The interactive effects of predators, resources, and disturbance on freshwater snail populations from the Everglades

The origins of population dynamics depend on interplay between abiotic and biotic factors; the relative importance of each changing across space and time. Predation is a central feature of ecological communities that removes individuals (consumption) and alters prey traits (non-consumptive). Resource quality mitigates non-consumptive predator effects by stimulating growth and reproduction. Disturbance resets predator-prey interactions by removing both. I integrate experiments, time-series analysis, and performance trials to examine the relative importance of these on the population dynamics of a snail species by studying a variety of their traits. A review of ninety-three published articles revealed that snail abundance was much less in the Everglades and similar ecosystems compared to all other freshwater ecosystems considered. Separating consumptive from non-consumptive (cues) predator effects at different phosphorous levels with an experiment determined that phosphorous stimulated, but predator cues inhibited snail growth (34% vs. 23%), activity (38% vs. 53%), and reproductive effort (99% vs. 90%) compared to controls. Cues induced taller shells and smaller openings and moved to refugia where they reduced periphyton by 8%. Consumptive predator effects were minor in comparison. In a reciprocal transplant cage experiment along a predator cue and phosphorous gradient created by a canal, snails grew 10% faster and produced 37% more eggs far from the canal (fewer cues) when fed phosphorous-enriched periphyton from near the canal. Time-series analysis at four sites and predator performance trials reveal that phosphorous-enriched regions support larger snail populations, seasonal drying removes snails at all sites, crayfish negatively affect populations in enriched regions, and molluscivorous fish consume snails in the wet season. Combining these studies reveals interplay between resources, predators, and seasonality that limit snail populations in the Everglades and lead to their low abundance compared to other freshwater ecosystems. Resource quality is emerging as the critical factor because improving resources profoundly improved growth and reproduction; seasonal drying and predation become important at times and places. This work contributes to the general understanding in ecology of the relative importance of different factors that structure populations and provides evidence that bolsters monitoring efforts to assess the Comprehensive Everglades Restoration Plan that show phosphorous enrichment is a major driver of ecosystem change.

Characterization, sources, and transformations of dissolved organic matter (DOM) in Florida coastal Everglades (FCE)

Dissolved organic matter (DOM) is one of the largest carbon reservoirs on this planet and is present in aquatic environments as a highly complex mixture of organic compounds. The Florida coastal Everglades (FCE) is one of the largest wetlands in the world. DOM in this system is an important biogeochemical component as most of the nitrogen (N) and phosphorous (P) are in organic forms. Achieving a better understanding of DOM dynamics in large coastal wetlands is critical, and a particularly important issue in the context of Everglades restoration. In this work, the environmental dynamics of surface water DOM on spatial and temporal scales was investigated. In addition, photo- and bio-reactivity of this DOM was determined, surface-to-groundwater exchange of DOM was investigated, and the size distribution of freshwater DOM in Everglades was assessed. The data show that DOM dynamics in this ecosystem are controlled by both hydrological and ecological drivers and are clearly different on spatial scales and variable seasonally. The DOM reactivity data, modeled with a multi-pool first order degradation kinetics model, found that fluorescent DOM in FCE is generally photo-reactive and bio-refractory. Yet the sequential degradation proved a “priming effect” of sunlight on the bacterial uptake and reworking of this subtropical wetland DOM. Interestingly, specific PARAFAC components were found to have different photo- and bio-degradation rates, suggesting a highly heterogeneous nature of fluorophores associated with the DOM. Surface-to-groundwater exchange of DOM was observed in different regions of the system, and compositional differences were associated with source and photo-reactivity. Lastly, the high degree of heterogeneity of DOM associated fluorophores suggested based on the degradation studies was confirmed through the EEM-PARAFAC analysis of DOM along a molecular size continuum, suggesting that the fluorescence characteristics of DOM are highly controlled by different size fractions and as such can exhibit significant differences in reactivity.

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.

Landscape patterns of periphyton in the Florida Everglades

Periphyton is an abundant and ubiquitous feature of the Florida Everglades, often forming thick mats that blanket shallow sediments and submersed plants. They are considered to be primary ecosystem engineers in the Everglades by forming and stabilizing soils, controlling concentrations of nutrients and gases, and supplying food and structure for other organisms. Distribution patterns are related to underlying physicochemical gradients as well as those hydrologic changes imposed by water management. Because communities respond rapidly to environmental change, their use has been advocated to provide indication of system degradation or restoration. The authors review studies on the distribution of periphyton in the Everglades, highlighting major findings relevant to water management, and also areas where additional exploration is necessary.

Rapid responses of vegetation to hydrological changes in Taylor Slough, Everglades National Park, Florida, USA

We analyzed the dynamics of freshwater marsh vegetation of Taylor Slough in eastern Everglades National Park for the 1979 to 2003 period, focusing on cover of individual plant species and on cover and composition of marsh communities in areas potentially influenced by a canal pump station (‘‘S332’’) and its successor station (‘‘S332D’’). Vegetation change analysis incorporated the hydrologic record at these sites for three intervals: pre-S332 (1961–1980), S332 (1980–1999), post-S332 (1999–2002). During S332 and post-S332 intervals, water level in Taylor Slough was affected by operations of S332 and S332D. To relate vegetation change to plot-level hydrological conditions in Taylor Slough, we developed a weighted averaging regression and calibration model (WA) using data from the marl prairies of Everglades National Park and Big Cypress National Preserve. We examined vegetation pattern along five transects. Transects 1–3 were established in 1979 south of the water delivery structures, and were influenced by their operations. Transects 4 and 5 were established in 1997, the latter west of these structures and possibly under their influence. Transect 4 was established in the northern drainage basin of Taylor Slough, beyond the likely zones of influence of S332 and S332D. The composition of all three southern transects changed similarly after 1979. Where muhly grass (Muhlenbergia capillaris var. filipes) was once dominant, sawgrass (Cladium jamaicense), replaced it, while where sawgrass initially predominated, hydric species such as spikerush (Eleocharis cellulosa Torr.) overtook it. Most of the changes in species dominance in Transects 1–3 occurred after 1992, were mostly in place by 1995–1996, and continued through 1999, indicating how rapidly vegetation in seasonal Everglades marshes can respond to hydrological modifications. During the post-S332 period, these long-term trends began reversing. In the two northern transects, total cover and dominance of both muhly grass and sawgrass increased from 1997 to 2003. Thus, during the 1990’s, vegetation composition south of S332 became more like that of long hydroperiod marshes, but afterward it partially returned to its 1979 condition, i.e., a community characteristic of less prolonged flooding. In contrast, the vegetation change along the two northern transects since 1997 showed little relationship to hydrologic status.

Hurricane disturbance and recovery of energy balance, CO2 fluxes and canopy structure in a mangrove forest of the Florida Everglades

Eddy covariance (EC) estimates of carbon dioxide (CO2) fluxes and energy balance are examined to investigate the functional responses of a mature mangrove forest to a disturbance generated by Hurricane Wilma on October 24, 2005 in the Florida Everglades. At the EC site, high winds from the hurricane caused nearly 100% defoliation in the upper canopy and widespread tree mortality. Soil temperatures down to -50 cm increased, and air temperature lapse rates within the forest canopy switched from statically stable to statically unstable conditions following the disturbance. Unstable conditions allowed more efficient transport of water vapor and CO2 from the surface up to the upper canopy layer. Significant increases in latent heat fluxes (LE) and nighttime net ecosystem exchange (NEE) were also observed and sensible heat fluxes (H) as a proportion of net radiation decreased significantly in response to the disturbance. Many of these impacts persisted through much of the study period through 2009. However, local albedo and MODIS (Moderate Resolution Imaging Spectro-radiometer) data (the Enhanced Vegetation Index) indicated a substantial proportion of active leaf area recovered before the EC measurements began 1 year after the storm. Observed changes in the vertical distribution and the degree of clumping in newly emerged leaves may have affected the energy balance. Direct comparisons of daytime NEE values from before the storm and after our measurements resumed did not show substantial or consistent differences that could be attributed to the disturbance. Regression analyses on seasonal time scales were required to differentiate the storm's impact on monthly average daytime NEE from the changes caused by interannual variability in other environmental drivers. The effects of the storm were apparent on annual time scales, and CO2 uptake remained approximately 250 g C m-2 yr-1 lower in 2009 compared to the average annual values measured in 2004-2005. Dry season CO2 uptake was relatively more affected by the disturbance than wet season values. Complex leaf regeneration dynamics on damaged trees during ecosystem recovery are hypothesized to lead to the variable dry versus wet season impacts on daytime NEE. In contrast, nighttime CO2 release (i.e., nighttime respiration) was consistently and significantly greater, possibly as a result of the enhanced decomposition of litter and coarse woody debris generated by the storm, and this effect was most apparent in the wet seasons compared to the dry seasons. The largest pre- and post-storm differences in NEE coincided roughly with the delayed peak in cumulative mortality of stems in 2007-2008. Across the hurricane-impacted region, cumulative tree mortality rates were also closely correlated with declines in peat surface elevation. Mangrove forest-atmosphere interactions are interpreted with respect to the damage and recovery of stand dynamics and soil accretion processes following the hurricane.

Physiological responses of red mangroves to the climate in the Florida Everglades

This manuscript reports the findings of physiological studies of red mangrove (Rhizophora mangle L.) conducted from June to August 2001 and from May to June 2003 in the Florida Everglades. In situ physiological measurements were made using environmentally controlled gas exchange systems. The field investigations were carried out to define how regional climate constrains mangrove physiology and ecosystem carbon assimilation. In addition, maximum carboxylation and photosynthetic active radiation (PAR) limited carbon assimilation capacities were investigated during the summer season to evaluate whether ecophysiological models developed for mesophyte plant species can be applied to mangroves. Under summertime conditions in the Florida Everglades, maximum foliar carbon dioxide (CO2) assimilation rates reached 18 μmol CO2 m−2 s−1. Peak molar stomatal conductance to water vapor (H2O) diffusion reached 300 mmol H2O m−2 s−1. Maximum carboxylation and PAR‐limited carbon assimilation rates at the foliage temperature of 30°C attained 76.1 ± 23.4 μmol CO2 m−2 s−1 and 128.1 ± 32.9 μmol (e−) m−2 s−1, respectively. Environmental stressors such as the presence of hypersaline conditions and high solar irradiance loading (>500 W m−2 or >1000 μmoles of photons m−2 s−1 of PAR) imposed sharp reductions in carbon assimilation rates and suppressed stomatal conductance. On the basis of both field observations and model analyses, it is also concluded that existing ecophysiological models need to be modified to consider the influences of hypersaline and high radiational loadings on the physiological responses of red mangroves.

Controls on mangrove forest-atmosphere carbon dioxide exchanges in western Everglades National Park

We report on net ecosystem production (NEP) and key environmental controls on net ecosystem exchange (NEE) of carbon dioxide (CO2) between a mangrove forest and the atmosphere in the coastal Florida Everglades. An eddy covariance system deployed above the canopy was used to determine NEE during January 2004 through August 2005. Maximum daytime NEE ranged from −20 to −25 mmol (CO2) m−2 s−1 between March and May. Respiration (Rd) was highly variable (2.81 ± 2.41 mmol (CO2) m−2 s−1), reaching peak values during the summer wet season. During the winter dry season, forest CO2 assimilation increased with the proportion of diffuse solar irradiance in response to greater radiative transfer in the forest canopy. Surface water salinity and tidal activity were also important controls on NEE. Daily light use efficiency was reduced at high (>34 parts per thousand (ppt)) compared to low (ppt) salinity by 46%. Tidal inundation lowered daytime Rd by ∼0.9 mmol (CO2) m−2 s−1 and nighttime Rd by ∼0.5 mmol (CO2) m−2 s−1. The forest was a sink for atmospheric CO2, with an annual NEP of 1170 ± 127 g C m−2 during 2004. This unusually high NEP was attributed to year‐round productivity and low ecosystem respiration which reached a maximum of only 3 g C m−2 d−1. Tidal export of dissolved inorganic carbon derived from belowground respiration likely lowered the estimates of mangrove forest respiration. These results suggest that carbon balance in mangrove coastal systems will change in response to variable salinity and inundation patterns, possibly resulting from secular sea level rise and climate change. Citation: Barr, J. G., V. Engel, J. D. Fuentes,

Periphyton as an indicator of restoration in the Everglades

Periphyton communities dominate primary production in much of the Florida Everglades wetland and therefore contribute to soil production, ecosystem metabolism and secondary production as well as the composition of dependent communities. Decades of research in the Everglades have supported research findings from other wetland types that cumulatively show that periphyton communities respond very rapidly to alterations in the two dominant drivers of wetland structure and function—hydrology and water quality. Hydrology controls periphyton productivity and composition by regulating moisture availability, substrate types available for colonization and supply of nutrients. Nutrients, particularly the limiting nutrient in this system, phosphorus (P), control levels of production and community composition. Because periphyton communities are well-established to be related to hydrology and water quality, an indicator was developed based on three periphyton attributes: abundance, quality (i.e., nutrient content) and community composition. This assessment tool offers a qualitative assessment of ecosystem response to potential changes in management activities at a time scale appropriate for active management. An example is provided of how the indicator can be used to assess the current water quality and hydrological conditions from high-density spatial surveys. Detected patterns of deterioration align with expectations derived from model predictions and known sources of nutrients and unnatural hydrologic regimes. If employed adaptively in ecosystem management, this tool can be used to both detect and react to change before the system has been irreparably altered.

Hydrologic measurements and implications for tree island formation within Everglades National Park

Tree islands in the Shark River Slough of the Everglades National Park (ENP), in the southern state of Florida in the United States, are part of a wetland system of densely vegetated ridges interspersed within relatively open sloughs. Human alteration of this system has had dramatic negative effects on the landscape of the region and restoration efforts will require adjusting the hydrology of the region to assure the preservation of these important ecologic features. The primary objectives of this study were to document the hydrology in the vicinity of tree islands in ENP by measuring velocities in time and space and by characterizing suspended sediments. The results of such measurements were interpreted with respect to factors that may limit tree island growth. The measurements were conducted in the vicinity of three tree islands known as Black Hammock (BH), Gumbo Limbo (GL), and an unnamed island that was named for this study as Satin Leaf (SL). Acoustical Doppler Velocity (ADV) meters were used for measuring the low velocities of the Everglades water flow. Properties of suspended sediments were characterized through measurements of particle size distribution, turbidity, concentration and particle density. Mean velocities observed at each of the tree islands varied from 0.9 to 1.4 cm/s. Slightly higher mean velocities were observed during the wet season (1.2–1.6 cm/s) versus the dry season (0.8–1.3 cm/s). Maximum velocities of more than 4 cm/s were measured in areas of Cladium jamaicense die-off and at the hardwood hammock (head) of the islands. At the island’s head, water is channelized around obstructions such as tree trunks in relatively rapid flow, which may limit the lateral extent of tree island growth. Channelization is facilitated by shade from the tree canopy, which limits the growth of underwater vegetation thereby minimizing the resistance to flow and limiting sediment deposition. Suspended sediment concentrations were low (0.5–1.5 mg/L) at all study sites and were primarily of organic origin. The mean particle size of the suspended sediments was 3 μm with a distribution that was exponential. Critical velocities needed to cause re-suspension of these particles were estimated to be above the actual velocities observed. Sediment transport within the water column appears to be at a near steady state during the conditions evaluated with low rates of sediment loss balanced by presumably the release of equivalent quantities of particles of organic origin. Existing hydrologic conditions do not appear to transport sufficient suspended sediments to result in the formation of tree islands. Of interest would be to collect hydrologic and sediment transport data during extreme hydrologic events to determine if enough sediment is transported under these conditions to promote sufficient sediment accumulations.

Pink shrimp as an indicator for restoration of everglades ecosystems

The pink shrimp, Farfantepenaeus duorarum, familiar to most Floridians as either food or bait shrimp, is ubiquitous in South Florida coastal and offshore waters and is proposed as an indicator for assessing restoration of South Florida's southern estuaries: Florida Bay, Biscayne Bay, and the mangrove estuaries of the lower southwest coast. Relationships between pink shrimp and salinity have been determined in both field and laboratory studies. Salinity is directly relevant to restoration because the salinity regimes of South Florida estuaries, critical nursery habitat for the pink shrimp, will be altered by changes in the quantity, timing, and distribution of freshwater inflow planned as part of the Comprehensive Everglades Restoration Project (CERP). Here we suggest performance measures based on pink shrimp density (number per square meter) in the estuaries and propose a restoration assessment and scoring scheme using these performance measures that can readily be communicated to managers, policy makers, and the interested public. The pink shrimp is an appropriate restoration indicator because of its ecological as well as its economic importance and also because scientific interest in pink shrimp in South Florida has produced a wealth of information about the species and relatively long time series of data on both juveniles in estuarine nursery habitats and adults on the fishing grounds. We suggest research needs for improving the pink shrimp performance measure.

Periphyton responses to eutrophication in the Florida Everglades: Cross-system patterns of structural and compositional change

We examined periphyton along transects in five Everglades marshes and related compositional and functional aspects to phosphorus(P ) gradients caused by enriched inflows. Results were compared to those of a P-addition experiment in a pristine Everglades marsh. While the water total P (TP) concentration was not related to P load in the marshes or experiment the concentration of TP in periphyton was strongly correlated with the distance from the P source. Increased P concentration in periphyton was associated with a loss of biomass,p articularly of the calcifying mat-forming matrix, regardless of the growth form of the periphyton (epiphytic, floating,or epilithic). Diatom species composition was also strongly related to P availability, but the TP optima of many species varied among marshes. Enriched periphyton communities were found 14 km downstream of P inputs to one marsh that has been receiving enhanced P loads for decades, where other studies using different biotic indicators show negligible change in the same marsh. Although recovery trajectories are unknown, periphyton indicators should serve as excellent metrics for the progression or amelioration of P-related effects in the 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.