Quantifying nitrogen cycling rates in freshwater marshes of the southern Everglades using nitrogen-15 tracer techniques

This dissertation research project addressed the question of how hydrologic restoration of the Everglades is impacting the nutrient dynamics of marsh ecosystems in the southern Everglades. These effects were analyzed by quantifying nitrogen (N) cycle dynamics in the region. I utilized stable isotope tracer techniques to investigate nitrogen uptake and cycling between the major ecosystem components of the freshwater marsh system. I recorded the natural isotopic signatures (δ15N and δ 13C) for major ecosystem components from the three major watersheds of the Everglades: Shark River Slough, Taylor Slough, and C-111 basin. Analysis of δ15 N and δ13C natural abundance data were used to demonstrate the spatial extent to which nitrogen from anthropogenic or naturally enriched sources is entering the marshes of the Everglades. In addition, I measured the fluxes on N between various ecosystem components at both near-canal and estuarine ecotone locations. Lastly, I investigated the effect of three phosphorus load treatments (0.00 mg P m-2, 6.66 mg P m-2, and 66.6 mg P m-2) on the rate and magnitude of ecosystem N-uptake and N-cycling. The δ15N and δ13C natural abundance data supported the hypothesis that ecosystem components from near-canal sites have heavier, more enriched δ 15N isotopic signatures than downstream sites. The natural abundance data also showed that the marshes of the southern Everglades are acting as a sink for isotopically heavier, canal-borne dissolved inorganic nitrogen (DIN) and a source for "new" marsh derived dissolved organic nitrogen (DON). In addition, the 15N mesocosm data showed the rapid assimilation of the 15N tracer by the periphyton component and the delayed N uptake by soil and macrophyte components in the southern Everglades.

Spatial scale and abundance patterns of large fish communities in freshwater marshes of the Florida Everglades

Anthropogenic habitat alterations and water-management practices have imposed an artificial spatial scale onto the once contiguous freshwater marshes of the Florida Everglades. To gain insight into how these changes may affect biotic communities, we examined whether variation in the abundance and community structure of large fishes (SL . 8 cm) in Everglades marshes varied more at regional or intraregional scales, and whether this variation was related to hydroperiod, water depth, floating mat volume, and vegetation density. From October 1997 to October 2002, we used an airboat electrofisher to sample large fishes at sites within three regions of the Everglades. Each of these regions is subject to unique watermanagement schedules. Dry-down events (water depth , 10 cm) occurred at several sites during spring in 1999, 2000, 2001, and 2002. The 2001 dry-down event was the most severe and widespread. Abundance of several fishes decreased significantly through time, and the number of days post-dry-down covaried significantly with abundance for several species. Processes operating at the regional scale appear to play important roles in regulating large fishes. The most pronounced patterns in abundance and community structure occurred at the regional scale, and the effect size for region was greater than the effect size for sites nested within region for abundance of all species combined, all predators combined, and each of the seven most abundant species. Non-metric multi-dimensional scaling revealed distinct groupings of sites corresponding to the three regions. We also found significant variation in community structure through time that correlated with the number of days post-dry-down. Our results suggest that hydroperiod and water management at the regional scale influence large fish communities of Everglades marshes.

Responses of sawgrass and spikerush to variation in hydrologic drivers and salinity in Southern Everglades marshes

Aboveground net primary production (ANPP) by the dominant macrophyte and plant community composition are related to the changing hydrologic environment and to salinity in the southern Everglades, FL, USA. We present a new non-destructive ANPP technique that is applicable to any continuously growing herbaceous system. Data from 16 sites, collected from 1998 to 2004, were used to investigate how hydrology and salinity controlled sawgrass (Cladium jamaicense Crantz.) ANPP. Sawgrass live biomass showed little seasonal variation and annual means ranged from 89 to 639 gdw m)2. Mortality rates were 20–35% of live biomass per 2 month sampling interval, for biomass turnover rates of 1.3–2.5 per year. Production by C. jamaicense was manifest primarily as biomass turnover, not as biomass accumulation. Rates typically ranged from 300 to 750 gdw m)2 year)1, but exceeded 1000 gdw m)2 year)1 at one site and were as high as 750 gdw m)2 year)1 at estuarine ecotone sites. Production was negatively related to mean annual water depth, hydroperiod, and to a variable combining the two (depth-days). As water depths and hydroperiods increased in our southern Everglades study area, sawgrass ANPP declined. Because a primary restoration goal is to increase water depths and hydroperiods for some regions of the Everglades, we investigated how the plant community responded to this decline in sawgrass ANPP. Spikerush (Eleocharis sp.) was the next most prominent component of this community at our sites, and 39% of the variability in sawgrass ANPP was explained by a negative relationship with mean annual water depth, hydroperiod, and Eleocharis sp. density the following year. Sawgrass ANPP at estuarine ecotone sites responded negatively to salinity, and rates of production were slow to recover after high salinity years. Our results suggest that ecologists, managers, and the public should not necessarily interpret a decline in sawgrass that may result from hydrologic restoration as a negative phenomenon.

Comparative study of periphyton community structure in long and short-hydroperiod Everglades marshes

The Florida Everglades is a mosaic of short and long-hydroperiod marshes that differ in the depth, duration, and timing of inundation. Algae are important primary producers in widespread Everglades’ periphyton mats, but relationships of algal production and community structure to hydrologic variability are poorly understood. We quantified differences in algal biomass and community structure between periphyton mats in 5 short and 6 long-hydroperiod marshes in Everglades National Park (ENP) in October 2000. We related differences to water depth and total phosphorus (TP) concentration in the water, periphyton and soils. Long and short-hydroperiod marshes differed in water depth (73 cm vs. 13 cm), periphyton TP concentrations (172μg g−1 vs. 107 μg g−1, respectively) and soil TP (284 μg g−1 vs. 145 μg g−1). Periphyton was abundant in both marshes, with short-hydroperiod sites having greater biomass than long-hydroperiod sites (2936 vs. 575 grams ash-free dry mass m−2). A total of 156 algal taxa were identified and separated into diatom (68 species from 21 genera) and “soft algae” (88 non-diatom species from 47 genera) categories for further analyses. Although diatom total abundance was greater in long-hydroperiod mats, diatom species richness was significantly greater in short- hydroperiod periphyton mats (62 vs. 47 diatom taxa). Soft algal species richness was greater in long-hydroperiod sites (81 vs. 67 soft algae taxa). Relative abundances of individual taxa were significantly different among the two site types, with soft algal distributions being driven by water depth, and diatom distributions by water depth and TP concentration in the water and periphyton. Periphyton communities differ between short and long-hydroperiod marshes, but because they share many taxa, alterations in hydroperiod could rapidly

Interactions between nutrient availability and hydroperiod shape macroinvertebrate communities in Florida Everglades marshes

Hydroperiod and nutrient status are known to influence aquatic communities in wetlands, but their joint effects are not well explored. I sampled floating periphyton mat and flocculent detritus (floc) infaunal communities using 6-cm diameter cores at short- and long-hydroperiod and constantly inundated sites across a range of phosphorus (P) availability (total phosphorus in soil, floc and periphyton). Differences in community structure between periphyton and floc microhabitats were greater than any variation attributable to hydroperiod, P availability, or other spatial factors. Multivariate analyses indicated community structure of benthic-floc infauna was driven by hydroperiod, although crowding (no. g−1 AFDM) of individual taxa showed no consistent responses to hydroperiod or P availability. In contrast, community structure of periphyton mat infauna was driven by P availability, while densities of mat infauna (no. m−2) were most influenced by hydroperiod (+correlations). Crowding of mat infauna increased significantly with P availability in short-hydroperiod marshes, but was constant across the P gradient in long-hydroperiod marshes. Increased abundance of floating-periphyton mat infauna with P availability at short-hydroperiod sites may result from a release from predation by small fish. Community structure and density were not different between long-hydroperiod and constantly inundated sites. These results have implications for the use of macroinvertebrates as indicators of water quality in wetlands and suggest the substrate sampled can influence interpretation of ecological responses observed in these communities.

Challenges in using siliceous subfossils as a tool for inferring past water level and hydroperiod in Everglades marshes

Successfully rehabilitating drained wetlands through hydrologic restoration is dependent on defining restoration targets, a process that is informed by pre-drainage conditions, as well as understanding linkages between hydrology and ecosystem structure. Paleoecological records can inform restoration goals by revealing long-term patterns of change, but are dependent on preservation of biomarkers that provide meaningful interpretations of environmental change. In the Florida Everglades, paleohydrological hind-casting could improve restoration forecasting, but frequent drying of marsh soils leads to poor preservation of many biomarkers. To determine the effectiveness of employing siliceous subfossils in paleohydrological reconstructions, we examined diatoms, plant and sponge silico-sclerids from three soil cores in the central Everglades marshes. Subfossil quality varied among cores, but the abundance of recognizable specimens was sufficient to infer 1,000–3,000 years of hydrologic change at decadal to centennial resolution. Phytolith morphotypes were linked to key marsh plant species to indirectly measure fluctuations in water depth. A modern dataset was used to derive diatom-based inferences of water depth and hydroperiod (R2 = 0.63, 0.47; RMSE = 14 cm, 120 days, respectively). Changes in subfossil quality and abundances at centennial time-scales were associated with mid-Holocene climate events including the Little Ice Age and Medieval Warm Period, while decadal-scale fluctuations in assemblage structure during the twentieth century suggested co-regulation of hydrology by cyclical climate drivers (particularly the Atlantic Multidecadal Oscillation) and water management changes. The successful reconstructions based on siliceous subfossils shown here at a coarse temporal scale (i.e., decadal to centennial) advocate for their application in more highly resolved (i.e., subdecadal) records, which should improve the ability of water managers to target the quantity and variability of water flows appropriate for hydrologic restoration.

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.

Tree Islands in the Shark Slough Landscape: Interactions of Vegetation, Hydrology, and Soils

Executive Summary: This report presents what we have learned about tree islands of Shark Slough and adjacent marshes of Everglades National Park (ENP), based on ecological studies carried out in these wetlands during the period 2000-2003. The tree islands of Shark Slough share many features with tree islands elsewhere in the Everglades. Their current composition and community structure is determined to a large extent by recent hydrology, as well as by disturbances (fire, freezes, hurricanes, man). Tree islands have historical, cultural, and biological values that are recognized by nearly all parties to the Comprehensive Everglades Restoration Plan (CERP). Maintaining and/or restoring the health of tree islands are major objectives of CERP. Consequently, there is a need within CERP for tools to assess the health of tree islands, and to relate these measures to the hydrologic regime to which they are exposed.

Accumulation and fate of mercury in an Everglades aquatic food web

This project examined the pathways of mercury (Hg) bioaccumulation and its relation to trophic position and hydroperiod in the Everglades. I described fish-diet differences across habitats and seasons by analyzing stomach contents of 4,000 fishes of 32 native and introduced species. Major foods included periphyton, detritus/algal conglomerate, small invertebrates, aquatic insects, decapods, and fishes. Florida gar, largemouth bass, pike killifish, and bowfin were at the top of the piscine food web. Using prey volumes, I quantitatively classified the fishes into trophic groups of herbivores, omnivores, and carnivores. Stable-isotope analysis of fishes and invertebrates gave an independent and similar assessment of trophic placement. Trophic patterns were similar to those from tropical communities. I tested for correlations of trophic position and total mercury. Over 4,000 fish, 620 invertebrate, and 46 plant samples were analyzed for mercury with an atomic-fluorescence spectrometer. Mercury varied within and among taxa. Invertebrates ranged from 25–200 ng g −1 ww. Small-bodied fishes varied from 78–>400 ng g −1 ww. Large predatory fishes were highest, reaching a maximum of 1,515 ng−1 ww. Hg concentrations in both fishes and invertebrates were positively correlated with trophic position. I examined the effects of season and hydroperiod on mercury in wild and caged mosquitofish at three pairs of marshes. Nine monthly collections of wild mosquitofish were analyzed. Hydroperiod-within-site significantly affected concentrations but it interacted with sampling period. To control for wild-fish dispersal, and to measure in situ uptake and growth, I placed captive-reared, neonate mosquitofish with mercury levels from 7–14 ng g−1 ww into field cages in the six study marshes in six trials. Uptake rates ranged from 0.25–3.61 ng g−1 ww d −1. As with the wild fish, hydroperiod-within-site was a significant main effect that also interacted with sampling period. Survival exceeded 80%. Growth varied with season and hydroperiod, with greatest growth in short-hydroperiod marshes. The results suggest that dietary bioaccumulation determined mercury levels in Everglades aquatic animals, and that, although hydroperiod affected mercury uptake, its effect varied with season. ^

Short and long hydroperiod Everglades periphyton mats: Community characterization and experimental hydroperiod manipulation

Hydroperiod, or the distribution, duration and timing of flooding affects both plant and animal distributions. The Florida Everglades is currently undergoing restoration that will result in altered hydroperiods. This study was conducted in Everglades National Park to document the variability in periphyton community structure and function between long and short hydroperiod Everglades marshes. Periphyton is an important primary producer and important food resource in the Everglades. Periphyton is also involved in marl soil formation and nutrient cycling. Although periphyton is an important component of the Everglades landscape, little is known about periphyton structural-functional variation between hydroperiods. ^ For this study diatoms, as well as fresh algae slides of diatoms, cyanobacteria and green algae were identified and enumerated. Short verse long hydroperiod soil and water column nutrients were compared. Short and long hydroperiod algal periphyton mat productivity rates were compared using BOD incubations. Experimental manipulations were performed to determine the effects of desiccation duration and rewetting on periphyton productivity, community structure, and nutrient flux. ^ Variation in periphyton community structure was significantly greater between hydroperiods than within hydroperiods. Short and long hydroperiod periphyton mats have the same algal species, it is the distribution and abundance that varies between hydroperiods. Long hydroperiod mats have greater diatom abundance while short hydroperiod mats have greater relative filamentous cyanobacterial abundance. ^ Long hydroperiod mats had greater net primary production (npp) than short hydroperiod mats. Short hydroperiod mats respond to rewetting more rapidly than do long hydroperiod mats. Dry short hydroperiod mats became net primary producers within 24 hours of rehydration. Increasing desiccation duration led to greater cyanobacterial abundance in long hydroperiod mats and decreased diatom abundance in both long and short hydroperiod mats. ^

Origin, transport and fate of organic matter in Florida Bay: A biomarker record of historical environmental changes

The Everglades are undergoing the world largest wetland restoration project with the aim of returning this system to hydrological conditions in place prior to anthropogenic modifications. Therefore, it is essential to know what these pristine conditions were. In this work, molecular marker (biomarker) distributions and carbon stable isotopic signatures in sediment samples were employed to assess historical environmental changes in Florida Bay over approximately the last 4000 years. Two biomarkers of terrestrial plants, particularly for mangroves (taraxerol and C29 n-alkane), combined with two seagrass proxies (the Paq and the C25/C 27 n-alkan-2-one ratio) revealed a sedimentary environmental shift from freshwater marshes to mangrove swamps and then to seagrass dominated marine ecosystems, likely as a result of sea-level rise in Florida Bay since the Holocene. The maximum values for the Paq and the C 25/C27 n-alkan-2-ones occurred during the 20th century, suggesting that the greatest abundance of seagrass cover is a recent rather than a historical, long-term phenomenon. The greater oscillation in frequency and amplitude for the biomarkers after 1900 potentially reflects an ecosystem under increasing anthropogenic stress. Several algal biomarkers such as C20 highly branched isoprenoids (HBIs), C 25 HBIs and dinosterol indicative of cyanobacteria, diatom and dinoflagellate organic matter inputs respectively, increased dramatically in the latter part of the 20th century and were attributed to recent anthropogenic changes in Florida Bay. ^ The highlight of this work is the development of HBIs as paleo-proxies. As biomarkers of diatoms, the C25 HBIs in the core from the central bay displayed the highest concentration at mid depth, reflecting strong historical inputs of diatom-derived sedimentary OM during that period. In fact, the depth profile of C25 HBIs coincided quite well with historical variations in diatom abundance and variations in diatom species composition in central Florida Bay based on the results of fossil diatom species analysis by microscopy. This study provides evidence that some C25 HBIs can be applied as biomarkers for certain diatom inputs in paleoenvironmental studies. The sources of C20 and C30 HBIs and their potential applicability as paleo-proxies were also investigated and their sources assessed based on their δ13C distributions. ^

Local biomass control on the composition and reactivity of particulate organic matter in aquatic environments

Freshwater ecosystems have been recognized as important components of the global carbon cycle, and the flux of organic matter (OM) from freshwater to marine environments can significantly affect estuarine and coastal productivity. The focus of this study was the assessment of carbon dynamics in two aquatic environments, namely the Florida Everglades and small prairie streams in Kansas, with the aim of characterizing the biogeochemistry of OM. In the Everglades, particulate OM (POM) is mostly found as a layer of flocculent material (floc). While floc is believed to be the main energy source driving trophic dynamics in this oligotrophic wetland, not much is known about its biogeochemistry. The objective of this study was to determine the origin/sources of OM in floc using biomarkers and pigment-based chemotaxonomy to assess specific biomass contributions to this material, on a spatial (freshwater marshes vs. mangrove fringe) and seasonal (wet vs. dry) scales. It was found that floc OM is derived from the local vegetation (mainly algal components and macrophyte litter) and its composition is controlled by seasonal drivers of hydrology and local biomass productivity. Photo-reactivity experiments showed that light exposure on floc resulted in photo-dissolution of POC with the generation of significant amounts of both dissolved OM (DOM) and nutrients (N & P), potentially influencing nutrient dynamics in this ecosystem. The bio-reactivity experiments determined as the amount and rate of CO2 evolution during incubation were found to vary on seasonal and spatial scales and were highly influenced by phosphorus limitation. Not much is known on OM dynamics in small headwater streams. The objective of this study was to determine carbon dynamics in sediments from intermittent prairie streams, characterized by different vegetation cover for their watershed (C4 grasses) vs. riparian zone (C3 plants). In this study sedimentary OM was characterized using a biomarker and compound specific carbon stable isotope approach. It was found that the biomarker composition of these sediments is dominated by higher plant inputs from the riparian zone, although inputs from adjacent prairie grasses were also apparent. Conflicting to some extent with the River Continuum Concept, sediments of the upper reaches contained more degraded OM, while the lower reaches were enriched in fresh material deriving from higher plants and plankton sources as a result of hydrological regimes and particle sorting.

Cascading ecological effects of low-level phosphorus enrichment in the Florida Everglades

Few studies have examined long-term ecological effects of sustained low-level nutrient enhancement on wetland biota. To determine sustained effects of phosphorus (P) addition on Everglades marshes we added P at low levels (5, 15, and 30 µg L-1 above ambient) for 5 yr to triplicate 100-m flow-through channels in pristine marsh. A cascade of ecological responses occurred in similar sequence among treatments. Although the rate of change increased with dosing level, treatments converged to similar enriched endpoints, characterized most notably by a doubling of plant biomass and elimination of native, calcareous periphyton mats. The full sequence of biological changes occurred without an increase in water total P concentration, which remained near ambient levels until Year 5. This study indicates that Everglades marshes have a near-zero assimilative capacity for P without a state change, that ecosystem responses to enrichment accumulate over time, and that downstream P transport mainly occurs through biota rather than the water column.

Tracking rates of ecotone migration due to salt-water encroachment using fossil mollusks in coastal South Florida

We determined the rate of migration of coastal vegetation zones in response to salt-water encroachment through paleoecological analysis of mollusks in 36 sediment cores taken along transects perpendicular to the coast in a 5.5 km2 band of coastal wetlands in southeast Florida. Five vegetation zones, separated by distinct ecotones, included freshwater swamp forest, freshwater marsh, and dwarf, transitional and fringing mangrove forest. Vegetation composition, soil depth and organic matter content, porewater salinity and the contemporary mollusk community were determined at 226 sites to establish the salinity preferences of the mollusk fauna. Calibration models allowed accurate inference of salinity and vegetation type from fossil mollusk assemblages in chronologically calibrated sediments. Most sediments were shallow (20–130 cm) permitting coarse-scale temporal inferences for three zones: an upper peat layer (zone 1) representing the last 30–70 years, a mixed peat-marl layer (zone 2) representing the previous ca. 150–250 years and a basal section (zone 3) of ranging from 310 to 2990 YBP. Modern peat accretion rates averaged 3.1 mm yr)1 while subsurface marl accreted more slowly at 0.8 mm yr)1. Salinity and vegetation type for zone 1 show a steep gradient with freshwater communities being confined west of a north–south drainage canal constructed in 1960. Inferences for zone 2 (pre-drainage) suggest that freshwater marshes and associated forest units covered 90% of the area, with mangrove forests only present along the peripheral coastline. During the entire pre-drainage history, salinity in the entire area was maintained below a mean of 2 ppt and only small pockets of mangroves were present; currently, salinity averages 13.2 ppt and mangroves occupy 95% of the wetland. Over 3 km2 of freshwater wetland vegetation type have been lost from this basin due to salt-water encroachment, estimated from the mollusk-inferred migration rate of freshwater vegetation of 3.1 m yr)1 for the last 70 years (compared to 0.14 m yr)1 for the pre-drainage period). This rapid rate of encroachment is driven by sea-level rise and freshwater diversion. Plans for rehydrating these basins with freshwater will require high-magnitude re-diversion to counteract locally high rates of sea-level rise.