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.

Comparative study of dissolved organic matter from groundwater and surface water in the Florida coastal Everglades using multi-dimensional spectrofluorometry combined with multivariate statistics

Dissolved organic matter (DOM) in groundwater and surface water samples from the Florida coastal Everglades were studied using excitation–emission matrix fluorescence modeled through parallel factor analysis (EEM-PARAFAC). DOM in both surface and groundwater from the eastern Everglades S332 basin reflected a terrestrial-derived fingerprint through dominantly higher abundances of humic-like PARAFAC components. In contrast, surface water DOM from northeastern Florida Bay featured a microbial-derived DOM signature based on the higher abundance of microbial humic-like and protein-like components consistent with its marine source. Surprisingly, groundwater DOM from northeastern Florida Bay reflected a terrestrial-derived source except for samples from central Florida Bay well, which mirrored a combination of terrestrial and marine end-member origin. Furthermore, surface water and groundwater displayed effects of different degradation pathways such as photodegradation and biodegradation as exemplified by two PARAFAC components seemingly indicative of such degradation processes. Finally, Principal Component Analysis of the EEM-PARAFAC data was able to distinguish and classify most of the samples according to DOM origins and degradation processes experienced, except for a small overlap of S332 surface water and groundwater, implying rather active surface-to-ground water interaction in some sites particularly during the rainy season. This study highlights that EEM-PARAFAC could be used successfully to trace and differentiate DOM from diverse sources across both horizontal and vertical flow profiles, and as such could be a convenient and useful tool for the better understanding of hydrological interactions and carbon biogeochemical cycling.

Sediment and Nutrient Deposition Associated with Hurricane Wilma in Mangroves of the Florida Coastal Everglades

The distribution of mangrove biomass and forest structure along Shark River estuary in the Florida Coastal Everglades (FCE) has been correlated with elevated total phosphorus concentration in soils thought to be associated with storm events. The passage of Hurricane Wilma across Shark River estuary in 2005 allowed us to quantify sediment deposition and nutrient inputs in FCE mangrove forests associated with this storm event and to evaluate whether these pulsing events are sufficient to regulate nutrient biogeochemistry in mangrove forests of south Florida. We sampled the spatial pattern of sediment deposits and their chemical properties in mangrove forests along FCE sites in December 2005 and October 2006. The thickness (0.5 to 4.5 cm) of hurricane sediment deposits decreased with distance inland at each site. Bulk density, organic matter content, total nitrogen (N) and phosphorus (P) concentrations, and inorganic and organic P pools of hurricane sediment deposits differed from surface (0–10 cm) mangrove soils at each site. Vertical accretion resulting from this hurricane event was eight to 17 times greater than the annual accretion rate (0.30± 0.03 cm year−1) averaged over the last 50 years. Total P inputs from storm-derived sediments were equivalent to twice the average surface soil nutrient P density (0.19 mg cm−3). In contrast, total N inputs contributed 0.8 times the average soil nutrient N density (2.8 mg cm−3). Allochthonous mineral inputs from Hurricane Wilma represent a significant source of sediment to soil vertical accretion rates and nutrient resources in mangroves of southwestern Everglades. The gradient in total P deposition to mangrove soils from west to east direction across the FCE associated with this storm event is particularly significant to forest development due to the P-limited condition of this carbonate ecosystem. This source of P may be an important adaptation of mangrove forests in the Caribbean region to projected impacts of sea-level rise.

Patterns of Root Dynamics in Mangrove Forests Along Environmental Gradients in the Florida Coastal Everglades, USA

Patterns of mangrove vegetation in two distinct basins of Florida Coastal Everglades (FCE), Shark River estuary and Taylor River Slough, represent unique opportunities to test hypotheses that root dynamics respond to gradients of resources, regulators, and hydroperiod. We propose that soil total phosphorus (P) gradients in these two coastal basins of FCE cause specific patterns in belowground biomass allocation and net primary productivity that facilitate nutrient acquisition, but also minimize stress from regulators and hydroperiod in flooded soil conditions. Shark River basin has higher P and tidal hydrology with riverine mangroves, in contrast to scrub mangroves of Taylor basin with more permanent flooding and lower P across the coastal landscape. Belowground biomass (0–90 cm) of mangrove sites in Shark River and Taylor River basins ranged from 2317 to 4673 g m-2, with the highest contribution (62–85%) of roots in the shallow root zone (0–45 cm) compared to the deeper root zone (45–90 cm). Total root productivity did not vary significantly among sites and ranged from 407 to 643 g m-2 y-1. Root production in the shallow root zone accounted for 57–78% of total production. Root turnover rates ranged from 0.04 to 0.60 y-1 and consistently decreased as the root size class distribution increased from fine to coarse roots, indicating differences in root longevity. Fine root biomass was negatively correlated with soil P density and frequency of inundation, whereas fine root turnover decreased with increasing soil N:P ratios. Lower P availability in Taylor River basin relative to Shark River basin, along with higher regulator and hydroperiod stress, confirms our hypothesis that interactions of stress from resource limitation and long duration of hydroperiod account for higher fine root biomass along with lower fine root production and turnover. Because fine root production and organic matter accumulation are the primary processes controlling soil formation and accretion in scrub mangrove forests, root dynamics in the P-limited carbonate ecosystem of south Florida have a major controlling role as to how mangroves respond to future impacts of sealevel rise.

Relating precipitation and water management to nutrient concentrations in the oligotrophic ‘‘upside-down’’ estuaries of the Florida Everglades

We present 8 yr of long-term water quality, climatological, and water management data for 17 locations in Everglades National Park, Florida. Total phosphorus (P) concentration data from freshwater sites (typically ,0.25 mmol L21, or 8 mg L21) indicate the oligotrophic, P-limited nature of this large freshwater–estuarine landscape. Total P concentrations at estuarine sites near the Gulf of Mexico (average ø0.5 m mol L21) demonstrate the marine source for this limiting nutrient. This ‘‘upside down’’ phenomenon, with the limiting nutrient supplied by the ocean and not the land, is a defining characteristic of the Everglade landscape. We present a conceptual model of how the seasonality of precipitation and the management of canal water inputs control the marine P supply, and we hypothesize that seasonal variability in water residence time controls water quality through internal biogeochemical processing. Low freshwater inflows during the dry season increase estuarine residence times, enabling local processes to control nutrient availability and water quality. El Nin˜o–Southern Oscillation (ENSO) events tend to mute the seasonality of rainfall without altering total annual precipitation inputs. The Nin˜o3 ENSO index (which indicates an ENSO event when positive and a La Nin˜a event when negative) was positively correlated with both annual rainfall and the ratio of dry season to wet season precipitation. This ENSO-driven disruption in seasonal rainfall patterns affected salinity patterns and tended to reduce marine inputs of P to Everglades estuaries. ENSO events also decreased dry season residence times, reducing the importance of estuarine nutrient processing. The combination of variable water management activities and interannual differences in precipitation patterns has a strong influence on nutrient and salinity patterns in Everglades estuaries.

Factors affecting the concentration and flux of materials in two southern Everglades mangrove wetlands

Concentrations and fluxes of C, N, and P were measured in dwarf and fringe mangrove wetlands along the Taylor River, Florida, USA from 1996 to 1998. Data from these studies revealed considerable spatial and temporal variability. Concentrations of C, N, and P in the dwarf wetland showed seasonal trends, while water source was better at explaining concentrations in the fringe wetland. The total and dissolved organic carbon (TOC and DOC), total nitrogen (TN), and total phosphorus (TP) content of both wetlands was higher during the wet season or when water was flowing to the south (Everglades source). Concentrations of nitrate plus nitrite (NOx –), ammonium (NH4 +), and soluble reactive phosphorus (SRP) in the fringe wetland were all highest during the dry season or northerly flow (bay source). Nutrient concentrations most effectively explained patterns of flux in both wetlands. Increased wetland uptake of a given constituent was usually a function of its availability in the water column. However, the release of NOx – from the dwarf wetland was related to the NH4 + concentration, suggesting a nitrification signal. Nitrogen flux in the dwarf wetland was also related to surface water salinity and temperature. Our findings indicate that freshwater Everglades marshes are an important source of dissolved organic matter to these wetlands, while Florida Bay may be a source of dissolved inorganic nutrients. Our data also suggest that temperature, salinity, and nutrient concentrations (as driven by season and water source) influence patterns of materials flux in this mangrove wetland. Applying long-term water quality data to the relationships we extracted from these flux data, we estimated that TN and TP were imported by the dwarf wetland 87 ± 10 and 48 ± 17% of the year, respectively. With Everglades restoration, modifications in freshwater delivery may have considerable effects on the exchanges of nutrients and organic matter in these transitional mangrove wetlands.

Eddy covariance and meteorological tower in a long-hydroperiod Everglades marsh during the height of the wet season, 2008 (at SRS-2), Shark River Slough

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. This image is made available for non-commercial or educational use only.

Temporally dependent C, N, and P dynamics associated with the decay of Rhizophora mangle L. leaf litter in oligotrophic mangrove wetlands of the Southern Everglades

We performed two litter decomposition experiments using nearly-senesced red mangrove (Rhizophora mangle L.) leaves collected from an Everglades dwarf mangrove wetland to understand the short-term (3 weeks) and long-term (1 year) changes in mass, as well as C-, N-, and P-content of decomposing leaf litter. We expected that leaves decomposing in this oligotrophic environment would be short-term sources of C, N, and P, but potential long-term sinks for N and P. In May 1998, we conducted a 3-week leaching experiment, incubating fresh, individual leaves in seawater for up to 21 days. From May 1997 to May 1998, leaf litter in mesh bags decomposed on the forest floor at two dwarf mangrove sites. Leaching accounted for about 33% loss of dry mass from R. mangle leaves after 3 weeks. Leaching losses were rapid, peaking by day 2, and large, with leachate concentrations of total organic carbon (TOC) and total phosphorus (TP) increasing by more than an order of magnitude after 3 weeks. Mean leaf C:N increased from 105 to 115 and N:P increased from a mean of 74 to 95 after 21 days, reflecting the relatively large leaching losses of N and P. Loss of mass in the litterbags leveled off after 4 months, with roughly 60%dry mass remaining (DMR) after nearly 1 year of decomposition. The mass of carbon in each litterbag declined significantly after 361 days, but the mass of nitrogen and phosphorus doubled, indicating long-term accumulation of these constituents into the detritus. Subsequently, the leaf C:N ratio dropped significantly from 90 to 34 after 361 days. Following an initial 44-day increase, leaf N:P decreased from 222 to 144, reflecting high accumulation of P relative to N. A review of several estuarine macrophyte decomposition studies reveals a trend in nitrogen accumulation through time regardless of site, but suggests no clear pattern for C and P. We believe that the increase in litter P observed in this study was indicative of the P-limited status of the greater Everglades ecosystem and that decomposing mangrove litter may represent a substantial phosphorus pool in the system.

Ecological indicators for system-wide assessment of the greater everglades ecosystem restoration program

Developing scientifically credible tools for measuring the success of ecological restoration projects is a difficult and a non-trivial task. Yet, reliable measures of the general health and ecological integrity of ecosystems are critical for assessing the success of restoration programs. The South Florida Ecosystem Restoration Task Force (Task Force), which helps coordinate a multi-billion dollar multi-organizational effort between federal, state, local and tribal governments to restore the Florida Everglades, is using a small set of system-wide ecological indicators to assess the restoration efforts. A team of scientists and managers identified eleven ecological indicators from a field of several hundred through a selection process using 12 criteria to determine their applicability as part of a system-wide suite. The 12 criteria are: (1) is the indicator relevant to the ecosystem? (2) Does it respond to variability at a scale that makes it applicable to the entire system? (3) Is the indicator feasible to implement and is it measureable? (4) Is the indicator sensitive to system drivers and is it predictable? (5) Is the indicator interpretable in a common language? (6) Are there situations where an optimistic trend with regard to an indicator might suggest a pessimistic restoration trend? (7) Are there situations where a pessimistic trend with regard to an indicator may be unrelated to restoration activities? (8) Is the indicator scientifically defensible? (9) Can clear, measureable targets be established for the indicator to allow for assessments of success? (10) Does the indicator have specificity to be able to result in corrective action? (11) What level of ecosystem process or structure does the indicator address? (12) Does the indicator provide early warning signs of ecological change? In addition, a two page stoplight report card was developed to assist in communicating the complex science inherent in ecological indicators in a common language for resource managers, policy makers and the public. The report card employs a universally understood stoplight symbol that uses green to indicate that targets are being met, yellow to indicate that targets have not been met and corrective action may be needed and red to represent that targets are far from being met and corrective action is required. This paper presents the scientific process and the results of the development and selection of the criteria, the indicators and the stoplight report card format and content. The detailed process and results for the individual indicators are presented in companion papers in this special issue of Ecological Indicators.

Five new species of yeasts from fresh water and marine habitats in the Florida Everglades

Yeast populations in the Shark River Slough of the Florida Everglades, USA, were examined during a 3-year period (2002–2005) at six locations ranging from fresh water marshes to marine mangroves. Seventy-four described species (33 ascomycetes and 41 basidiomycetes) and an approximately equal number of undescribed species were isolated during the course of the investigation. Serious human pathogens, such as Candida tropicalis, were not observed, which indicates that their presence in coastal waters is due to sources of pollution. Some of the observed species were widespread throughout the fresh water and marine habitats, whereas others appeared to be habitat restricted. Species occurrence ranged from prevalent to rare. Five representative unknown species were selected for formal description. The five species comprise two ascomycetes: Candida sharkiensis sp. nov. (CBS 11368T) and Candida rhizophoriensis sp. nov. (CBS 11402T) (Saccharomycetales, Metschnikowiaceae), and three basidiomycetes: Rhodotorula cladiensis sp. nov. (CBS 10878T) in the Sakaguchia clade (Cystobasidiomycetes), Rhodotorula evergladiensis sp. nov. (CBS 10880T) in the Rhodosporidium toruloides clade (Microbotryomycetes, Sporidiobolales) and Cryptococcus mangaliensis sp. nov. (CBS 10870T) in the Bulleromyces clade (Agaricomycotina, Tremellales).

Seasonal plant water uptake patterns in the saline southeast Everglades ecotone

The purpose of this study was to determine the seasonal water use patterns of dominant macrophytes coexisting in the coastal Everglades ecotone. We measured the stable isotope signatures in plant xylem water of Rhizophora mangle, Cladium jamaicense, and Sesuvium portulacastrum during the dry (DS) and wet (WS) seasons in the estuarine ecotone along Taylor River in Everglades National Park, FL, USA. Shallow soilwater and deeper groundwater salinity was also measured to extrapolate the salinity encountered by plants at their rooting zone. Average soil water oxygen isotope ratios (δ 18O) was enriched (4.8 ± 0.2‰) in the DS relative to the WS (0.0 ± 0.1‰), but groundwater δ 18O remained constant between seasons (DS: 2.2 ± 0.4‰; WS: 2.1 ± 0.1‰). There was an inversion in interstitial salinity patterns across the soil profile between seasons. In the DS, shallow water was euhaline [i.e., 43 practical salinity units (PSU)] while groundwater was less saline (18 PSU). In the WS, however, shallow water was fresh (i.e., 0 PSU) but groundwater remained brackish (14 PSU). All plants utilized 100% (shallow) freshwater during the WS, but in the DS R. mangle switched to a soil–groundwater mix (δ 55% groundwater) while C. jamaicense and S. portulacastrum continued to use euhaline shallow water. In the DS, based on δ 18O data, the roots of R. mangle roots were exposed to salinities of 25.4 ± 1.4 PSU, less saline than either C. jamaicense(39.1 ± 2.2 PSU) or S. portulacastrum (38.6 ± 2.5 PSU). Although the salinity tolerance of C. jamaicense is not known, it is unlikely that long-term exposure to high salinity is conducive to the persistence of this freshwater marsh sedge. This study increases our ecological understanding of how water uptake patterns of individual plants can contribute to ecosystem levels changes, not only in the southeast saline Everglades, but also in estuaries in general in response to global sea level rise and human-induced changes in freshwater flows.

Spatial and temporal patterns of aboveground net primary productivity (ANPP) along two freshwater-estuarine transects in the Florida Coastal Everglades

We present here a 4-year dataset (2001–2004) on the spatial and temporal patterns of aboveground net primary production (ANPP) by dominant primary producers (sawgrass, periphyton, mangroves, and seagrasses) along two transects in the oligotrophic Florida Everglades coastal landscape. The 17 sites of the Florida Coastal Everglades Long Term Ecological Research (FCE LTER) program are located along fresh-estuarine gradients in Shark River Slough (SRS) and Taylor River/C-111/Florida Bay (TS/Ph) basins that drain the western and southern Everglades, respectively. Within the SRS basin, sawgrass and periphyton ANPP did not differ significantly among sites but mangrove ANPP was highest at the site nearest the Gulf of Mexico. In the southern Everglades transect, there was a productivity peak in sawgrass and periphyton at the upper estuarine ecotone within Taylor River but no trends were observed in the C-111 Basin for either primary producer. Over the 4 years, average sawgrass ANPP in both basins ranged from 255 to 606 g m−2 year−1. Average periphyton productivity at SRS and TS/Ph was 17–68 g C m−2 year−1 and 342–10371 g C m−2 year−1, respectively. Mangrove productivity ranged from 340 g m−2 year−1 at Taylor River to 2208 g m−2 year−1 at the lower estuarine Shark River site. Average Thalassia testudinum productivity ranged from 91 to 396 g m−2 year−1 and was 4-fold greater at the site nearest the Gulf of Mexico than in eastern Florida Bay. There were no differences in periphyton productivity at Florida Bay. Interannual comparisons revealed no significant differences within each primary producer at either SRS or TS/Ph with the exception of sawgrass at SRS and the C−111 Basin. Future research will address difficulties in assessing and comparing ANPP of different primary producers along gradients as well as the significance of belowground production to the total productivity of this ecosystem.

Seasonal fish community variation in headwater mangrove creeks in the southwestern Everglades: an examination of their role as dry-down refuges

The connectivity between the fish community of estuarine mangroves and that of freshwater habitats upstream remains poorly understood. In the Florida Everglades, mangrove-lined creeks link freshwater marshes to estuarine habitats downstream and may act as dry-season refuges for freshwater fishes. We examined seasonal dynamics in the fish community of ecotonal creeks in the southwestern region of Everglades National Park, specifically Rookery Branch and the North and watson rivers. Twelve low-order creeks were sampled via electrofishing, gill nets, and minnow traps during the wet season, transition period, and dry season in 2004-2005. Catches were greater in Rookery Branch than in the North and watson rivers, particularly during the transition period. Community composition varied seasonally in Rookery Branch, and to a greater extent for the larger species, reflecting a pulse of freshwater taxa into creeks as marshes upstream dried periodically. The pulse was short-lived, a later sample showed substantial decreases in freshwater fish numbers. No evidence of a similar influx was seen in the North and watson rivers, which drain shorter hydroperiod marshes and exhibit higher salinities. These results suggest that head-water creeks can serve as important dry-season refugia. Increased freshwater flow resulting from Everglades restoration may enhance this connectivity.

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.