Forest Resource Islands in a Sub-tropical Marsh: Soil–Site Relationships in Everglades Hardwood Hammocks

Spatial heterogeneity in soils is often characterized by the presence of resource-enriched patches ranging in size from a single shrub to wooded thickets. If the patches persist long enough, the primary constraint on production may transition from one limiting environmental factor to another. Tree islands that are scattered throughout the Florida Everglades basin comprise nutrient-enriched patches, or resource islands, in P-limited oligotrophic marshes. We used principal component analysis and multiple regressions to characterize the belowground environment (soil, hydrology) of one type of tree island, hardwood hammocks, and examined its relationship with the three structural variables (basal area, biomass, and canopy height) indicative of site productivity. Hardwood hammocks in the southern Everglades grow on two distinct soil types. The first, consisting of shallow, organic, relatively low-P soils, is common in the seasonally flooded Marl Prairie landscape. In contrast, hammocks on islands embedded in long hydroperiod marsh have deeper, alkaline, mineral soils with extremely high P concentrations. However, this edaphic variation does not translate simply into differences in forest structure and production. Relative water depth was unrelated to all measures of forest structure and so was soil P, but the non-carbonate component of the mineral soil fraction exhibited a strong positive relationship with canopy height. The development of P-enriched forest resource islands in the Everglades marsh is accompanied by the buildup of a mineral soil; however, limitations on growth in mature islands appear to differ substantively from those that dominate incipient stages in the transformation from marsh to forest. Key words: resource island; tree

The White Ibis and Wood Stork as indicators for restoration of the everglades ecosystem

Large numbers of colonially nesting herons, egrets, ibises, storks and spoonbills were one of the defining natural phenomena of the historical Everglades. Reproduction of these species has been tracked over at least a century, and some clear responses to dramatic anthropogenic hydrological alterations have been established. These include a marked decline in nesting populations of several species, and a movement of colonies away from the over-drained estuarine region. Ponding in a large portion of the freshwater marsh has favored species that hunt by sight in deep water (egrets, cf. 25–45 cm), while tactile feeders (ibises and storks) that depend on concentrated prey in shallow water (5–25 cm) have become proportionately much less common. There has been a marked increase in the interval between exceptionally large breeding aggregations of White Ibises (Eudocimus albus). Loss of short hydroperiod wetlands on the margins of the Everglades have delayed nest initiations 1–2 months by Wood Storks (Mycteria americana) resulting in poor nesting success. These responses are consistent with mechanisms that involve foraging, and the availability and production of prey animals, and each of the relationships is highly dependent on hydrology. Here, we define a group of characteristics about wading bird dynamics (= indicators) that collectively track the specific ecological relationships that supported ibises and storks in the past. We suggest four metrics as indicators of restoration success: timing of nesting by storks, the ratio of nesting ibises + storks to Great Egrets, the proportion of all nests located in the estuarine/freshwater ecotone, and the interval between years with exceptionally large ibis nestings. Each of these metrics has historical (e.g., predrainage) data upon which to base expectations for restoration, and the metrics have little measurement error relative to the large annual variation in numbers of nests. In addition to the strong scientific basis for the use of these indicators, wading birds are also a powerful tool for public communication because they have strong aesthetic appeal, and their ecological relationships with water are intuitively understandable. In the interests of communicating with the public and decision-makers, we integrate these metrics into a single-page annual “traffic-light” report card for wading bird responses. Collectively, we believe these metrics offer an excellent chance of detecting restoration of the ecosystem functions that supported historical wading bird nesting patterns.

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.

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.

Temporal Population Genetic Structure of Eastern Mosquitofish in a Dynamic Aquatic Landscape

We analyzed the effect of periodic drying in the Florida Everglades on spatiotemporal population genetic structure of eastern mosquitofish (Gambusia holbrooki). Severe periodic drying events force individuals from disparate sources to mix in dry season relatively deep-water refuges. In 1996 (a wet year) and 1999 (a dry year), we sampled mosquitofish at 20 dry-season refuges distributed in 3 water management regions and characterized genetic variation for 10 allozyme and 3 microsatellite loci. In 1996, most of the ecosystem did not dry, whereas in 1999, many of our sampling locations were isolated by expanses of dried marsh surface. In 1996, most spatial genetic variation was attributed to heterogeneity within regions. In 1999, spatial genetic variation within regions was not significant. In both years, a small but significant amount of variation (less than 1% of the total variation) was partitioned among regions. Variance was consistently greater than zero among long-hydroperiod sites within a region, but not among short-hydroperiod sites within a region, where hydroperiod was measured as time since last marsh surface dry-down forcing fishes into local refuges. In 1996, all sites were in Hardy–Weinberg equilibrium. In 1999, we observed fewer heterozygotes than expected for most loci and sites suggesting a Wahlund effect arising from fish leaving areas that dried and mixing in deep-water refuges.

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.

Disturbance frequency and community structure in a twenty-five year intervention study

Models of community regulation commonly incorporate gradients of disturbance inversely related to the role of biotic interactions in regulating intermediate trophic levels. Higher trophic-level organisms are predicted to be more strongly limited by intermediate levels of disturbance than are the organisms they consume. We used a manipulation of the frequency of hydrological disturbance in an intervention analysis to examine its effects on small-fish communities in the Everglades, USA. From 1978 to 2002, we monitored fishes at one long-hydroperiod (average 350 days) and at one short-hydroperiod (average 259 days; monitoring started here in 1985) site. At a third site, managers intervened in 1985 to diminish the frequency and duration of marsh drying. By the late 1990s, the successional dynamics of density and relative abundance at the intervention site converged on those of the long-hydroperiod site. Community change was manifested over 3 to 5 years following a dry-down if a site remained inundated; the number of days since the most recent drying event and length of the preceding dry period were useful for predicting population dynamics. Community dissimilarity was positively correlated with the time since last dry. Community dynamics resulted from change in the relative abundance of three groups of species linked by life-history responses to drought. Drought frequency and intensity covaried in response to hydrological manipulation at the landscape scale; community-level successional dynamics converged on a relatively small range of species compositions when drought return-time extended beyond 4 years. The density of small fishes increased with diminution of drought frequency, consistent with disturbance-limited community structure; less-frequent drying than experienced in this study (i.e., longer return times) yields predator-dominated regulation of small-fish communities in some parts of the Everglades.

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.

Biomarker Assessment of Spatial and Temporal Changes in the Composition of Flocculent Material (Floc) in the Subtropical Wetland of the Florida Coastal Everglades

Flocculent material (floc) is an important energy source in wetlands. In the Florida Everglades, floc is present in both freshwater marshes and coastal environments and plays a key role in food webs and nutrient cycling. However, not much is known about its environmental dynamics, in particular its biological sources and bio-reactivity. We analysed floc samples collected from different environments in the Florida Everglades and applied biomarkers and pigment chemotaxonomy to identify spatial and seasonal differences in organic matter sources. An attempt was made to link floc composition with algal and plant productivity. Spatial differences were observed between freshwater marsh and estuarine floc. Freshwater floc receives organic matter inputs from local periphyton mats, as indicated by microbial biomarkers and chlorophyll-a estimates. At the estuarine sites, the floc is dominated by mangrove as well as diatom inputs from the marine end-member. The hydroperiod (duration and depth of inundation) at the freshwater sites influences floc organic matter preservation, where the floc at the short-hydroperiod site is more oxidised likely due to periodic dry-down conditions. Seasonal differences in floc composition were not consistent and the few that were observed are likely linked to the primary productivity of the dominant biomass (periphyton in the freshwater marshes and mangroves in the estuarine zone). Molecular evidence for hydrological transport of floc material from the freshwater marshes to the coastal fringe was also observed. With the on-going restoration of the Florida Everglades, it is important to gain a better understanding of the biogeochemical dynamics of floc, including its sources, transformations and reactivity.

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. ^

Defining the role of floating periphyton mats in shaping food-web dynamics in the Florida Everglades

Expansive periphyton mats are a striking characteristic of the Florida Everglades. Floating periphyton mats are home to a diverse macroinvertebrate community dominated by chironomid and ceratopogonid larvae and amphipods that use the mat as both a food resource and refuge from predation. While this periphyton complex functions as a self-organizing system, it also serves as a base for trophic interactions with larger organisms. The purpose of my research was to quantify variation in the macroinvertebrate community inhabiting floating periphyton mats, describe the role of mats in shaping food-web dynamics, and describe how these trophic interactions change with eutrophication. ^ I characterized the macroinvertebrate community inhabiting periphyton through a wet-season by describing spatial variation on scales from 0.2 m to 3 km. Floating periphyton mats contained a diverse macroinvertebrate community, with greater taxonomic richness and higher densities of many taxa than adjacent microhabitats. Macroinvertebrate density increased through the wet season as periphyton mats developed. While some variation was noted among sites, spatial patterns were not observed on smaller scales. I also sampled ten sites representing gradients of hydroperiod and nutrient (P) levels. The density of macroinvertebrates inhabiting periphyton mats increased with increasing P availability; however, short-hydroperiod P-enriched sites had the highest macroinvertebrate density. This pattern suggests a synergistic interaction of top-down and bottom-up effects. In contrast, macroinvertebrate density was lower in benthic floc, where it was negatively correlated with hydroperiod. ^ I used two types of mesocosms (field cages and tanks) to manipulate large consumers (fish and grass shrimp) with inclusion/exclusion cages over an experimental P gradient. In most cases, periphyton mats served as an effective predation refuge. Macroinvertebrates were consumed more frequently in P-enriched treatments, where mats were also heavily grazed. Macroinvertebrate densities decreased with increasing P in benthic floc, but increased with enrichment in periphyton mats until levels were reached that caused disassociation of the mat. ^ This research documents several indirect trophic interactions that can occur in complex habitats, and emphasizes the need to characterize dynamics of all microhabitats to fully describe the dynamics of an ecosystem. ^

Vegetation:environment relationships and water management in Shark Slough, Everglades National Park

The hydrologic regime of Shark Slough, the most extensive long hydroperiod marsh in Everglades National Park, is largely controlled by the location, volume, and timing of water delivered to it through several control structures from Water Conservation Areas north of the Park. Where natural or anthropogenic barriers to water flow are present, water management practices in this highly regulated system may result in an uneven distribution of water in the marsh, which may impact regional vegetation patterns. In this paper, we use data from 569 sampling locations along five cross-Slough transects to examine regional vegetation distribution, and to test and describe the association of marsh vegetation with several hydrologic and edaphic parameters. Analysis of vegetation:environment relationships yielded estimates of both mean and variance in soil depth, as well as annual hydroperiod, mean water depth, and 30-day maximum water depth within each cover type during the 1990’s. We found that rank abundances of the three major marsh cover types (Tall Sawgrass, Sparse Sawgrass, and Spikerush Marsh) were identical in all portions of Shark Slough, but regional trends in the relative abundance of individual communities were present. Analysis also indicated clear and consistent differences in the hydrologic regime of three marsh cover types, with hydroperiod and water depths increasing in the order Tall Sawgrass , Sparse Sawgrass , Spikerush Marsh. In contrast, soil depth decreased in the same order. Locally, these differences were quite subtle; within a management unit of Shark Slough, mean annual values for the two water depth parameters varied less than 15 cm among types, and hydroperiods varied by 65 days or less. More significantly, regional variation in hydrology equaled or exceeded the variation attributable to cover type within a small area. For instance, estimated hydroperiods for Tall Sawgrass in Northern Shark Slough were longer than for Spikerush Marsh in any of the other regions. Although some of this regional variation may reflect a natural gradient within the Slough, a large proportion is the result of compartmentalization due to current water management practices within the marsh.We conclude that hydroperiod or water depth are the most important influences on vegetation within management units, and attribute larger scale differences in vegetation pattern to the interactions among soil development, hydrology and fire regime in this pivotal portion of Everglades.

Seasonal Fish Dispersal in Ephemeral Wetlands of the Florida Everglades

We hypothesized that fishes in short-hydroperiod wetlands display pulses in activity tied to seasonal flooding and drying, with relatively low activity during intervening periods. To evaluate this hypothesis, sampling devices that funnel fish into traps (drift fences) were used to investigate fish movement across the Everglades, U.S.A. Samples were collected at six sites in the Rocky Glades, a seasonally flooded karstic habitat located on the southeastern edge of the Everglades. Four species that display distinct recovery patterns following drought in long-hydroperiod wetlands were studied: eastern mosquitofish (Gambusia holbrooki) and flagfish (Jordanella floridae) (rapid recovery); and bluefin killifish (Lucania goodei) and least killifish (Heterandria formosa) (slow recovery). Consistent with our hypothesized conceptual model, fishes increased movement soon after flooding (immigration period) and just before drying (emigration period), but decreased activity in the intervening foraging period. We also found that eastern mosquitofish and flagfish arrived earlier and showed stronger responses to hydrological variation than either least killifish or bluefin killifish. We concluded that these fishes actively colonize and escape ephemeral wetlands in response to flooding and drying, and display species-specific differences related to flooding and drying that reflect differences in dispersal ability. These results have important implications for Everglades fish metacommunity dynamics.

A Geospatial Database of Tree Islands within the Mustang Corner Fire Incident of 2008

Fire, which affects community structure and composition at all trophic levels, is an integral component of the Everglades ecosystem (Wade et al. 1980; Lockwood et al. 2003). Without fire, the Everglades as we know it today would be a much different place. This is particularly true for the short-hydroperiod marl prairies that predominate on the eastern and western flanks of Shark River Slough, Everglades National Park (Figure 1). In general, fire in a tropical or sub-tropical grassland community favors the dominance of C4 grasses over C3 species (Roscoe et al. 2000; Briggs et al. 2005). Within this pyrogenic graminoid community also, periodic natural fires, together with suitable hydrologic regime, maintain and advance the dominance of C4 vs C3 graminoids (Sah et al. 2008), and suppress the encroachment of woody stems (Hanan et al. 2009; Hanan et al. unpublished manuscript) originating from the tree islands that, in places, dominate the landscape within this community. However, fires, under drought conditions and elevated fuel loads, can spread quickly throughout the landscape, oxidizing organic soils, both in the prairie and in the tree islands, and, in the process, lead to shifts in vegetation composition. This is particularly true when a fire immediately precedes a flood event (Herndon et al. 1991; Lodge 2005; Sah et al. 2010), or if so much soil is consumed during the fire that the hydrologic regime is permanently altered as a result of a decrease in elevation (Zaffke 1983).

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

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