Carbon and Nitrogen Stable Isotopic Patterns in South Florida Coastal Ecosystems: Modern and Paleoceanographic Perspectives

Long term management plans for restoration of natural flow conditions through the Everglades increase the importance of understanding potential nutrient impacts of increased freshwater delivery on coastal biogeochemistry. The present study sought to increase understanding of the coastal marine system of South Florida under modern conditions and through the anthropogenic changes in the last century, on scales ranging from individual nutrient cycle processes to seasonal patterns in organic material (OM) under varying hydrodynamic regime, to century scale analysis of sedimentary records. In all applications, carbon and nitrogen stable isotopic compositions of OM were examined as natural recorders of change and nutrient cycling in the coastal system. High spatial and temporal variability in stable isotopic compositions were observed on all time scales. During a transient phytoplankton bloom, ä15N values suggested nitrogen fixation as a nutrient source supporting enhanced productivity. Seasonally, particulate organic material (POM) from ten sites along the Florida Reef Tract and in Florida Bay demonstrated variable fluctuations dependent on hydrodynamic setting. Three separate intra-annual patterns were observed, yet statistical differences were observed between groupings of Florida Bay and Atlantic Ocean sites. The POM ä15N values ranged on a quarterly basis by 7‰, while ä13C varied by 22‰. From a sediment history perspective, four cores collected from Florida Bay further demonstrated the spatial and temporal variability of the system in isotopic composition of bulk OM over time. Source inputs of OM varied with location, with terrestrial inputs dominating proximal to Everglades freshwater discharge, seagrasses dominating in open estuary cores, and a marine mixture of phytoplankton and seagrass in a core from the boundary zone between Florida Bay and the Gulf of Mexico. Significant shifts in OM geochemistry were observed coincident with anthropogenic events of the 20th century, including railroad and road construction in the Florida Keys and Everglades, and also the extensive drainage changes in Everglades hydrology. The sediment record also preserved evidence of the major hurricanes of the last century, with excursions in geochemical composition coincident with Category 4-5 storms.

Spatiotemporal Variation in Abundance and Social Structure of Bottlenose Dolphins in the Florida Coastal Everglades

Bottlenose dolphins (Tursiops truncatus) are large-bodied predators that are locally abundant in the coastal Everglades. Because of their potential to exert strong top-down effects on their communities, it is important to understand how spatiotemporal variation in biotic and abiotic factors affects the abundance and behavior of dolphins. This study combined two years of transect surveys with photographic identification methods to assess spatiotemporal variation in the abundance and group sizes of bottlenose dolphins across four large regions of the coastal Everglades including the Shark and Harney Rivers, Whitewater Bay, and coastal oceans of the Gulf of Mexico and Florida Bay. Dolphin abundance was similar across wet and dry seasons, except in river habitats where abundances were higher during the dry season. Group sizes were largest in Florida Bay and open water. Dolphins may be relatively resilient to abiotic changes in the coastal Everglades, with the possible exception of river habitats.

The exchange of carbon, nitrogen, and phosphorus in dwarf and fringe mangroves of the oligotrophic southern Everglades

Water management has altered both the natural timing and volume of freshwater delivered to Everglades National Park. This is especially true for Taylor Slough and the C-111 basin, as hypersaline events in Florida Bay have been linked to reduced freshwater flow in this area. In light of recent efforts to restore historical flows to the eastern Everglades, an understanding of the impact of this hydrologic shift is needed in order to predict the trajectory of restoration. I conducted a study to assess the importance of season, water chemistry, and hydrologic conditions on the exchange of nutrients in dwarf and fringe mangrove wetlands along Taylor Slough. I also performed mangrove leaf decomposition studies to determine the contribution of biotic and abiotic processes to mass loss, the effect of salinity and season on degradation rates, and the importance of this litter component as a rapid source of nutrients. ^ Dwarf mangrove wetlands consistently imported total nutrients (C, N, and P) and released NO2− +NO3 −, with enhanced release during the dry season. Ammonium flux shifted from uptake to release over the study period. Dissolved phosphate activity was difficult to discern in either wetland, as concentrations were often below detection limits. Fluxes of dissolved inorganic nitrogen in the fringe wetland were positively related to DIN concentrations. The opposite was found for total nitrogen in the fringe wetland. A dynamic budget revealed a net annual export of TN to Florida Bay that was highest during the wet season. Simulated increases and decreases in freshwater flow yielded reduced exports of TN to Florida Bay as a result of changes in subsystem and water flux characteristics. Finally, abiotic processes yielded substantial nutrient and mass losses from senesced leaves with little influence of salinity. Dwarf mangrove leaf litter appeared to be a considerable source of nutrients to the water column of this highly oligotrophic wetland. To summarize, nutrient dynamics at the subsystem level were sensitive to short-term changes in hydrologic and seasonal conditions. These findings suggest that increased freshwater flow has the potential to lead to long-term, system-level changes that may reach as far as eastern Florida Bay. ^

The role of disturbance and resource availability in structuring plant communities

Disturbances alter competitive hierarchies by reducing populations and altering resource regimes. The interaction between disturbance and resource availability may strongly influence the structure of plant communities, as observed in the recolonization of seagrass beds in outer Florida Bay that were denuded by sea-urchin overgrazing. There is no consensus concerning the interaction between disturbance and resource availability on competition intensity (CI). On the other hand, species diversity is dependent on both factors. Peaks in species diversity have been observed to occur when both resource availability and disturbance intensity are high, thus implying that CI is low. Based on this supposition of previous models, I presented the resource-disturbance hypothesis as a graphical model to make predictions of CI as a function of both disturbance intensity and the availability of a limiting resource. The predictions of this model were tested in two experiments within a seagrass community in south Florida, in which transplants of Halodule wrightii were placed into near-monocultures of Syringodium filiforme in a full-factorial array. In the first experiment, two measures of relative CI were calculated based on the changes in the short-shoot number (SS) and of rhizome length (RHL) on the transplants. Both light and disturbance were identified as important factors, though the interaction between light * disturbance was not significant. Relative CISS ranged between 0.2 and 1.0 for the high light and high disturbance treatments and the relative CIRHL < 0 for the same treatments, though results were not significantly different due to high variability and low sample size. These results, including a contour schematic using six data points from the different treatment combinations, preliminarily suggests that the resource-disturbance hypothesis may be used may be used as a next step in developing our understanding of the mechanisms involved in structuring plant communities. Furthermore, the focus of the model is on the outcome of CI, which may be a useful predictor of changes in species diversity. Further study is needed to confirm the results of this study and validate the usefulness of this model in other systems. ^

Stable isotopes reveal complex changes in trophic relationships following nutrient addition in a coastal marine ecosystem

Complex links between the top-down and bottomup forces that structure communities can be disrupted by anthropogenic alterations of natural habitats.We used relative abundance and stable isotopes to examine changes in epifaunal food webs in seagrass (Thalassia testudinum) beds following 6 months of experimental nutrient addition at two sites in Florida Bay (USA) with different ambient fertility. At a eutrophic site, nutrient addition did not strongly affect food web structure, but at a nutrient-poor site, enrichment increased the abundances of crustacean epiphyte grazers, and the diets of these grazers became more varied. Benthic grazers did not change in abundance but shifted their diet away from green macroalgae + associated epiphytes and towards an opportunistic seagrass (Halodule wrightii) that occurred only in nutrient addition treatments. Benthic predators did not change in abundance, but their diets were more varied in enriched plots. Food chain length was short and unaffected by site or nutrient treatment, but increased food web complexity in enriched plots was suggested by increasingly mixed diets. Strong bottom-up modifications of food web structure in the nutrient-limited site and the limited top-down influences of grazers on seagrass epiphyte biomass suggest that, in this system, the bottom-up role of nutrient enrichment can have substantial impacts on community structure, trophic relationships, and, ultimately, the productivity values of the ecosystem.

Long-Term Effects of Adding Nutrients to an Oligotrophic Coastal Environment

Management of ecological disturbances requires an understanding of the time scale and dynamics of community responses to disturbance events. To characterize long-term seagrass bed responses to nutrient enrichment, we established six study sites in Florida Bay, USA. In 24 plots (0.25 m2) at each site, we regularly added nitrogen (N) and phosphorus (P) in a factorial design for 7 years. Five of the six sites exhibited strong P limitation. Over the first 2 years, P enrichment increased Thalassia testudinum cover in the three most P-limited sites. After 3 years, Halodule wrightii began to colonize many of the P-addition plots, but the degree of colonization was variable among sites, possibly due to differences in the supply of viable propagules. Thalassia increased its allocation to aboveground tissue in response to P enrichment; Halodule increased in total biomass but did not appear to change its aboveground: belowground tissue allocation. Nutrient enrichment did not cause macroalgal or epiphytic overgrowth of the seagrass. Nitrogen retention in the study plots was variable but relatively low, whereas phosphorus retention was very high, often exceeding 100% of the P added as fertilizer over the course of our experiments. Phosphorus retentions exceeding 100% may have been facilitated by increases in Thalassia aboveground biomass, which promoted the settlement of suspended particulate matter containing phosphorus. Our study demonstrated that lowintensity press disturbance events such as phosphorus enrichment can initiate a slow, ramped successional process that may alter community structure over many years.

Variable responses within epiphytic and benthic microalgal communities to nutrient enrichment

We examined the spatial extent of nitrogen (N) and phosphorus (P) limitation of each of the major benthic primary producer groups in Florida Bay (seagrass, epiphytes, macroalgae, and benthic microalgae) and characterized the shifts in primary producer community composition following nutrient enrichment. We established 24 permanent 0.25-m2 study plots at each of six sites across Florida Bay and added N and P to the sediments in a factorial design for 18 mo. Tissue nutrient content of the turtlegrass Thalassia testudinum revealed a spatial pattern in P limitation, from severe limitation in the eastern bay (N:P > 96:1), moderate limitation in two intermediate sites (approximately 63:1), and balanced with N availability in the western bay (approximately 31:1). P addition increased T. testudinum cover by 50-75% and short-shoot productivity by up to 100%, but only at the severely P-limited sites. At sites with an ambient N:P ratio suggesting moderate P limitation, few seagrass responses to nutrients occurred. Where ambient T. testudinum tissue N:P ratios indicated N and P availability was balanced, seagrass was not affected by nutrient addition but was strongly influenced by disturbance (currents, erosion). Macroalgal and epiphytic and benthic microalgal biomass were variable between sites and treatments. In general, there was no algal overgrowth of the seagrass in enriched conditions, possibly due to the strength of seasonal influences on algal biomass or regulation by grazers. N addition had little effect on any benthic primary producers throughout the bay. The Florida Bay benthic primary producer community was P limited, but P-induced alterations of community structure were not uniform among primary producers or across Florida Bay and did not always agree with expected patterns of nutrient limitation based on stoichiometric predictions from field assays of T. testudinum tissue N:P ratios.

Red mangroves emit hydrocarbons

The objective of this study is to investigate hydrocarbon species and amounts released by red mangrove foliage and determine if these quantities warrant future research on atmospheric chemical processing of these compounds. The field investigation took place during July 2001 at Key Largo, Florida Bay, Florida. Foliage still attached to plants was enclosed in cuvettes while air of known flow rates circulated around leaves to study, hydrocarbon emissions. Cuvette air samples underwent gas chromatographic analyses to determine species and amounts of hydrocarbons released by mangrove foliage. Red mangrove foliage emits isoprene and trace amounts of the monoterpenes of alpha-pinene, beta-pinene, camphene, and d-limonene. The mangrove flowers released these latter compounds in amounts ranging from 0.5 to 10 mg (monoterpene) per gram of dry biomass per hour. These fluxes are normalized to, the foliage temperature of 30 degreesC. When normalized to the foliage temperature of 30 degreesC and light levels of 1000 mumol m(-2) s(-1), isoprene emission rates as high as 0.092 +/- 0.109 mug (isoprene) per gram of dry biomass per hour were measured. Compared to terrestrial forest ecosystems, red mangroves are low isoprene emitters. During peak flowering periods in the summertime, however, red mangroves may emit sufficient amounts of monoterpenes to alter ground-level ozone concentrations and contribute to biogenic aerosol formation.

Nutrient enrichment, grazer identity, and their effects on epiphytic algal assemblages: field experiments in subtropical turtlegrass Thalassia testudinum meadows

We tested the relative importance of top-down and bottom-up effects by experimentally evaluating the combined and separate effects of nutrient availability and grazer species composition on epiphyte communities and seagrass condition in Florida Bay. Although we succeeded in substantially enriching our experimental cylinders, as indicated by elevated nitrogen concentrations in epiphytes and seagrass leaves, we did not observe any major increases in epiphyte biomass or major loss of Thalassia testudinum by algal overgrowth. Additionally, we did not detect any strong grazer effects and found very few significant nutrient-grazer interactions. While this might suggest that there was no important differential response to nutrients by individual grazer species or by various combinations of grazers, our results were complicated by the lack of significant differences between control and grazer treatments, and as such, these results are best explained by the presence of unwanted amphipod grazers (mean = 471 ind. m–2) in the control cylinders. Our estimates of grazing rates and epiphyte productivities indicate that amphipods in the control cylinders could have lowered epiphyte biomass to the same level that the experimental grazers did, thus effectively transforming the control treatments into grazer treatments. If so, our experiments suggest that the effects of invertebrate grazing (and those of amphipods alone) were stronger than the effects of nutrient enrichment on epiphytic algae, and that it does not require a large density

The potential role of plant oxygen and sulphide dynamics in die-off events of the tropical seagrass, Thalassia testudinum

1 Oxygen and sulphide dynamics were examined, using microelectrode techniques, in meristems and rhizomes of the seagrass Thalassia testudinum at three different sites in Florida Bay, and in the laboratory, to evaluate the potential role of internal oxygen variability and sulphide invasion in episodes of sudden die-off. The sites differed with respect to shoot density and sediment composition, with an active die-off occurring at only one of the sites. 2 Meristematic oxygen content followed similar diel patterns at all sites with high oxygen content during the day and hyposaturation relative to the water column during the night. Minimum meristematic oxygen content was recorded around sunrise and varied among sites, with values close to zero at the die-off site. 3 Gaseous sulphide was detected within the sediment at all sites but at different concentrations among sites and within the die-off site. Spontaneous invasion of sulphide into Thalassia rhizomes was recorded at low internal oxygen partial pressure during darkness at the die-off site. 4 A laboratory experiment showed that the internal oxygen dynamics depended on light availability, and hence plant photosynthesis, and on the oxygen content of the water column controlling passive oxygen diffusion from water column to leaves and belowground tissues in the dark. 5 Sulphide invasion only occurred at low internal oxygen content, and the rate of invasion was highly dependent on the oxygen supply to roots and rhizomes. Sulphide was slowly depleted from the tissues when high oxygen partial pressures were re-established through leaf photosynthesis. Coexistence of sulphide and oxygen in the tissues and the slow rate of sulphide depletion suggest that sulphide reoxidation is not biologically mediated within the tissues of Thalassia. 6 Our results support the hypothesis that internal oxygen stress, caused by low water column oxygen content or poor plant performance governed by other environmental factors, allows invasion of sulphide and that the internal plant oxygen and sulphide dynamics potentially are key factors in the episodes of sudden die-off in beds of Thalassia testudinum . Root anoxia followed by sulphide invasion may be a more general mechanism determining the growth and survival of other rooted plants in sulphate-rich aquatic environments.

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.

Negative relationships between the nutrient and carbohydrate content of the seagrass Thalassia testudinum

This study documents relationships between plant nutrient content and rhizome carbohydrate content of a widely distributed seagrass species, Thalassia testudinum, in Florida. Five distinct seagrass beds were sampled for leaf nitrogen, leaf phosphorus, and rhizome carbohydrate content from 1997 to 1999. All variables displayed marked intra- and inter- regional variation. Elemental ratios (mean N:P ± S.E.) were lowest for Charlotte Harbor (9.9 ± 0.2) and highest for Florida Bay (53.5 ± 0.9), indicating regional shifts in the nutrient content of plant material. Rhizome carbohydrate content (mean ± S.E.) was lowest for Anclote Keys (21.8 ± 1.6 mg g−1 FM), and highest for Homosassa Bay (40.7 ± 1.7 mg g−1 FM). Within each region, significant negative correlations between plant nutrient and rhizome carbohydrate content were detected; thus, nutrient-replete plants displayed low carbohydrate content, while nutrient-deplete plants displayed high carbohydrate content. Spearman's rank correlations between nutrient and carbohydrate content varied from a minimum in Tampa Bay (ρ = −0.2) to a maximum in Charlotte Harbor (ρ = −0.73). Linear regressions on log-transformed data revealed similar trends. This consistent trend across five distinct regions suggests that nutrient supply may play an important role in the regulation of carbon storage within seagrasses. Here we present a new hypothesis for studies which aim to explain the carbohydrate dynamics of benthic plants.

Biogeochemical effects of iron availability on primary producers in a shallow marine carbonate environment

We completed a synoptic survey of iron, phosphorus, and sulfur concentrations in shallow marine carbonate sediments from south Florida. Total extracted iron concentrations typically were 50 μmol g-1 dry weight (DW) and tended to decrease away from the Florida mainland, whereas total extracted phosphorus concentrations mostly were 10 μmol g-1 DW and tended to decrease from west to east across Florida Bay. Concentrations of reduced sulfur compounds, up to 40 μmol g-1 DW, tended to covary with sediment iron concentrations, suggesting that sulfide mineral formation was iron-limited. An index of iron availability derived from sediment data was negatively correlated with chlorophyll a concentrations in surface waters, demonstrating the close coupling of sediment-water column processes. Eight months after applying a surface layer of iron oxide granules to experimental plots, sediment iron, phosphorus, and sulfur were elevated to a depth of 10 cm relative to control plots. Biomass of the seagrass Thalassia testudinum was not different between control and iron addition plots, but individual shoot growth rates were significantly higher in experimental plots after 8 months. Although the iron content of leaf tissues was significantly higher from iron addition plots, no difference in phosphorus content of T. testudinum leaves was observed. Iron addition altered plant exposure to free sulfide, documented by a significantly higher δ34S of leaf tissue from experimental plots relative to controls. Iron as a buffer to toxic sulfides may promote individual shoot growth, but phosphorus availability to plants still appears to limit production in carbonate sediments.