Across-scale patterning of plant-soil-water interactions surrounding tree islands in Southern Everglades landscapes

The freshwater Everglades is a complex system containing thousands of tree islands embedded within a marsh-grassland matrix. The tree island-marsh mosaic is shaped and maintained by hydrologic, edaphic and biological mechanisms that interact across multiple scales. Preserving tree islands requires a more integrated understanding of how scale-dependent phenomena interact in the larger freshwater system. The hierarchical patch dynamics paradigm provides a conceptual framework for exploring multi-scale interactions within complex systems. We used a three-tiered approach to examine the spatial variability and patterning of nutrients in relation to site parameters within and between two hydrologically defined Everglades landscapes: the freshwater Marl Prairie and the Ridge and Slough. Results were scale-dependent and complexly interrelated. Total carbon and nitrogen patterning were correlated with organic matter accumulation, driven by hydrologic conditions at the system scale. Total and bioavailable phosphorus were most strongly related to woody plant patterning within landscapes, and were found to be 3 to 11 times more concentrated in tree island soils compared to surrounding marshes. Below canopy resource islands in the slough were elongated in a downstream direction, indicating soil resource directional drift. Combined multi-scale results suggest that hydrology plays a significant role in landscape patterning and also the development and maintenance of tree islands. Once developed, tree islands appear to exert influence over the spatial distribution of nutrients, which can reciprocally affect other ecological processes.

Molecular characterization of Cladium peat from the Florida Everglades: biomarker associations with humic fractions

The accumulation and preservation of peat soils in Everglades freshwater marshes and mangrove swamps is an essential process in the ecological functioning of these ecosystems. Human intervention and climate change have modified nutrient dynamics and hydroperiod in the Everglades and peat loss due to such anthropogenic activities is evident. However, not much is known on the molecular level regarding the biogeochemical characteristics, which allow peat to be preserved in the Everglades. Lipid biomarkers trapped within or bound to humic-type structures can provide important geochemical information regarding the origin and microbial transformation of OM in peat. Four lipid fractions obtained from a Cladium peat, namely the freely extractable fraction and those associated with humin, humic acid, and fulvic acid fractions, showed clear differences in their molecular distribution suggesting different OM sources and structural and diagenetic states of the source material. Both, higher plant derived and microbial lipids were found in association with these humic-type substances. Most biomarker distributions suggest an increment in the microbial/terrestrial lipid ratio from the free to humin to humic to fulvic fractions. Microbial reworking of lipids, and the incorporation of microbial biomarkers into the humic-type fractions was evident, as well as the preservation of diagenetic byproducts. The lipid distribution associated with the fulvic acids suggests a high degree of microbial reworking for this fraction. Evidence for this 3D structure was obtained through the presence of the relatively high abundance of α,ω-dicarboxylic acids and phenolic and benzenecarboxylic compounds. The increment in structural complexity of the phenolic and benzencarboxylic compounds in combination with the reduction in the carbon chain length of the dicarboxylic acids from the free to fulvic fraction suggests the latter to be structurally the most stable, compacted and diagenetically altered substrate. This analytical approach can now be applied to peat samples from other areas within the Everglades ecosystem, affected differently by human intervention with the aim to assess changes in organic matter preservation.

Everglades Periphyton: A Biogeochemical Perspective

Periphyton is an important component of the Everglades biogeochemical cycle but remains poorly understood. From a biogeochemical perspective, periphyton is a dense aggregation of diverse microorganisms (autotrophic and heterotrophic) and particles (mineral and detrital) imbedded within an extracellular matrix. The authors synthesize Everglades periphyton biogeochemistry and diversity at the ecosystem and community scales. The primary regulator of biogeochemical processes (material flux, transformation, and storage) is photosynthesis, which controls oxidation-reduction potentials and heterotrophic metabolism. Eutrophication and hydrologic alterations have resulted in fundamental periphyton biogeochemical differences. Elucidation of these processes is required to predict and interpret responses to ecosystem restoration.

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

The Role of the Everglades Mangrove Ecotone Region (EMER) in Regulating Nutrient Cycling and Wetland Productivity in South Florida

The authors summarize the main findings of the Florida Coastal Everglades Long-Term Ecological Research (FCE-LTER) program in the EMER, within the context of the Comprehensive Everglades Restoration Plan (CERP), to understand how regional processes, mediated by water flow, control population and ecosystem dynamics across the EMER landscape. Tree canopies with maximum height <3 m cover 49% of the EMER, particularly in the SE region. These scrub/dwarf mangroves are the result of a combination of low soil phosphorus (P < 59 μg P g dw−1) in the calcareous marl substrate and long hydroperiod. Phosphorus limits the EMER and its freshwater watersheds due to the lack of terrigenous sediment input and the phosphorus-limited nature of the freshwater Everglades. Reduced freshwater delivery over the past 50 years, combined with Everglades compartmentalization and a 10 cm rise in coastal sea level, has led to the landward transgression (1.5 km in 54 years) of the mangrove ecotone. Seasonal variation in freshwater input strongly controls the temporal variation of nitrogen and P exports (99%) from the Everglades to Florida Bay. Rapid changes in nutrient availability and vegetation distribution during the last 50 years show that future ecosystem restoration actions and land use decisions can exert a major influence, similar to sea level rise over the short term, on nutrient cycling and wetland productivity in the EMER.

Biogeochemistry of Nitrogen Across the Everglades Landscape

Compared to phosphorus (P), nitrogen (N) has received little attention across the Everglades landscape. Despite this lack of attention, N plays important roles in many Everglades systems, including being a significant pollutant in Florida Bay and the Gulf of Mexico, the limiting nutrient in highly P-impacted areas, and an important substrate for microbial metabolism. Storage and transport of N throughout the Everglades is dominated by organic forms, including peat soils and dissolved organic N in the water column. In general, N sources are highest in the northern areas; however, atmospheric deposition and active N2 fixation by the periphyton components are a significant N source throughout most systems. Many of the processes involved in the wetland N cycle remain unmeasured for most of the Everglades systems. In particular, the lack of in situ rates for N2 fixation and denitrification prevent the construction of system-level budgets, especially for the Southern mangrove systems where N export into Florida Bay is critical. There is also the potential for several novel N processes (e.g., Anammox) with an as yet undetermined importance for nitrogen cycling and function of the Everglades ecosystem. Phosphorus loading alters the N cycle by stimulating organic N mineralization with resulting flux of ammonium and DON, and at elevated P concentrations, by increasing rates of N2 fixation and N assimilation. Restoration of hydrology has a potential for significantly impacting N cycling in the Everglades both in terms of affecting N transport, but also by altering aerobic-anaerobic transitions at the soil-water interface or in areas with seasonal drawdowns (e.g., marl prairies). Based on the authors’ understanding of N processes, much more research is necessary to adequately predict potential impacts from hydrologic restoration, as well as the function of Everglades systems as sinks, sources, and transformers of N in the South Florida landscape.

Changes in Community Structure of Sediment Bacteria Along the Florida Coastal Everglades Marsh–Mangrove–Seagrass Salinity Gradient

Community structure of sediment bacteria in the Everglades freshwater marsh, fringing mangrove forest, and Florida Bay seagrass meadows were described based on polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE) patterns of 16S rRNA gene fragments and by sequencing analysis of DGGE bands. The DGGE patterns were correlated with the environmental variables by means of canonical correspondence analysis. There was no significant trend in the Shannon–Weiner index among the sediment samples along the salinity gradient. However, cluster analysis based on DGGE patterns revealed that the bacterial community structure differed according to sites. Not only were these salinity/vegetation regions distinct but the sediment bacteria communities were consistently different along the gradient from freshwater marsh, mangrove forest, eastern-central Florida Bay, and western Florida Bay. Actinobacteria- and Bacteroidetes/Chlorobi-like DNA sequences were amplified throughout all sampling sites. More Chloroflexi and members of candidate division WS3 were found in freshwater marsh and mangrove forest sites than in seagrass sites. The appearance of candidate division OP8-like DNA sequences in mangrove sites distinguished these communities from those of freshwater marsh. The seagrass sites were characterized by reduced presence of bands belonging to Chloroflexi with increased presence of those bands related to Cyanobacteria, γ-Proteobacteria, Spirochetes, and Planctomycetes. This included the sulfate-reducing bacteria, which are prevalent in marine environments. Clearly, bacterial communities in the sediment were different along the gradient, which can be explained mainly by the differences in salinity and total phosphorus.

Molecular characterization of proteinaceous material in the Florida coastal Everglades

Dissolved organic nitrogen (DON) is the least known component of the nitrogen cycle, in part as a result of the lack of adequate analytical methods for its molecular characterization. In this study proteinaceous material in DON, collected at six geomorphologically different sites in the Florida coastal Everglades, was characterized by amino acid analysis and protein gel electrophoresis. The amino acid composition of the samples suggests that the canal DON was more degraded and subject to higher microbial inputs than the mangrove marshwater and marine end-member stations. Sodium dodecyl sulfate polyacrylamide gel electrophoresis (SDS-PAGE) and two-dimensional polyacrylamide gel electrophoresis (2D-PAGE) results supported this observation as distinctly different protein profiles were obtained for the canal waters compared to samples collected at other stations. These preliminary results highlight the potential of combining amino acid and intact protein analysis to fingerprint the sources of DON in different aquatic environments, and show SDS-PAGE as a potentially useful method to characterize DON.

Effects of hydrologic and water quality drivers on periphyton dynamics in the southern Everglades

Everglades periphyton mats are tightly-coupled autotrophic (algae and cyanobacteria) and heterotrophic (eubacteria, fungi and microinvertebrates) microbial assemblages. We investigated the effect of water column total phosphorus and nitrogen concentrations, water depth and hydroperiod on periphyton of net production, respiration, nutrient content, and biomass. Our study sites were located along four transects that extended southward with freshwater sheetflow through sawgrass-dominated marsh. The water source for two of the transects were canal-driven and anchored at canal inputs. The two other transects were rain-driven (ombrotrophic) and began in sawgrass-dominated marsh. Periphyton dynamics were examined for upstream and downstream effects within and across the four transects. Although all study sites were characterized as short hydroperiod and phosphorus-limited oligotrophic, they represent gradients of hydrologic regime, water source and water quality of the southern Everglades. Average periphyton net production of 1.08 mg C AFDW−1 h−1 and periphyton whole system respiration of 0.38 mg C AFDW−1 h−1 rates were net autotrophic. Biomass was generally highest at ombrotrophic sites and sites downstream of canal inputs. Mean biomass over all our study sites was high, 1517.30 g AFDW m−2. Periphyton was phosphorus-limited. Average periphyton total phosphorus content was 137.15 μg P g−1 and average periphyton total N:P ratio was 192:1. Periphyton N:P was a sensitive indicator of water source. Even at extremely low mean water total phosphorus concentrations ( ≤ 0.21 μmol l−1), we found canal source effects on periphyton dynamics at sites adjacent to canal inputs, but not downstream of inflows. These canal source effects were most pronounced at the onset of wet season with initial rewetting. Spatial and temporal variability in periphyton dynamics could not solely be ascribed to water quality, but was often associated with both hydrology and water source.

Recent and Historic Drivers of Landscape Change in the Everglades Ridge, Slough, and Tree Island Mosaic

More than half of the original Everglades extent formed a patterned peat mosaic of elevated ridges, lower and more open sloughs, and tree islands aligned parallel to the dominant flow direction. This ecologically important landscape structure remained in a dynamic equilibrium for millennia prior to rapid degradation over the past century in response to human manipulation of the hydrologic system. Restoration of the patterned landscape structure is one of the primary objectives of the Everglades restoration effort. Recent research has revealed that three main drivers regulated feedbacks that initiated and maintained landscape structure: the spatial and temporal distribution of surface water depths, surface and subsurface flow, and phosphorus supply. Causes of recent degradation include but are not limited to perturbations to these historically important controls; shifts in mineral and sulfate supply may have also contributed to degradation. Restoring predrainage hydrologic conditions will likely preserve remaining landscape pattern structure, provided a sufficient supply of surface water with low nutrient and low total dissolved solids content exists to maintain a rainfall-driven water chemistry. However, because of hysteresis in landscape evolution trajectories, restoration of areas with a fully degraded landscape could require additional human intervention.

Benthic diatom assemblages as indicators of water quality in the Everglades and three tropical karstic wetlands

Limestone-based (karstic) freshwater wetlands of the Everglades, Belize, Mexico, and Jamaica are distinctive in having a high biomass of CaCO3-rich periphyton mats. Diatoms are common components of these mats and show predictable responses to environmental variation, making them good candidates for assessing nutrient enrichment in these naturally ultraoligotrophic wetlands. However, aside from in the Everglades of southern Florida, very little research has been done to document the diatoms and their environmental preferences in karstic Caribbean wetlands, which are increasingly threatened by eutrophication. We identified diatoms in periphyton mats collected during wet and dry periods from the Everglades and similar freshwater karstic wetlands in Belize, Mexico, and Jamaica. We compared diatom assemblage composition and diversity among locations and periods, and the effect of the limiting nutrient, P, on species composition among locations. We used periphyton-mat total P (TP) as a metric of availability. A total of 176 diatom species in 45 genera were recorded from the 4 locations. Twenty-three of these species, including 9 that are considered indicative of Everglades diatom flora, were found in all 4 locations. In Everglades and Caribbean sites, we identified assemblages and indicator species associated with low and high periphyton-mat TP and calculated TP optima and tolerances for each indicator species. TP optima and tolerances of indicator species differed between the Everglades and the Caribbean, but weighted averaging models predicted periphyton-mat TP concentrations from diatom assemblages at Everglades (R2  =  0.56) and Caribbean (R2  =  0.85) locations. These results show that diatoms can be effective indicators of water quality in karstic wetlands of the Caribbean, but application of regionally generated transfer functions to distant sites provides less reliable estimates than locally developed functions.

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.

Legacy and Fate of Mercury and Methylmercury in the Florida Everglades

Mass inventories of total Hg (THg) and methylmercury (MeHg) and mass budgets of Hg newly deposited during the 2005 dry and wet seasons were constructed for the Everglades. As a sink for Hg, the Everglades has accumulated 914, 1138, 4931, and 7602 kg of legacy THg in its 4 management units, namely Water Conservation Area (WCA) 1, 2, 3, and the Everglades National Park (ENP), respectively, with most Hg being stored in soil. The current annual Hg inputs account only for 1−2% of the legacy Hg. Mercury transport across management units during a season amounts to 1% or less of Hg storage, except for WCA 2 where inflow inputs can contribute 4% of total MeHg storage. Mass budget suggests distinct spatiality for cycling of seasonally deposited Hg, with significantly lower THg fluxes entering water and floc in ENP than in the WCAs. Floc in WCAs can retain a considerable fraction (around 16%) of MeHg produced from the newly deposited Hg during the wet season. This work is important for evaluating the magnitude of legacy Hg contamination and for predicting the fate of new Hg in the Everglades, and provides a methodological example for large-scale studies on Hg cycling in wetlands.

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.

Nonnative African jewelfish are more fit but not bolder at the invasion front: a trait comparison across an Everglades range expansion

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. Any opinions, findings, conclusions, or recommendations expressed in the material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.