Hydroperiod and seasonal effects on fish decomposition in an oligotrophic Everglades marsh

In the Everglades, the majority of fish detrital inputs occur during the dry scason, when waterlevel drawdown reduces aquatic habitat. While these mortality events are highly seasonal, the remineralization and recycling of fish detrital nutrients may represent an important stimulus to the ecosystem in the following wet season. The goal of this study was to quantify the rate of detrital fish decomposition during three periods of the year to determine seasonal variations in decomposition patterns in this ecosystem. A multiple regression analysis showed that hydroperiod and water depth both played a role in determining fish decomposition rates within this ecosystem. Decomposition rates ranged from a low of 13% day−1 in December 2000 to a high of 50% day−1 in April 2001, the height of the dry season. Phosphorus analysis showed that Gambusia holbrooki, the dominant small fish species in the Everglades, contains 7.169±1.46 mg P g−1 wet fish weight. Based on the observed decomposition rates and the average biomass added, the estimafed daily flux of phosphorus from the experimental detrital loading ranged from a low of 27.04 mg P day−1 to a high of 108.14 mg P day−1 during the decomposition period. We estimated that these inputs could represent an input of 43 μg P m−2 day−1 to the total temporal Everglades phosphorus budget. Although much of this phosphorus is likely incorporated into the macroinvertebrate pool, detrital inputs peak during the dry season when nutrients are most likely to be incorporated into the soil and occur when decomposition of vegetative material is moisture-limited. These inputs may therefore play an important role in stimulating vegetative production during the early wet season.

Quantifying the responses of calcareous periphyton crusts to rehydration: A microcosm study (Florida Everglades)

We examined the high-resolution temporal dynamics of recovery of dried periphyton crusts following rapid rehydration in a phosphorus (P)-limited short hydroperiod Everglades wetland. Crusts were incubated in a greenhouse in tubs containing water with no P or exogenous algae to mimic the onset of the wet season in the natural marsh when heavy downpours containing very low P flood the dry wetland. Algal and bacterial productivity were tracked for 20 days and related to compositional changes and P dynamics in the water. A portion of original crusts was also used to determine how much TP could be released if no biotic recovery occurred. Composition was volumetrically dominated by cyanobacteria (90%) containing morphotypes typical of xeric environments. Algal and bacterial production recovered immediately upon rehydration but there was a net TP loss from the crusts to the water in the first 2 days. By day 5, however, cyanobacteria and other bacteria had re-absorbed 90% of the released P. Then, water TP concentration reached a steady-state level of 6.6 μg TP/L despite water TP concentration through evaporation. Phosphomonoesterase (PMEase) activity was very high during the first day after rehydration due to the release of a large pre-existing pool of extracellular PMEase. Thereafter, the activity dropped by 90% and increased gradually from this low level. The fast recovery of desiccated crusts upon rehydration required no exogenous P or allogenous algae/bacteria additions and periphyton largely controlled P concentration in the water.

Ecosystem structure, nutrient dynamics, and hydrologic relationships in tree islands of the southern Everglades, Florida, USA

Tree islands are an important structural component of many graminoid-dominated wetlands because they increase ecological complexity in the landscape. Tree island area has been drastically reduced with hydrologic modifications within the Everglades ecosystem, yet still little is known about the ecosystem ecology of Everglades tree islands. As part of an ongoing study to investigate the effects of hydrologic restoration on short hydroperiod marshes of the southern Everglades, we report an ecosystem characterization of seasonally flooded tree islands relative to locations described by variation in freshwater flow (i.e. locally enhanced freshwater flow by levee removal). We quantified: (1) forest structure, litterfall production, nutrient utilization, soil dynamics, and hydrologic properties of six tree islands and (2) soil and surface water physico-chemical properties of adjacent marshes. Tree islands efficiently utilized both phosphorus and nitrogen, but indices of nutrient-use efficiency indicated stronger P than N limitation. Tree islands were distinct in structure and biogeochemical properties from the surrounding marsh, maintaining higher organically bound P and N, but lower inorganic N. Annual variation resulting in increased hydroperiod and lower wet season water levels not only increased nitrogen use by tree species and decreased N:P values of the dominant plant species (Chrysobalanus icaco), but also increased soil pH and decreased soil temperature. When compared with other forested wetlands, these Everglades tree islands were among the most nutrient efficient, likely a function of nutrient immobilization in soils and the calcium carbonate bedrock. Tree islands of our study area are defined by: (1) unique biogeochemical properties when compared with adjacent short hydroperiod marshes and other forested wetlands and (2) an intricate relationship with marsh hydrology. As such, they may play an important and disproportionate role in nutrient and carbon cycling in Everglades wetlands. With the loss of tree islands that has occurred with the degradation of the Everglades system, these landscape processes may have been altered. With this baseline dataset, we have established a long-term ecosystem-scale experiment to follow the ecosystem trajectory of seasonally flooded tree islands in response to hydrologic restoration of the southern Everglades.

Relationships Between Hydrology and Soils Describe Vegetation Patterns in Seasonally Flooded Tree Islands of the Southern Everglades, Florida

Tree island ecosystems are important and distinct features of Florida Everglades wetlands. We described the inter-relationships among abiotic factors describing seasonally flooded tree islands and characterized plant–soil relationships in tree islands occurring in a relatively unimpacted area of the Everglades. We used Principal Components Analysis (PCA) to reduce our multi-factor dataset, quantified forest structure and vegetation nutrient dynamics, and related these vegetation parameters to PCA summary variables using linear regression analyses. We found that, of the 21 abiotic parameters used to characterize the ecosystem structure of seasonally flooded tree islands, 13 parameters were significantly correlated with four principal components, and they described 78% of the variance among the study islands. Most variation was described by factors related to soil oxidation and hydrology, exemplifying the sensitivity of tree island structure to hydrologic conditions. PCA summary variables describing tree island structure were related to variability in Chrysobalanus icaco (L.) canopy cover, Ilex cassine (L.) and Salix caroliniana (Michx.) canopy cover, Myrica cerifera (L.) plot frequency, litter turnover, % phosphorus resorption of co-dominant species, and nitrogen nutrient-use efficiency. This study supported findings that vegetation characteristics can be sensitive indicators of variability in tree island ecosystem structure. This study produced valuable, information which was used to recommend ecological targets (i.e. restoration performance measures) for seasonally flooded tree islands in more impacted regions of the Everglades landscape.

Biogeochemical Processes on Tree Islands in the Greater Everglades: Initiating a New Paradigm

Scientists’ understanding of the role of tree islands in the Everglades has evolved from a plant community of minor biogeochemical importance to a plant community recognized as the driving force for localized phosphorus accumulation within the landscape. Results from this review suggest that tree transpiration, nutrient infiltration from the soil surface, and groundwater flow create a soil zone of confluence where nutrients and salts accumulate under the head of a tree island during dry periods. Results also suggest accumulated salts and nutrients are flushed downstream by regional water flows during wet periods. That trees modulate their environment to create biogeochemical hot spots and strong nutrient gradients is a significant ecological paradigm shift in the understanding of the biogeochemical processes in the Everglades. In terms of island sustainability, this new paradigm suggests the need for distinct dry-wet cycles as well as a hydrologic regime that supports tree survival. Restoration of historic tree islands needs further investigation but the creation of functional tree islands is promising.

Maintaining tree islands in the Florida Everglades: nutrient redistribution is the key

The Florida Everglades is an oligotrophic wetland system with tree islands as one of its most prominent landscape features. Total soil phosphorus concentrations on tree islands can be 6 to 100 times greater than phosphorus levels in the surrounding marshes and sloughs, making tree islands nutrient hotspots. Several mechanisms are believed to redistribute phosphorus to tree islands: subsurface water flows generated by evapotranspiration of trees, higher deposition rates of dry fallout, deposition of guano by birds and other animals, groundwater upwelling, and bedrock mineralization by tree exudates. A conceptual model is proposed, in which the focused redistribution of limiting nutrients, especially phosphorus, onto tree islands controls their maintenance and expansion. Because of increased primary production and peat accretion rates, the redistribution of phosphorus can result in an increase in both tree island elevation and size. Human changes to hydrology have greatly decreased the number and size of tree islands in parts of the Everglades. The proposed model suggests that the preservation of existing tree islands, and ultimately of the Everglades landscape, requires the maintenance of these phosphorus redistribution mechanisms.

Historic primary producer communities linked to water quality and hydrologic changes in the northern Everglades

The northern Everglades Water Conservation Areas have experienced recent ecological shifts in primary producer community structure involving marl periphyton mats and dense Typha-dominated macrophyte stands. Multiple investigations have identified phosphorus (P) as a driver of primary producer community structure, but effects of water impoundment beginning in the 1950s and changes in water hardness [e.g., (CaCO3)] have also been identified as a concern. In an effort to understand pre-1950, primary producer community structure and identify community shifts since 1950, we measured pigment proxies on three sediment cores collected in Water Conservation Area-2A (WCA-2A) along a phosphorus enrichment gradient. Photosynthetic pigments, sediment total phosphorus content (TP), organic matter, total organic carbon and nitrogen were used to infer historic primary producer communities and changes in water quality and hydrology regulating those communities. Excess 210Pb was used to establish historic dates for the sediment cores. Results indicate the northern area of WCA-2A increased marl deposition and increased algal abundance ca. 1920. This increase in (presumably) calcareous periphyton before intensive agriculture and impoundment suggest canal-derived calcium inputs and to some extent early drainage effects played a role in initiating this community shift. The northern area community then shifted to Typha dominance around 1965. The areas to the south in WCA-2A experienced increased marl deposition and algal abundance around or just prior to 1950s impoundment, the precise timing limited by core age resolution. Continued increases in algal abundance were evident after 1950, coinciding with impoundment and deepening of canals draining into WCA-2A, both likely increasing water mineral and nutrient concentrations. The intermediate site developed a Typha-dominated community ca. 1995 while the southern-most core site WCA-2A has yet to develop Typha dominance. Numerous studies link sediment TP >650 mg P/kg to marsh habitat degradation into Typha-dominance. The northern and intermediate cores where Typha is currently support this previous research by showing a distinct shift in the sediment record to Typha dominance corresponding to sediment TP between 600 and 700 mg P/kg. These temporal and spatial differences are consistent with modern evidence showing water-column gradients in mineral inputs (including Ca, carbonates, and phosphorus) altering primary producer community structure in WCA-2A, but also suggest hydroperiod has an effect on the mechanisms regulating periphyton development and Typha dominance.

Indirect and direct controls of macroinvertebrates and small fish by abiotic factors and trophic interactions in the Florida Everglades

1. The roles of nutrients, disturbance and predation in regulating consumer densities have long been of interest, but their indirect effects have rarely been quantified in wetland ecosystems. The Florida Everglades contains gradients of hydrological disturbance (marsh drying) and nutrient enrichment (phosphorus), often correlated with densities of macroinvertebrate infauna (macroinvertebrates inhabiting periphyton), small fish and larger invertebrates, such as snails, grass shrimp, insects and crayfish. However, most causal relationships have yet to be quantified. 2.  We sampled periphyton (content and community structure) and consumer (small omnivores, carnivores and herbivores, and infaunal macroinvertebrates inhabiting periphyton) density at 28 sites spanning a range of hydrological and nutrient conditions and compared our data to seven a priori structural equation models. 3.  The best model included bottom-up and top-down effects among trophic groups and supported top-down control of infauna by omnivores and predators that cascaded to periphyton biomass. The next best model included bottom-up paths only and allowed direct effects of periphyton on omnivore density. Both models suggested a positive relationship between small herbivores and small omnivores, indicating that predation was unable to limit herbivore numbers. Total effects of time following flooding were negative for all three consumer groups even when both preferred models suggested positive direct effects for some groups. Total effects of nutrient levels (phosphorus) were positive for consumers and generally larger than those of hydrological disturbance and were mediated by changes in periphyton content. 4.  Our findings provide quantitative support for indirect effects of nutrient enrichment on consumers, and the importance of both algal community structure and periphyton biomass to Everglades food webs. Evidence for top-down control of infauna by omnivores was noted, though without substantially greater support than a competing bottom-up-only model.

A preliminary analysis of the correlation of food-web characteristics with hydrology and nutrient gradients in the southern Everglades

We estimated trophic position and carbon source for three consumers (Florida gar, Lepisosteus platyrhincus; eastern mosquitofish, Gambusia holbrooki; and riverine grass shrimp, Palaemonetes paludosus) from 20 sites representing gradients of productivity and hydrological disturbance in the southern Florida Everglades, U.S.A. We characterized gross primary productivity at each site using light/dark bottle incubation and stem density of emergent vascular plants. We also documented nutrient availability as total phosphorus (TP) in floc and periphyton, and the density of small fishes. Hydrological disturbance was characterized as the time since a site was last dried and the average number of days per year the sites were inundated for the previous 10 years. Food-web attributes were estimated in both the wet and dry seasons by analysis of δ15N (trophic position) and δ13C (food-web carbon source) from 702 samples of aquatic consumers. An index of carbon source was derived from a two-member mixing model with Seminole ramshorn snails (Planorbella duryi) as a basal grazing consumer and scuds (amphipods Hyallela azteca) as a basal detritivore. Snails yielded carbon isotopic values similar to green algae and diatoms, while carbon values of scuds were similar to bulk periphyton and floc; carbon isotopic values of cyanobacteria were enriched in C13compared to all consumers examined. A carbon source similar to scuds dominated at all but one study site, and though the relative contribution of scud-like and snail-like carbon sources was variable, there was no evidence that these contributions were a function of abiotic factors or season. Gar consistently displayed the highest estimated trophic position of the consumers studied, with mosquitofish feeding at a slightly lower level, and grass shrimp feeding at the lowest level. Trophic position was not correlated with any nutrient or productivity parameter, but did increase for grass shrimp and mosquitofish as the time following droughts increased. Trophic position of Florida gar was positively correlated with emergent plant stem density.

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.

Biogeochemical Effects of Simulated Sea Level Rise on Carbon Loss in an Everglades Mangrove Peat Soil

Saltwater intrusion and inundation can affect soil microbial activity, which regulates the carbon (C) balance in mangroves and helps to determine if these coastal forests can keep pace with sea level rise (SLR). This study evaluated the effects of increased salinity (+15 ppt), increased inundation (−8 cm), and their combination, on soil organic C loss from a mangrove peat soil (Everglades, Florida, USA) under simulated tides. Soil respiration (CO2 flux), methane (CH4) flux, dissolved organic carbon (DOC) production, and porewater nutrient concentrations were quantified. Soil respiration was the major pathway of soil organic C loss (94–98%) and was approximately 90% higher in the control water level than the inundated treatment under elevated salinity. Respiration rate increased with water temperature, but depended upon salinity and tidal range. CH4 flux was minimal, while porewater DOC increased with a concomitant, significant decline in soil bulk density under increased inundation. Porewater ammonium increased (73%) with inundation and soluble reactive phosphorus increased (32%) with salinity. Overall, the decline in soil organic C mineralization from combined saltwater intrusion and prolonged inundation was not significant, but results suggest SLR could increase this soil’s susceptibility to peat collapse and accelerate nutrient and DOC export to adjacent Florida Bay.

Effect of Water Management on Interannual Variation in Bulk Soil Properties from the Eastern Coastal Everglades

We examined interannual variation in soil properties from wetlands occurring in adjacent drainage basins from the southeastern Everglades. Triplicate 10-cm soil cores were collected, homogenized, and analyzed during the wet season 2006–2010 from five freshwater sawgrass wetland marshes and three estuarine mangrove forests. Soil bulk density from the Taylor Slough basin ranged from 0.15 gm-cm−3 to 0.5 gm-cm−3, was higher than from the Panhandle basin every year, and generally increased throughout the study period. Organic matter as a percent loss on ignition ranged from 7 % to 12 % from freshwater marshes and from 13 % to 56 % from estuarine mangroves. Extractable iron in soils was similar among drainage basins and wetland types, typically ranging from 0.6 to 2.0 g Fe kg−1. In contrast, inorganic sulfur was on average over four times higher from estuarine soils relative to freshwater, and was positively correlated with soil organic matter. Finally total soil phosphorus (P) was lower in freshwater soils relative to estuarine soils (84 ± 5 versus 326 ± 32 mg P kg−1). Total P from the freshwater marshes in the Panhandle basin rose throughout the study period from 54.7 ± 8.4 to 107 ± 17 mg P kg−1, a possible outcome of differences in water management between drainage basins.

Boundary Effects on Benthic Microbial Phosphorus Concentrations and Diatom Beta Diversity in a Hydrologically-modified, Nutrient-limited Wetland

Water flow and flooding duration in wetlands influence the structure and productivity of microbial communities partly through their influence on nutrient loading. The effect of flow-regulated nutrient loads is especially relevant for microbial communities in nutrient-poor settings, where delivery controls nutrient uptake rates and the intensity of microbial interactions. We examined the effect of hydrologic history and proximity to water sources on nutrient enrichment of benthic microbial assemblages (periphyton) and on their diatom species composition, along the artificial boundaries of Taylor Slough, a historically phosphorus-depleted drainage of the Florida Everglades. Concentrations of phosphorus in periphyton declined from the wetland boundary near inflow structures to 100-m interior, with spatial and temporal variability in rates dependent on proximity to and magnitude of water flow. Phosphorus availability influenced the beta diversity of diatom assemblages, with higher values near inflow structures where resources were greatest, while interior sites and reference transects contained assemblages with constant composition of taxa considered endemic to the Everglades. This research shows how hydrologic restoration may have unintended consequences when incoming water quality is not regulated, including a replacement of distinctive microbial assemblages by ubiquitous, cosmopolitan ones.

Benthic Exchange of C, N, and P Along the Estuarine Ecotone of Lower Taylor Slough, Florida (USA): Effect of Seasonal Flows and Phosphorus Availability

The southern Everglades and Florida Bay have experienced a nearly 50 % reduction in freshwater flow resulting in increased salinity and landward expansion of mangrove forest. Given the marine end-member is a natural source of P to this region, it is necessary to understand the interactions between inflows and P availability in controlling the exchange of materials across the mangrove ecotone. From 2007 to 2008, we used sediment core incubations to quantify fluxes of dissolved inorganic N and P and dissolved organic carbon (DOC) in three ecotone areas (dwarf mangrove, pond, and bay). Experiments were repeated seasonally over 2 years involving P-enriched surface water as a factor. We saw consistent uptake of soluble reactive P (SRP), DOC, and nitrate + nitrite (N+N) by the soils/sediments and release of ammonium (NH4 +) from soils/sediments to the water column across all sites and seasons. P enrichment had no discernible effect on DIN or DOC flux, suggesting that rapid P uptake may have been more geochemically mediated. However, uptake of added P occurred across all sites and seasons, reflecting high uptake capacity in this carbonate system and the potential of the mangrove ecotone to sequester P as it becomes more available.

Trajectories of Vegetation Response to Water Management in Taylor Slough, Everglades National Park, Florida

Ecosystem management practices that modify the major drivers and stressors of an ecosystem often lead to changes in plant community composition. This paper examines how closely the trajectory of vegetation change in seasonally-flooded wetlands tracks management-induced alterations in hydrology and soil characteristics. We used trajectory analysis, a multivariate method designed to test hypotheses about rates and directions of community change, to examine vegetation shifts in response to changes in water management practices within the Taylor Slough basin of Everglades National Park. We summarized vegetation data by non-metric multidimensional scaling ordination, and examined the time trajectory of each site along environmental vectors representing hydrology and soil phosphorus gradients. In the Taylor Slough basin, vegetation change trajectories closely followed the hydrologic changes caused by the operation of water pumps and detention ponds adjacent to the canals. We also observed a shift in vegetation composition along a vector of increasing soil phosphorus, which suggests the need for implementing measures to avoid P-enrichment in southern Everglades marl prairies. This study indicates that shifts in vegetation composition in response to changes in hydrologic conditions and associated parameters may be detected through trajectory analysis, thereby providing feedback for adaptive management of wetland ecosystems.