Iron Additions Reduce Sulfate Reduction Rates and Improve Seagrass Growth on Organic-Enriched Carbonate Sediments

Here we demonstrate, through experimental iron additions to a Mediterranean seagrass meadow, that iron plays a pivotal role in seagrass systems on carbonate sediments, directly through its role as a limiting nutrient, and indirectly by stimulating phosphorus recycling through the activity of the enzyme alkaline phosphatase and by buffering the development of reduced conditions in sediments. Iron additions were performed throughout the active root zone (30 cm depth) to two Posidonia oceanica meadows, one on organic-enriched sediments and one on organic poor sediments (Reference). Seagrass growth, nutrient incorporation and sediment biogeochemical conditions were followed for four months. Iron additions had positive effects on seagrass growth (leaf production increased with 55%) and nutrient incorporation (increased 46–91%) in the organic-enriched site, increasing to levels found at the Reference site. There was no effect of iron additions in the Reference seagrass meadow suggesting that iron was not the most important controlling factor at this site. The iron pools were about two times higher compared to the organic-enriched site. The main effect on the sediment biogeochemical conditions at the organic-enriched site was a suppression of sulfate reduction activity to the levels encountered at the Reference site (6.7 mmol m⁻²d⁻¹ vs. 4.7–5.9 mmol m⁻²d⁻¹). This suggests that the sulfide stress on the seagrasses was removed and that the iron availability increased due to reduced precipitation of iron-sulfides and thus improving seagrass growth conditions in these organic-enriched sediments.     

The effects of manipulation of sedimentary iron and organic matter on sediment biogeochemistry and seagrasses in a subtropical carbonate environment

The microbial metabolism of organic matter (OM) in seagrass beds can create sulfidic conditions detrimental to seagrass growth; iron (Fe) potentially has ameliorating effects through titration of the sulfides and the precipitation of iron-sulfide minerals into the sediment. In this study, the biogeochemical effects of Fe availability and its interplay with sulfur and OM on sulfide toxicity, phosphorous (P) availability, seagrass growth and community structure were tested. The availability of Fe and OM was manipulated in a 2 × 2 factorial experiment arranged in a Latin square, with four replicates per treatment. The treatments included the addition of Fe, the addition of OM, the addition of both Fe and OM as well as no addition. The experiment was conducted in an oligotrophic, iron-deficient seagrass bed. Fe had an 84.5% retention efficiency in the sediments with the concentration of Fe increasing in the seagrass leaves over the course of the experiment. Porewater chemistry was significantly altered with a dramatic decrease in sulfide levels in Fe addition plots while sulfide levels increased in the OM addition treatments. Phosphorus increased in seagrass leaves collected in the Fe addition plots. Decreased sulfide stress was evidenced by heavier δ³⁴S in leaves and rhizomes from plots to which Fe was added. The OM addition negatively affected seagrass growth but increased P availability; the reduced sulfide stress in Fe added plots resulted in elevated productivity. Fe availability may be an important determinant of the impact that OM has on seagrass vitality in carbonate sediments vegetated with seagrasses.     

Changes in biomass and spatial distribution of <Emphasis Type="Italic">Elodea nuttallii</Emphasis> (Planch.) St. John,an invasive submerged plant,in oligomesotrophic Lake Kizaki from 1999 to 2002

Distribution of pure Elodea nuttallii vegetation was surveyed from 1999 to 2002, immediately after the most recent expansion of the species in Lake Kizaki, Japan. During 2001 and 2002, areas of E. nuttallii vegetation rapidly diminished and the summer plant height decreased wherever the vegetation remained. The organic matter content, total phosphorus, and extracted P of the sediment from the vegetation bed were measured. A linear relationship was observed between the extracted P in the sediment and the biomass. The extracted P significantly decreased in the shallow littoral vegetation bed, where the biomass clearly diminished. A fertilization experiment using the shallow littoral sediment collected in the vegetation bed was conducted in 2001. In this experiment, apical shoots of E. nuttallii were planted in pots with fertilized sediment (nitrogen, phosphorus, and potassium additions). The growth of E. nuttallii shoots was significantly enhanced by enrichment with phosphorus alone. The ecological implication of sediment phosphorus limitation is discussed in relation to the cause of decline in the E. nuttallii population in Lake Kizaki.     

Seasonal oscillation of microbial iron and sulfate reduction in saltmarsh sediments (Sapelo Island, GA, USA)

Seasonal variations in anaerobic respiration pathways were investigated at three saltmarsh sites using chemical data, sulfate reduction rate measurements, enumerations of culturable populations of anaerobic iron-reducing bacteria (FeRB), and quantification of in situ 16S rRNA hybridization signals targeted for sulfate-reducing bacteria (SRB). Bacterial sulfate reduction in the sediments followed seasonal changes in temperature and primary production of the saltmarsh, with activity levels lowest in winter and highest in summer. In contrast, a dramatic decrease in the FeRB population size was observed during summer at all sites. The collapse of FeRB populations during summer was ascribed to high rates of sulfide production by SRB, resulting in abiotic reduction of bioavailable Fe(III) (hydr)oxides. To test this hypothesis, sediment slurry incubations at 10, 20 and 30 °C were carried out. Increases in temperature and labile organic carbon availability (acetate or lactate additions) increased rates of sulfate reduction while decreasing the abundance of culturable anaerobic FeRB. These trends were not reversed by the addition of amorphous Fe(III) (hydr)oxides to the slurries. However, when sulfate reduction was inhibited by molybdate, no decline in FeRB growth was observed with increasing temperature. Addition of dissolved sulfide adversely impacted propagation of FeRB whether molybdate was added or not. Both field and laboratory data therefore support a sulfide-mediated limitation of microbial iron respiration by SRB. When total sediment respiration rates reach their highest levels during summer, SRB force a decline in the FeRB populations. As sulfate reduction activity slows down after the summer, the FeRB are able to recover.     

Modification of animal habitat by large plants: mechanisms by which seagrasses influence clam growth

Summary Field experiments withMercenaria mercenaria in a relatively high-energy environment demonstrated that clams on unvegetated sand flats failed to grow during autumn while those within seagrass beds grew substantially. Clam growth rates at the seagrass margin that first receives the faster-flowing, flood-tidal currents were about 25% less than at the opposite edge. In a second experiment, pruning, which reduced average blade length by 50–75%, was shown to enhance near-bottom current velocities and to reduce shell growth ofMercenaria during summer by about 50%. As in the first experiment, clams in the unvegetated sand flats exhibited no net growth. Clam mortality, caused mostly by predatory crabs and whelks, was much higher on sand flats than in seagrass beds and intermediate in clipped seagrass. Although consistent with some previous reports, these growth results are still surprising given that they contradict the generalization that suspension feeders grow faster under more rapid current regimes.Three types of indirect interactions might explain the observed effect of seagrass on growth of buried clams: (1) altering food supply; (2) changing the intensity of biological disturbance on feeding clams; and/or (3) affecting the physical stability of the sediments. Previous research on this question has focused almost exclusively on processes that alter food supply rates. In this study, food concentrations, as indicated by suspended chla, were 30% higher inside than outside one seagrass bed, whereas chla concentrations in two other beds were not different from those on adjacent sand flats. This result is sufficient to show that more intense food depletion was not induced by the reduction in flow velocities under the seagrass canopy. Nevertheless, the possible small difference in food concentrations between vegetated and unvegetated bottom seems insufficient to explain the absence of growth of sand-flat clams, especially given the virtual lack of food limitation among suspension feeders in this system. Two data sets demonstrated that the effects of biological disturbance agents cannot be ignored. An outdoor laboratory experiment showed that even in the absence of physical contact between predator and prey the presence of a whelk reduces the amount of time spent feeding byMercenaria. This result suggests that sand flats, where predation rates are higher, may be sites of lower clam growth than seagrass beds because of greater consumer interference with clam feeding. Furthermore, clam siphons are proportionately larger inside seagrass than on sand flats, implying that siphon nipping may not be as intense inside seagrass. This process, too, would reduce net growth of sand-flat clams. Finally, no explicit test was conducted of the hypothesis that enhanced sediment transport in the absence of flow baffling and root binding by seagrass inhibits net growth of clams on high-energy sand flats. Nevertheless, this is a reasonable explanation for the pattern of enhanced growth of seagrass clams, and could serve to explain the otherwise unexplained pattern of lower clam growth at the edge of the seagrass bed that experiences the faster flood-tidal current velocities. Each broad process, changing fluid dynamics, altering consumer access, and varying sediment stability, represents a mechanism whereby habitat structure, provided by the dominant plant, has an important indirect influence on the functional value of the habitat for resident animals.     

DIFFERENTIAL EFFECTS OF SEDIMENTS ON SURVIVAL AND GROWTH OF FUCUS SERRATUS EMBRYOS (FUCALES,PHAEOPHYCEAE)1

Recruitment of seaweeds through small reproductive stages is limited on sediment inundated rocky shores and largely unsuccessful in soft sediment environments. Burial in sediment has several potentially negative effects for seaweed propagules, and these effects were differentiated in a laboratory experiment. We investigated how light deprivation, sediment type (grain size, organic content, and origin), and sediment chemistry (oxygen presence and toxicity through hydrogen sulfide) affected survivorship and growth of Fucus serratus L. embryos. Presence of hydrogen sulfide had overriding negative impacts on both survivorship and growth of Fucus embryos, independently of sediment type and light availability. In contrast, simple anaerobiosis generally did not impair survival or growth of the embryos. Fine sediments, 3 mm thick, significantly reduced embryo survivorship, presumably through accumulation of metabolic waste products in the immediate vicinity of the embryos as a consequence of constrained diffusion. This effect was equally pronounced in the presence of a 1‐mm layer of organically rich biodeposits. Irradiance levels did not affect survival of embryos but influenced growth. Decreasing thickness and increasing coarseness of sediments together represented a gradient of enhanced light penetration and diffusion. Growth of embryos increased along this gradient. In nature, soft sediment environments with organically enriched muds (e.g. tidal flats and salt marshes) represent habitats least favorable for colonization through small reproductive stages of seaweeds.     

Influence of external zinc and phosphorus supply on Cd uptake by rice (Oryza sativa L.) seedlings with root surface iron plaque

Rice seedlings were grown in hydroponic culture to determine the effects of external Zn and P supply on plant uptake of Cd in the presence or absence of iron plaque on the root surfaces. Iron plaque was induced by supplying 50 mg l⁻¹ Fe²⁺ in the nutrient solution for 2 day. Then 43-day-old seedlings were exposed to 10 μmol l⁻¹ Cd together with 10 μmol l⁻¹ Zn or without Zn (Zn–Cd experiment), or to 10 μmol l⁻¹ Cd with 1.0 mmol l⁻¹ P or without P (P–Cd experiment) for another 2 day. The seedlings were then harvested and the concentrations of Fe, Zn, P and Cd in dithionite–citrate–bicarbonate (DCB) extracts and in roots and shoots were determined. The dry weights of roots and shoots of seedlings treated with 50 mg l⁻¹ Fe were significantly lower than when no Fe was supplied. Adsorption of Cd, Zn and P on the iron plaque increased when Fe was supplied but Cd concentrations in DCB extracts were unaffected by external Zn or P supply levels. Cd concentrations in shoots and roots were lower when Fe was supplied. Zn additions decreased Cd concentrations in roots but increased Cd concentrations in shoots, whereas P additions significantly increased shoot and root Cd concentrations and this effect diminished when Fe was supplied. The percentage of Cd in DCB extracts was significantly lower than in roots or shoots, accounting for up to 1.8–3.8% of the plant total Cd, while root and shoot Cd were within the ranges 57–76% and 21–40% respectively in the two experiments. Thus, the main barrier to Cd uptake seemed to be the root tissue and the contribution of iron plaque on root surfaces to plant Cd uptake was minor. The changes in plant Cd uptake were not due to Zn or P additions altering Cd adsorption on iron plaque, but more likely because Zn or P interfered with Cd uptake by the roots and translocation to the shoots.     

Diurnal changes in pore water sulfide concentrations in the seagrass Thalassia testudinum beds: the effects of seagrasses on sulfide dynamics

The dynamics of the seagrass-sulfide interaction were examined in relation to diel changes in sediment pore water sulfide concentrations in Thalassia testudinum beds and adjacent bare areas in Corpus Christi Bay and lower Laguna Madre, Texas, USA, during July 1996. Pore water sulfide concentrations in seagrass beds were significantly higher than in adjacent bare areas and showed strong diurnal variations; levels significantly decreased during mid-day at shallow sediment depths (0-10 cm) containing high below-ground tissue biomass and surface area. In contrast, diurnal variations in sediment sulfide concentrations were absent in adjacent bare patches, and at deeper (>10 cm) sediment depths characterized by low below-ground plant biomass or when the grasses were experimentally shaded. These observations suggest that the mid-day depressions in sulfide levels are linked to the transport of photosynthetically produced oxygen to seagrass below-ground tissues that fuels sediment sulfide oxidation. Lower sulfide concentrations in bare areas are likely a result of low sulfate reduction rates due to low organic matter available for remineralization. Further, high reoxidation rates due to rapid exchange between anoxic pore water and oxic overlying water are probably stimulated in bare areas by higher current velocity on the sediment surface than in seagrass beds. The dynamics of pore water sulfides in seagrass beds suggest no toxic sulfide intrusion into below-ground tissues during photosynthetic periods and demonstrate that the sediment chemical environment is considerably modified by seagrasses. The reduced sediment sulfide levels in seagrass beds during photosynthetic periods will enhance seagrass production through reduced sulfide toxicity to seagrasses and sediment microorganisms related to the nutrient cycling.     

Assessing the effect of genetic diversity on the early establishment of the threatened seagrass Posidonia australis using a reciprocal‐transplant experiment

Two common goals for restoration are rapid plant establishment and long‐term plant persistence. The success of transplanted populations may be jeopardized if the donor transplants are not genetically diverse, and/or poorly matched to their new environment. Here, we test the effects of local adaptation and plot‐level genetic diversity on the early establishment phase of a threatened seagrass species, Posidonia australis, by performing a reciprocal transplant experiment across two genetically and geographically distinct populations in southeastern Australia. Posidonia australis is a long‐lived, slow‐growing species that has no seed bank, and the successful transplantation of live shoots and seedlings is the only available restoration method. Our results show a strong effect of local adaptation and genetic diversity on P. australis survivorship and performance over the first 6 months following transplantation. High‐genetic diversity plots displayed higher survival rates and exhibited reduced productivity and increased carbohydrate reserves within the rhizome. This suggests that high‐diversity plots included shoots that were conserving energy stores by actively reducing growth rates during the early stages of transplantation. The lowest diversity plots exhibited high leaf and root productivity and corresponding low carbohydrate reserves. This may be a sign of stress in the low‐diversity transplants, potentially explaining the very low survival rate. We suggest that future restoration efforts source donor transplants from multiple local sources to ensure both local adaptation and sufficient genetic diversity to increase the likelihood of early establishment success.     

Positive Feedbacks in Seagrass Ecosystems: Implications for Success in Conservation and Restoration

Abstract Seagrasses are threatened by human activity in many locations around the world. Their decline is often characterized by sudden ecosystem collapse from a vegetated to a bare state. In the 1930s, such a dramatic event happened in the Dutch Wadden Sea. Before the shift, large seagrass beds (Zostera marina) were present in this area. After the construction of a large dam and an incidence of the “wasting disease” in the early 1930s, these meadows became virtually extinct and never recovered despite restoration attempts. We investigated whether this shift could be explained as a critical transition between alternative stable states, and whether the lack of recovery could be due to the high resilience of the new turbid state. We analyzed the depth distribution of the historical meadows, a long-term dataset of key factors determining turbidity and a minimal model based on these data. Results demonstrate that recovery was impossible because turbidity related to suspended sediment was too high, probably because turbidity was no longer reduced by seagrass itself. Model simulations on the positive feedback suggest indeed the robust occurrence of alternative stable states and a high resilience of the current turbid state. As positive feedbacks are common in seagrasses, our findings may explain both the worldwide observed collapses and the low success rate of restoration attempts of seagrass habitats. Therefore, appreciation of ecosystem resilience may be crucial in seagrass ecosystem management.     

Seagrass beds and intertidal invertebrates: an experimental test of the role of habitat structure

The majority of field experiments have been carried out on relatively small spatial and short temporal scales, but some of the most interesting ecological processes operate at much larger scales. However, large-scale experiments appropriate to the landscape, often have to be carried out with minimal plot replication and hence reduced statistical power. Here, we report the results of such a large scale, un-replicated field experiment on the Mondego estuary, Portugal, which nevertheless provides compelling evidence of the importance of habitat structure for invertebrate community composition and dynamics. In this estuary, seagrass beds have suffered a dramatic decline over the last 20 years, associated with changes in invertebrate assemblages. In addition, the most abundant species in the system, Hydrobia ulvae, displays distinctly different population structures in those sites. The aim of the field experiment was to test the hypothesis that these differences are related to enhanced survival of snails due to protection from avian or fish predators so that they can grow to larger body sizes in the more complex habitat provided by seagrass. We tested this hypothesis through a large-scale experiment using artificial seagrass beds over a 12-month period. Adult snail densities were higher in the artificial bed plots compared to controls. However, these differences emerged only slowly, related to snail growth rate. This suggests that protection from epibenthic predators can have a significant effect on population structure and hence biomass and productivity of key species in this system. However, the invertebrate assemblage in artificial seagrass plots and the natural seagrass bed, remained statistically separate by the end of the experiment. Handling editor: K. Martens     

Habitat-dependent growth in a Caribbean sea urchin <Emphasis Type="Italic">Tripneustes ventricosus</Emphasis>: the importance of food type

The sea urchin Tripneustes ventricosus is a common, yet relatively poorly known, grazer of seagrass beds and coral reefs throughout the Caribbean. We compared the size and abundance of urchins between adjacent seagrass and coral reef habitats (where macroalgae are the dominant primary producers). We also conducted a laboratory experiment comparing the growth rate of juvenile urchins fed a diet of either macroalgae or seagrass. Reef urchins had significantly larger test diameter than those in the seagrass on some sampling dates. This size difference may be at least partially explained by diet, because laboratory-reared urchins fed macroalgae grew significantly faster than those fed seagrass. The seagrass population, however, was stable over time, whereas the reef population exhibited strong fluctuations in abundance. Overall, our study indicates that both the seagrass and coral reef habitats are capable of supporting healthy, reproductive populations of T. ventricosus. Each, however, appears to offer a distinct advantage: faster growth on the reef and greater population stability in the seagrass.     

Sulfur cycling and seagrass (Posidonia oceanica) status in carbonate sediments

Sulfur cycling was investigated in carbonate-rich and iron-poor sediments vegetated with Posidonia oceanica in oligotrophic Mediterranean around Mallorca Island, Spain, to quantify sulfate reduction and pools of sulfide in seagrass sediments. The oxygen penetration depth was low (< 4.5 mm) and sulfate reduction rates were relatively high (0.7–12 mmol m^–2d^–1). The total pools of reduced sulfides were remarkably low (< 5 mol S m^–2) indicating a fast turnover of reduced sulfides in these iron-poor sediments. The sulfate reduction rates were generally higher in vegetated compared to bare sediments possible due to enhanced sedimentation of sestonic material inside the seagrass meadows. The sulfate reduction rates were positively correlated with the seasonal variation in water temperature and negatively correlated with the shoot density indicating that the microbial activity was controlled by temperature and release of oxygen from the roots. The pools of reduced sulfides were low in these iron-poor sediments leading to high oxygen consumption for reoxidation. The sediments were highly anoxic as shown by relatively low oxygen penetration depths (< 4.5 mm) in these low organic sediments. The net shoot recruitment rate was negative in sediments enriched with organic matter, suggesting that organic matter enrichment may be an important factor for seagrass status in these iron-depleted carbonate sediments.     

Seagrasses and eutrophication

This review summarizes the historic, correlative field evidence and experimental research that implicate cultural eutrophication as a major cause of seagrass disappearance. We summarize the underlying physiological responses of seagrass species, the potential utility of various parameters as indicators of nutrient enrichment in seagrasses, the relatively sparse available information about environmental conditions that exacerbate eutrophication effects, and the better known array of indirect stressors imposed by nutrient over-enrichment that influence seagrass growth and survival. Seagrass recovery following nutrient reductions is examined, as well as the status of modeling efforts to predict seagrass response to changing nutrient regimes.The most common mechanism invoked or demonstrated for seagrass decline under nutrient over-enrichment is light reduction through stimulation of high-biomass algal overgrowth as epiphytes and macroalgae in shallow coastal areas, and as phytoplankton in deeper coastal waters. Direct physiological responses such as ammonium toxicity and water-column nitrate inhibition through internal carbon limitation may also contribute. Seagrass decline under nutrient enrichment appears to involve indirect and feedback mechanisms, and is manifested as sudden shifts in seagrass abundance rather than continuous, gradual changes in parallel with rates of increased nutrient additions. Depending on the species, interactions of high salinity, high temperature, and low light have been shown to exacerbate the adverse effects of nutrient over-enrichment. An array of indirect effects of nutrient enrichment can accelerate seagrass disappearance, including sediment re-suspension from seagrass loss, increased system respiration and resulting oxygen stress, depressed advective water exchange from thick macroalgal growth, biogeochemical alterations such as sediment anoxia with increased hydrogen sulfide concentrations, and internal nutrient loading via enhanced nutrient fluxes from sediments to the overlying water. Indirect effects on trophic structure can also be critically important, for example, the loss of herbivores, through increased hypoxia/anoxia and other habitat shifts, that would have acted as “ecological engineers” in promoting seagrass survival by controlling algal overgrowth; and shifts favoring exotic grazers that out-compete seagrasses for space. Evidence suggests that natural seagrass population shifts are disrupted, slowed or indefinitely blocked by cultural eutrophication, and there are relatively few known examples of seagrass meadow recovery following nutrient reductions.Reliable biomarkers as early indicators of nutrient over-enriched seagrass meadows would benefit coastal resource managers in improving protective measures. Seagrasses can be considered as “long-term" integrators (days to weeks) of nutrient availability, especially through analyses of their tissue content, and of activities of enzymes such as nitrate reductase and alkaline phosphatase. The ratio of leaf nitrogen content to leaf mass has also shown promise as a “nutrient pollution indicator” for the seagrass Zostera marina, with potential application to other species. In modeling efforts, seagrass response to nutrient loading has proven difficult to quantify beyond localized areas because long-term data consistent in quality are generally lacking, and high inter-annual variability in abundance and productivity depending upon stochastic meteorological and hydrographic conditions.Efforts to protect remaining seagrass meadows from damage and loss under eutrophication, within countries and across regions, are generally lacking or weak and ineffective. Research needs to further understand about seagrasses and eutrophication should emphasize experimental studies to assess the response of a wider range of species to chronic, low-level as well as acute, pulsed nutrient enrichment. These experiments should be conducted in the field or in large-scale mesocosms following appropriate acclimation, and should emphasize factor interactions (N, P, C; turbidity; temperature; herbivory) to more closely simulate reality in seagrass ecosystems. They should scale up to address processes that occur over larger scales, including food-web dynamics that involve highly mobile predators and herbivores. Without any further research, however, one point is presently very clear: Concerted local and national actions, thus far mostly lacking, are needed worldwide to protect remaining seagrass meadows from accelerating cultural eutrophication in rapidly urbanizing coastal zones.     

Development of a ‘sediment-stress’ functional-level indicator for the seagrass Halophila ovalis

Understanding mechanistic relationships between seagrass and their environmental stressors should be considered for effective management of estuaries and may inform on why change has occurred. We aimed to develop indicators for seagrass health in response to sediment conditions for the Swan-Canning Estuary, south-west Australia. This article describes the development of a new sediment-stress indicator, relating aspects of seagrass productivity with sediment sulfur dynamics. Sulfur stable isotope ratio and total sulfur were measured monthly within the roots, rhizomes and leaves of Halophila ovalis, and significantly varied across sites and months. The growth of seagrass over the summer months appeared restricted by sediment condition, with growth of seagrass lower when sediment derived sulfur and/or total sulfur within rhizome of leaf tissues was higher. H. ovalis appeared quite tolerant of sulfide intrusion within the root compartment, but growth was compromised when sulfide breached the root–rhizome barrier. The tightest correlation between potential sulfur metrics and seagrass growth was observed for the ratio (δ³⁴S_leaf + 30)/(TS_leaf), and it is this ratio that we propose may be a useful sediment-stress indicator for seagrass. The study also highlights that sediment condition needs to be considered at the meadow scale.     

Methane emission and sulfide levels increase in tropical seagrass sediments during temperature stress: A mesocosm experiment

Climate change‐induced ocean warming is expected to greatly affect carbon dynamics and sequestration in vegetated shallow waters, especially in the upper subtidal where water temperatures may fluctuate considerably and can reach high levels at low tides. This might alter the greenhouse gas balance and significantly reduce the carbon sink potential of tropical seagrass meadows. In order to assess such consequences, we simulated temperature stress during low tide exposures by subjecting seagrass plants (Thalassia hemprichii) and associated sediments to elevated midday temperature spikes (31, 35, 37, 40, and 45°C) for seven consecutive days in an outdoor mesocosm setup. During the experiment, methane release from the sediment surface was estimated using gas chromatography. Sulfide concentration in the sediment pore water was determined spectrophotometrically, and the plant's photosynthetic capacity as electron transport rate (ETR), and maximum quantum yield (Fv/Fm) was assessed using pulse amplitude modulated (PAM) fluorometry. The highest temperature treatments (40 and 45°C) had a clear positive effect on methane emission and the level of sulfide in the sediment and, at the same time, clear negative effects on the photosynthetic performance of seagrass plants. The effects observed by temperature stress were immediate (within hours) and seen in all response variables, including ETR, Fv/Fm, methane emission, and sulfide levels. In addition, both the methane emission and the size of the sulfide pool were already negatively correlated with changes in the photosynthetic rate (ETR) during the first day, and with time, the correlations became stronger. These findings show that increased temperature will reduce primary productivity and increase methane and sulfide levels. Future increases in the frequency and severity of extreme temperature events could hence reduce the climate mitigation capacity of tropical seagrass meadows by reducing CO₂ sequestration, increase damage from sulfide toxicity, and induce the release of larger amounts of methane.     

Seagrasses are negatively affected by organic matter loading and Arenicola marina activity in a laboratory experiment

When two ecosystem engineers share the same natural environment, the outcome of their interaction will be unclear if they have contrasting habitat-modifying effects (e.g., sediment stabilization vs. sediment destabilization). The outcome of the interaction may depend on local environmental conditions such as season or sediment type, which may affect the extent and type of habitat modification by the ecosystem engineers involved. We mechanistically studied the interaction between the sediment-stabilizing seagrass Zostera noltii and the bioturbating and sediment-destabilizing lugworm Arenicola marina, which sometimes co-occur for prolonged periods. We investigated (1) if the negative sediment destabilization effect of A. marina on Z. noltii might be counteracted by positive biogeochemical effects of bioirrigation (burrow flushing) by A. marina in sulfide-rich sediments, and (2) if previously observed nutrient release by A. marina bioirrigation could affect seagrasses. We tested the individual and combined effects of A. marina presence and high porewater sulfide concentrations (induced by organic matter addition) on seagrass biomass in a full factorial lab experiment. Contrary to our expectations, we did not find an effect of A. marina on porewater sulfide concentrations. A. marina activities affected the seagrass physically as well as by pumping nutrients, mainly ammonium and phosphate, from the porewater to the surface water, which promoted epiphyte growth on seagrass leaves in our experimental set-up. We conclude that A. marina bioirrigation did not alleviate sulfide stress to seagrasses. Instead, we found synergistic negative effects of the presence of A. marina and high sediment sulfide levels on seagrass biomass.     

Understanding invasion as a process: the case of <Emphasis Type="Italic">Phalaris arundinacea</Emphasis> in wet prairies

Invasive plants that most threaten biodiversity are those that rapidly form a monospecific stand, like the clonal grass, Phalaris arundinacea. Understanding complex and potentially interacting factors that are common in urban and agricultural landscapes and underlie rapid invasions requires an experimental, factorial approach. We tested the effects of flooding and nutrient and sediment additions (3 × 3 × 3 = 27 treatments, plus a control with no additions) on invasion of Phalaris into mesocosms containing wet prairie vegetation. We discovered a three-step invasion and degradation process: (1) initially, resident native species declined with prolonged flooding and sediment additions, and (2) prolonged flooding, sedimentation, and nutrients accelerated Phalaris aboveground growth; biomass rose to 430 times that of the control within just two growing seasons. The dramatic expansion of Phalaris in the second year resulted in the formation of monospecific stands in over one-third of the treatments, as (3) native species continued their decline in year 2. Disturbances acted alone and in combination to make the resident wetland community more invasible and Phalaris more aggressive, leading to monospecific stands. Yet, Phalaris did not always “win”: under the least disturbed conditions, the resident plant canopy remained dense and vigorous and Phalaris remained small. When anthropogenic disturbances coincide with increases in the gross supply of resources, more tolerant, fast-growing, and morphologically plastic plants like Phalaris can invade very rapidly. The fluctuating resource hypothesis should thus be refined to consider the role of interacting disturbances in facilitating invasions.     

Large- and small-scale effects of habitat structure on rates of predation: how percent coverage of seagrass affects rates of predation and siphon nipping on an infaunal bivalve

Landscape ecology, predominantly a terrestrial discipline, considers the effect of large-scale (tens of meters to kilometers) spatial patterns of habitats on ecological processes such as competition, predation, and flow of energy. In this study, a landscape-ecology approach was applied to a marine soft-sediment environment to examine rates of predation and transfer of secondary production in and around vegetated habitats. Seagrass beds naturally occur in a variety of spatial configurations from patches 1–10s of meters across with interspersed unvegetated sediments (i.e., patchy coverage) to more continuous coverage with little or no bare sediment. I designed experiments to address how percent coverage of seagrass in a 100-m² area of seafloor, and the spatial arrangement (degree of patchiness or fragmentation) of an equal area (100 m²) of vegetation affected predation (lethal) and siphon nipping (sublethal) intensity on an infaunal bivalve, Mercenaria mercenaria (hard clam). Measures of seagrass density and biomass with different percent coverage of seagrass were also made. When clams were placed in both the vegetated and unvegetated portions of the seafloor nearly twice as many clams were recovered live with 99% seagrass cover than with 23% seagrass cover, while survivorship was intermediate with 70% cover. Cropping of clam siphons from both the vegetated and unvegetated sediments was also affected by the amount of seagrass cover in a 100-m² area of seafloor: mean adjusted siphon weights were approximately 76% heavier from the 99% seagrass cover treatment than from the 70% or 23% cover treatments. Survivorship of clams placed within an equal area of seagrass in very patchy, patchy, and continuous spatial configurations was 40% higher in the continuous seagrass treatment than in either of the two patchy treatments. This study demonstrates that transfer of secondary production in the form of predation and cropping on an infaunal organism is altered as the percent cover of seagrass changes. While large-scale changes in the amount and spatial patterning of vegetation may affect habitat utilization patterns and foraging HGLoopbehavior, increased seagrass density and biomass with increased percent coverage of seagrass limit any conclusions concerning predator foraging behavior and feeding success in response to patch shapes and sizes. Instead, local changes in seagrass characteristics provide the most compelling explanation for the observed results.     

Nutrient availability in the sediment and the reciprocal effects between the native seagrass Cymodocea nodosa and the introduced rhizophytic alga Caulerpa taxifolia

Two reciprocal experiments testing for the effects of nutrient addition in the sediment and competitive interactions between the native seagrass Cymodocea nodosa (Ucria) Ascherson and the introduced alga Caulerpa taxifolia (Vahl) C. Agardh were performed. This study was conducted for 13 months (August 1995 until September 1996) in a bay on the south coast of Elba Island (Italy). Each experiment consisted of the manipulation of the level of nutrients (addition vs. control) and the manipulation of the neighbours (presence vs. removal). Response variables were blade density and size for one experiment and shoot density and leaf length of seagrass in the other. Results indicated that the presence of Caulerpa taxifolia did not affect significantly Cymodocea nodosa shoot density and the increased nutrient availability in the sediment did not alter this pattern. Neither the removal of the canopy of the seagrass nor the fertilization of the sediment has influenced significantly the density of the alga. Both species, where co-occurring, show larger size than where the neighbour is removed. Hence, results of this study suggest that the two species on the long term are likely to coexist and that the high nutrient supply of the sediment would not enhance the probability of success neither of the seagrass nor of the alga. Predictions made on the basis of short-term results, that high nutrient loads of the substratum would have represented an even more suitable condition for C. taxifolia to colonize C. nodosa beds and that on the long-term the alga has a high probability of success, did not occur.