRS and TS/Ph was 17–68 g C m−2 year−1 and 342–10371 g C m−2 year−1, respectively. Mangrove productivity ranged from 340 g m−2 year−1 at Taylor River to 2208 g m−2 year−1 at the lower estuarine Shark River site. Average Thalassia testudinum productivity ranged from 91 to 396 g m−2 year−1 and was 4-fold greater at the site nearest the Gulf of Mexico than in eastern Florida Bay. There were no differences in periphyton productivity at Florida Bay. Interannual comparisons revealed no significant differences within each primary producer at either SRS or TS/Ph with the exception of sawgrass at SRS and the C−111 Basin. Future research will address difficulties in assessing and comparing ANPP of different primary producers along gradients as well as the significance of belowground production to the total productivity of this ecosystem.

Projected Reorganization of Florida Bay Seagrass Communities in Response to the Increased Freshwater Inflow of Everglades Restoration

Historic changes in water-use management in the Florida Everglades have caused the quantity of freshwater inflow to Florida Bay to decline by approximately 60% while altering its timing and spatial distribution. Two consequences have been (1) increased salinity throughout the bay, including occurrences of hypersalinity, coupled with a decrease in salinity variability, and (2) change in benthic habitat structure. Restoration goals have been proposed to return the salinity climates (salinity and its variability) of Florida Bay to more estuarine conditions through changes in upstream water management, thereby returning seagrass species cover to a more historic state. To assess the potential for meeting those goals, we used two modeling approaches and long-term monitoring data. First, we applied the hydrological mass balance model FATHOM to predict salinity climate changes in sub-basins throughout the bay in response to a broad range of freshwater inflow from the Everglades. Second, because seagrass species exhibit different sensitivities to salinity climates, we used the FATHOM-modeled salinity climates as input to a statistical discriminant function model that associates eight seagrass community types with water quality variables including salinity, salinity variability, total organic carbon, total phosphorus, nitrate, and ammonium, as well as sediment depth and light reaching the benthos. Salinity climates in the western sub-basins bordering the Gulf of Mexico were insensitive to even the largest (5-fold) modeled increases in freshwater inflow. However, the north, northeastern, and eastern sub-basins were highly sensitive to freshwater inflow and responded to comparatively small increases with decreased salinity and increased salinity variability. The discriminant function model predicted increased occurrences ofHalodule wrightii communities and decreased occurrences of Thalassia testudinum communities in response to the more estuarine salinity climates. The shift in community composition represents a return to the historically observed state and suggests that restoration goals for Florida Bay can be achieved through restoration of freshwater inflow from the Everglades.

Occurrence and distribution of novel botryococcene hydrocarbons in freshwater wetlands of the Florida Everglades

A high abundance of isoprenoid hydrocarbons, the botryococcenes, with carbon numbers from 32 to 34 were detected in the Florida Everglades freshwater wetlands. These compounds were present in varying amounts up to 106 μg/gdw in periphyton, 278 μg/gdw in floc, and 46 μg/gdw in soils. Their structures were determined based on comparison to standards, interpretation of their mass spectra and those of their hydrogenation products, and comparison of Kovats indexes to those reported in the literature. A total of 26 cyclic and acyclic botryococcenes with 8 skeletons were identified, including those with fewer degrees of unsaturation, which are proposed as early diagenetic derivatives from the natural products. This is the first report that botryococcenes occur in the Everglades freshwater wetlands. Their potential biogenetic sources from green algae and cyanobacteria were examined, but neither contained botryococcenes. Thus, the source implication of botryococcenes in this ecosystem needs further study.

Modeling seasonal dynamics of small fish cohorts in fluctuating freshwater marsh landscapes

Small-bodied fishes constitute an important assemblage in many wetlands. In wetlands that dry periodically except for small permanent waterbodies, these fishes are quick to respond to change and can undergo large fluctuations in numbers and biomasses. An important aspect of landscapes that are mixtures of marsh and permanent waterbodies is that high rates of biomass production occur in the marshes during flooding phases, while the permanent waterbodies serve as refuges for many biotic components during the dry phases. The temporal and spatial dynamics of the small fishes are ecologically important, as these fishes provide a crucial food base for higher trophic levels, such as wading birds. We develop a simple model that is analytically tractable, describing the main processes of the spatio-temporal dynamics of a population of small-bodied fish in a seasonal wetland environment, consisting of marsh and permanent waterbodies. The population expands into newly flooded areas during the wet season and contracts during declining water levels in the dry season. If the marsh dries completely during these times (a drydown), the fish need refuge in permanent waterbodies. At least three new and general conclusions arise from the model: (1) there is an optimal rate at which fish should expand into a newly flooding area to maximize population production; (2) there is also a fluctuation amplitude of water level that maximizes fish production, and (3) there is an upper limit on the number of fish that can reach a permanent waterbody during a drydown, no matter how large the marsh surface area is that drains into the waterbody. Because water levels can be manipulated in many wetlands, it is useful to have an understanding of the role of these fluctuations.

Molecular characterization of dissolved organic matter in freshwater wetlands of the Florida Everglades

In this study, the molecular composition of dissolved organic matter (DOM), collected from wetlands of the Southern Everglades, was examined using a variety of analytical techniques in order to characterize its sources and transformation in the environment. The methods applied for the characterization of DOM included fluorescence spectroscopy, solid state 13C CPMAS NMR spectroscopy, and pyrolysis-GC/MS. The relative abundance of protein-like components and carbohydrates increased from the canal site to more remote freshwater marsh sites suggesting that significant amounts of non-humic DOM are autochthonously produced within the freshwater marshes, and are not exclusively introduced through canal inputs. Such in situ DOM production is important when considering how DOM from canals is processed and transported to downstream estuaries of Florida Bay.

Comparison of snail density, standing stock, and body size between Caribbean karst wetlands and other freshwater ecosystems

Synthesizing data from multiple studies generates hypotheses about factors that affect the distribution and abundance of species among ecosystems. Snails are dominant herbivores in many freshwater ecosystems, but there is no comprehensive review of snail density, standing stock, or body size among freshwater ecosystems. We compile data on snail density and standing stock, estimate body size with their quotient, and discuss the major pattern that emerges. We report data from 215 freshwater ecosystems taken from 88 studies that we placed into nine categories. Sixty-five studies reported density, seven reported standing stock, and 16 reported both. Despite the breadth of studies, spatial and temporal sampling scales were limited. Researchers used 25 different sampling devices ranging in area from 0.0015 to 2.5 m2. Most ecosystem categories had similar snail densities, standing stocks, and body sizes suggesting snails shared a similar function among ecosystems. Caribbean karst wetlands were a striking exception with much lower density and standing stock, but large body size. Disparity in body size results from the presence of ampullariids in Caribbean karst wetlands suggesting that biogeography affects the distribution of taxa, and in this case size, among aquatic ecosystems. We propose that resource quality explains the disparity in density and standing stock between Caribbean karst wetlands and other categories. Periphyton in Caribbean karst wetlands has high carbon-to-phosphorous ratios and defensive characteristics that inhibit grazers. Unlike many freshwater ecosystems where snails are key grazers, we hypothesize that a microbial loop captures much of the primary production in Caribbean karst wetlands.

A Hydrological Budget (2002–2008) for a Large Subtropical Wetland Ecosystem Indicates Marine Groundwater Discharge Accompanies Diminished Freshwater Flow

Water budget parameters are estimated for Shark River Slough (SRS), the main drainage within Everglades National Park (ENP) from 2002 to 2008. Inputs to the water budget include surface water inflows and precipitation while outputs consist of evapotranspiration, discharge to the Gulf of Mexico and seepage losses due to municipal wellfield extraction. The daily change in volume of SRS is equated to the difference between input and outputs yielding a residual term consisting of component errors and net groundwater exchange. Results predict significant net groundwater discharge to the SRS peaking in June and positively correlated with surface water salinity at the mangrove ecotone, lagging by 1 month. Precipitation, the largest input to the SRS, is offset by ET (the largest output); thereby highlighting the importance of increasing fresh water inflows into ENP for maintaining conditions in terrestrial, estuarine, and marine ecosystems of South Florida.

Determining Spatial and Temporal Inputs of Freshwater, Including Submarine Groundwater Discharge, to a Subtropical Estuary Using Geochemical Tracers, Biscayne Bay, South Florida

Geochemical mixing models were used to decipher the dominant source of freshwater (rainfall, canal discharge, or groundwater discharge) to Biscayne Bay, an estuary in south Florida. Discrete samples of precipitation, canal water, groundwater, and bay surface water were collected monthly for 2 years and analyzed for salinity, stable isotopes of oxygen and hydrogen, and Sr2+/Ca2+ concentrations. These geochemical tracers were used in three separate mixing models and then combined to trace the magnitude and timing of the freshwater inputs to the estuary. Fresh groundwater had an isotopic signature (δ 18O = −2.66‰, δD −7.60‰) similar to rainfall (δ 18O = −2.86‰, δD = −4.78‰). Canal water had a heavy isotopic signature (δ 18O = −0.46‰, δD  = −2.48‰) due to evaporation. This made it possible to use stable isotopes of oxygen and hydrogen to separate canal water from precipitation and groundwater as a source of freshwater into the bay. A second model using Sr2+/Ca2+ ratios was developed to discern fresh groundwater inputs from precipitation inputs. Groundwater had a Sr2+/Ca2+ ratio of 0.07, while precipitation had a dissimilar ratio of 0.89. When combined, these models showed a freshwater input ratio of canal/precipitation/groundwater of 37%:53%:10% in the wet season and 40%:55%:5% in the dry season with an error of ±25%. For a bay-wide water budget that includes saltwater and freshwater mixing, fresh groundwater accounts for 1–2% of the total fresh and saline water input.

Controls on Ecosystem Carbon Dioxide Exchange in Short- and Long-Hydroperiod Florida Everglades Freshwater Marshes

Although freshwater wetlands are among the most productive ecosystems on Earth, little is known of carbon dioxide (CO2) exchange in low latitude wetlands. The Everglades is an extensive, oligotrophic wetland in south Florida characterized by short- and long-hydroperiod marshes. Chamber-based CO2 exchange measurements were made to compare the marshes and examine the roles of primary producers, seasonality, and environmental drivers in determining exchange rates. Low rates of CO2 exchange were observed in both marshes with net ecosystem production reaching maxima of 3.77 and 4.28 μmol CO2 m−2 s−1 in short- and long-hydroperiod marshes, respectively. Fluxes of CO2 were affected by seasonality only in the short-hydroperiod marsh, where flux rates were significantly lower in the wet season than in the dry season. Emergent macrophytes dominated fluxes at both sites, though this was not the case for the short-hydroperiod marsh in the wet season. Water depth, a factor partly under human control, significantly affected gross ecosystem production at the short-hydroperiod marsh. As Everglades ecosystem restoration proceeds, leading to deeper water and longer hydroperiods, productivity in short-hydroperiod marshes will likely be more negatively affected than in long-hydroperiod marshes. The Everglades stand in contrast to many freshwater wetlands because of ecosystem-wide low productivity rates.

Relating Freshwater Flow with Estuarine Water Quality in the Southern Everglades Mangrove Ecotone

Florida Bay is more saline than it was historically, and reduced freshwater flows may lead to more phosphorus inputs to the mangrove ecotone from the marine end-member. This is important given plans to restore freshwater flow into eastern Florida Bay. We investigated the relationships between salinity, nutrients, and hydrologic variables in the mangrove ecotone of Taylor Slough. We expected that total phosphorus (TP) would increase with salinity, reflecting a downstream marine source, while total nitrogen (TN) would increase with flow in the mangrove ecotone. Despite expectations of increased flows improving the ecological health of lower Taylor Slough and Florida Bay, total nitrogen (TN) and total phosphorus (TP) dynamics may shift in response to new conditions of flow and salinity as well as organic carbon, N, and P availability. Our results showed that TP concentrations are more discharge-driven while TN is more variable and potentially derived from different sources along the flow path from the freshwater Everglades marshes to Florida Bay. Increased flow of freshwater through Taylor Slough will likely decrease TP concentrations in this historically oligotrophic and P-limited ecosystem. However, more studies along the mangrove ecotone is needed to understand how increased flows will affect nitrogen dynamics relative to phosphorus.

Dissolved Black Nitrogen (DBN) in Freshwater Environments

Biomass burning results in the formation and accumulation of pyrogenic products such as black carbon (BC) and black nitrogen (BN) in soils. The ubiquitous presence of pyrogenic products in natural dissolved organic matter (DOM) and potential implications in global carbon cycling have recently been reported. However, little is known about the environmental dynamics or the importance in the global N cycle of dissolved BN (DBN; or heteroaromatic N). Here we report the coupling between DBN and dissolved BC (DBC) in ultrafiltered DOM from six headwater streams across a climatic region of North America, suggesting similar combustion sources, and that DOC may play an important role in the translocation of soil BN to the dissolved phase. The export of potentially recalcitrant riverine DBN to the ocean may affect the biogeochemical cycling of N and possibly the microbial community structure in aquatic environments.

Woody Plant Invasion into the Freshwater Marl Prairie Habitat of the Cape Sable Seaside Sparrow: Final Report

In the fall of 2005, U.S. Fish and Wildlife Services (USFWS) contracted with Florida International University (FIU) to study the physical and biological drivers underlying the distribution of woody plant species in the marl prairie habitat of the Cape Sable Seaside Sparrow (CSSS). This report presents what we have learned about woody plant encroachment based on studies carried out during the period 2006-2008. The freshwater marl prairie habitat currently occupied by the Cape Sable seaside sparrow (CSSS; Ammodramus maritimus mirabilis) is a dynamic mosaic comprised of species-rich grassland communities and tree islands of various sizes, densities and compositions. Landscape heterogeneity and the scale of vegetative components across the marl prairie is primarily determined by hydrologic conditions, biological factors (e.g. dispersal and growth morphology), and disturbances such as fire. The woody component of the marl prairie landscape is subject to expansion through multiple positive feedback mechanisms, which may be initiated by recent land use change (e.g. drainage). Because sparrows are known to avoid areas where the woody component is too extensive, a better understanding of invasion dynamics is needed to ensure proper management.

Contribution of lipophilic secondary metabolites to the toxicity of strains of freshwater cyanobacterial harmful algal blooms, identified using the zebrafish (Danio rerio) embryo as a model for vertebrate development

Cyanobacteria ("blue-green algae") are known to produce a diverse repertoire of biologically active secondary metabolites. When associated with so-called "harmful algal blooms", particularly in freshwater systems, a number of these metabolites have been associated—as "toxins", or commonly "cyanotoxins"—with human and animal health concerns. In addition to the known water-soluble toxins from these genera (i.e. microcystins, cylindrospermopsin, and saxitoxins), our studies have shown that there are metabolites within the lipophilic extracts of these strains that inhibit vertebrate development in zebrafish embryos. Following these studies, the zebrafish embryo model was implemented in the bioassay-guided purification of four isolates of cyanobacterial harmful algal blooms, namely Aphanizomenon, two isolates of Cylindrospermopsis, and Microcystis, in order to identify and chemically characterize the bioactive lipophilic metabolites in these isolates. ^ We have recently isolated a group of polymethoxy-1-alkenes (PMAs), as potential toxins, based on the bioactivity observed in the zebrafish embryos. Although PMAs have been previously isolated from diverse cyanobacteria, they have not previously been associated with relevant toxicity. These compounds seem to be widespread across the different genera of cyanobacteria, and, according to our studies, suggested to be derived from the polyketide biosynthetic pathway which is a common synthetic route for cyanobacterial and other algal toxins. Thus, it can be argued that these metabolites are perhaps important contributors to the toxicity of cyanobacterial blooms. In addition to the PMAs, a set of bioactive glycosidic carotenoids were also isolated because of their inhibition of zebrafish embryonic development. These pigmented organic molecules are found in many photosynthetic organisms, including cyanobacteria, and they have been largely associated with the prevention of photooxidative damage. This is the first indication of these compounds as toxic metabolites and the hypothesized mode of action is via their biotransformation to retinoids, some of which are known to be teratogenic. Additional fractions within all four isolates have been shown to contain other uncharacterized lipophilic toxic metabolites. This apparent repertoire of lipophilic compounds may contribute to the toxicity of these cyanobacterial harmful algal blooms, which were previously attributed primarily to the presence of the known water-soluble toxins.^