Interspecific variation in the elemental and stable isotope content of seagrasses in South Florida

The elemental (C, N, and P) and isotope (δ13C, δ15N) content of leaves of the seagrasses Thalassia testudinum, Halodule wrightii, and Syringodium filiforme were measured across a 10 000 km2 survey of the seagrass communities of South Florida, USA, in 1999 and 2000. Trends at local and broad spatial scales were compared to examine interspecific variation in the seagrass characteristics often used as ecological indicators. The elemental and stable isotope contents of all species were variable and demonstrated marked interspecific variation. At broad spatial scales, mean N:P ratios were lowest for T. testudinum (36.5 ± 1.1) and S. filiforme (38.9 ± 1.3), and highest for H. wrightii (44.1 ± 1.8). Stable carbon isotope ratios (δ13C) were highest for S. filiforme (–6.2 ± 0.2‰), intermediate for T. testudinum (–8.6 ± 0.2‰), and lowest for H. wrightii (–10.6 ± 0.3‰). Stable nitrogen isotopes (δ15N) were heaviest for T. testudinum (2.0 ± 0.1‰), and lightest for H. wrightii (1.0 ± 0.3‰) and S. filiforme (1.6 ± 0.2‰). Site depth was negatively correlated to δ13C for all species, while δ15N was positively correlated to depth for H. wrightii and S. filiforme. Similar trends were observed in local comparisons, suggesting that taxon-specific physiological/ecological properties strongly control interspecific variation in elemental and stable isotope content. Temporal trends in δ13C were measured, and revealed that interspecific variation was displayed throughout the year. This work documents interspecific variation in the nutrient dynamics of 3 common seagrasses in South Florida, indicating that interpretation of elemental and stable isotope values needs to be species specific

Novel methodology for in situ carbon dioxide enrichment of benthic ecosystems

Future climate change will likely represent a major stress to shallow aquatic and coastal marine communities around the world. Most climate change research, particularly in regards to increased pCO2 and ocean acidification, relies on ex situ mesocosm experimentation, isolating target organisms from their environment. Such mesocosms allow for greater experimental control of some variables, but can often cause unrealistic changes in a variety of environmental factors, leading to “bottle effects.” Here we present an in situ technique of altering dissolved pCO2within nearshore benthic communities (e.g., macrophytes, algae, and/or corals) using submerged clear, open-top chambers. Our technique utilizes a flow-through design that replicates natural water flow conditions and minimizes caging effects. The clear, open-top design additionally ensures that adequate light reaches the benthic community. Our results show that CO2 concentrations and pH can be successfully manipulated for long durations within the open-top chambers, continuously replicating forecasts for the year 2100. Enriched chambers displayed an average 0.46 unit reduction in pH as compared with ambient chambers over a 6-month period. Additionally, CO2 and HCO3 – concentrations were all significantly higher within the enriched chambers. We discuss the advantages and disadvantages of this technique in comparison to other ex situ mesocosm designs used for climate change research

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

Mechanisms of Bicarbonate Use Influence the Photosynthetic Carbon Dioxide Sensitivity of Tropical Seagrasses

The photosynthetic bicarbonate () use properties of three widely distributed tropical seagrasses were compared using a series of laboratory experiments. Photosynthetic rates of Thalassia testudinum, Halodule wrightii, and Syringodium filiforme were monitored in an enclosed chamber while being subjected to shifts in pH and dissolved inorganic carbon. Specific mechanisms of seagrass use were compared by examining the photosynthetic effects of the carbonic anhydrase inhibitor acetazolamide (AZ). All seagrasses increased photosynthetic rates with reduced pH, suggesting a large effect of dissolved aqueous carbon dioxide (CO2(aq)). However, there was considerable interspecific variation in pH response. T. testudinum was highly sensitive, increasing photosynthetic rates by 100% as the pH was reduced from 8.2 to 7.4, whereas rates in H. wrightii and S. filiforme increased by only 20% over a similar range, and displayed prominent photosynthetic plateaus, indicating an increased capacity for use. Additional incubations that manipulated [] under constant [CO2(aq)] support these findings, as only H. wrightii and S. filiforme increased photosynthetic rates with increasing []. T. testudinum responded to AZ addition, indicating that carbonic anhydrase enzymes facilitate limited use. H. wrightii and S. filiforme showed no response to AZ, suggesting alternate, more efficient mechanisms of use. Estimated kinetic parameters, Ks(CO2) and Vmax, revealed interspecific variation and further support these conclusions. Variation in photosynthetic pH responses and AZ sensitivity indicate distinctions in the carbon use properties of seagrasses exposed to similar environmental conditions. These results suggest that not all seagrasses will similarly respond to future increases in CO2(aq) availability. Attention towards potential shifts in competitive interactions within multispecific seagrass beds is warranted

Modeled CO2 Emissions from Coastal Wetland Transitions to Other Land Uses: Tidal Marshes, Mangrove Forests, and Seagrass Beds

The sediments of coastal wetlands contain large stores of carbon which are vulnerable to oxidation once disturbed, resulting in high levels of CO2 emissions that may be avoided if coastal ecosystems are conserved or restored. We used a simple model to estimate CO2 emissions from mangrove forests, seagrass beds, and tidal marshes based on known decomposition rates for organic matter in these ecosystems under either oxic or anoxic conditions combined with assumptions of the proportion of sediment carbon being deposited in either oxic or anoxic environments following a disturbance of the habitat. Our model found that over 40 years after disturbance the cumulative CO2 emitted from tidal marshes, mangrove forests, and seagrass beds were ∼70–80% of the initial carbon stocks in the top meter of the sediment. Comparison of our estimates of CO2 emissions with empirical studies suggests that (1) assuming 50% of organic material moves to an oxic environment after disturbance gives rise to estimates that are similar to CO2 emissions reported for tidal marshes; (2) field measurements of CO2 emissions in disturbed mangrove forests were generally higher than our modeled emissions that assumed 50% of organic matter was deposited in oxic conditions, suggesting higher proportions of organic matter may be exposed to oxic conditions after disturbance in mangrove ecosystems; and (3) the generally low observed rates of CO2 emissions from disturbed seagrasses compared to our estimates, assuming removal of 50% of the organic matter to oxic environments, suggests that lower proportions may be exposed to oxic conditions in seagrass ecosystems. There are significant gaps in our knowledge of the fate of wetland sediment carbon in the marine environment after disturbance. Greater knowledge of the distribution, form, decomposition, and emission rates of wetland sediment carbon after disturbance would help to improve models

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

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\u27s 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

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 \u3e 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

Light Attenuation in Estuarine Mangrove Lakes

Submerged aquatic vegetation (SAV) cover has declined in brackish lakes in the southern Everglades characterized by low water transparencies, emphasizing the need to evaluate the suitability of the aquatic medium for SAV growth and to identify the light attenuating components that contribute most to light attenuation. Underwater attenuation of downwards irradiance of photosynthetically active radiation (PAR) was determined over a three year period at 42 sites in shallow (\u3c2 m depth) mangrove-surrounded lakes in two sub-estuaries in the coastal Everglades, Florida USA. Turbidity, chromophoric dissolved organic matter (CDOM), and phytoplankton chlorophyll a (chl a) were measured concurrently and their respective contributions to the light attenuation rate were estimated. Light transmission to the benthos relative to literature estimates of minimum requirements for SAV growth indicated that the underwater light environment was often unsuitable for SAV. Light attenuation rates (n = 417) corrected for solar elevation angles ranged from 0.16 m-1 to 9.83 m-1 with a mean of 1.73 m-1. High concentrations of CDOM with high specific light absorption contributed the most to light attenuation followed by turbidity and chl a. CDOM alone sufficiently reduces light transmission beyond the estimated limits for SAV growth, making it difficult for ecosystem managers to increase SAV abundance by management activities. Light limitation of SAV in these areas may be a persistent feature because of their proximity to CDOM source materials from the surrounding mangrove swamp. Increasing freshwater flow into these areas may dilute CDOM concentrations and improve the salinity and light climate for SAV communities

Effect of Seagrass on Current Speed: Importance of Flexibility vs. Shoot Density

Water flow through seagrass beds transports nutrients, affects sediment stability and chemistry, and imposes hydrodynamic forces on shoots that alter canopy configuration. Past studies done under diverse conditions yielded conflicting results about the effects of shoot density on flow through seagrass bed canopies. We used eelgrass, Zostera marina, to study how the density of flexible shoots affect the hydrodynamics of seagrass beds in unidirectional water flow. By exposing randomly-arranged shoots of uniform length to current velocities controlled in a flume, the effects of shoot density and distance downstream from the bed edge could be determined without confounding factors. Comparison of velocity profiles within beds to those upstream of beds showed that flow was slower in the beds. However, shoot density, downstream distance, and current velocity did not affect the percent reduction in flow velocity in a bed. Turbulence enhances mixing of substances carried in the water. Here, turbulence intensity (index of the importance of turbulent velocity fluctuations relative to average current velocity) was lower when ambient flow was faster, but was not affected by shoot density or downstream position, Drag (hydrodynamic force on a shoot that bends it over in the flow direction) provides another measure of how the canopy affects flow experienced by a shoot. Drag was not affected by current velocity, shoot density, or downstream position in the bed. Gaps between shoots can enhance light and flow penetration into the canopy, but when shoots are bent over by flow, they can cover gaps. Faster ambient currents caused greater gap closure, which at each current speed was greater for high shoot densities. Thus, canopy gap closure did not correlate with percent flow reduction in grass beds or with drag on individual shoots, both of which were independent of shoot density and ambient current velocity. Since changing shoot density does not affect the flow in a grass bed exposed to a given ambient current, our results are inconsistent with the hypothesis that the high shoot densities observed in grass beds in habitats exposed to rapid flow are due to a direct, adaptive response of the grass to the flow environment