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Multi-specific seagrass meadow assemblages dominate most tropical intertidal regions but the relative role of environmental stress in determining distribution patterns is still uncertain. Here we combine observational and experimental approaches to examine aerial exposure as a factor driving species occurrence patterns in intertidal meadows of the Andaman archipelago, where up to 6 seagrass species co-occur. In the studied meadow, patterns of exposure did not map onto distance from the coast, instead creating a patchy matrix of exposure, based on fine-scale bathymetric differences. Distributional surveys showed that seagrass species were similarly patchy, often tracking the degree of aerial exposure during low tide. While some species (Halophila ovalis, Halophila minor, and Thalassia hemprichii) frequently occurred in submerged or subtidal areas and were rarely found in completely exposed areas, other species (Cymodocea rotundata, Halophila beccarii, and Halodule uninervis) also occupied areas that were subject to partial or complete aerial exposure during low tide. To confirm this pattern, we used field-based transplant experiments, employing a natural gradient of tidal exposure to subject six seagrass species to different desiccation exposure times. After a month, H. beccarii and H. uninervis transplants survived in areas that sustained more than 3 h of aerial tidal exposure without significant mortality, compared with other species (H. ovalis, H. minor, T. hemprichii, C. rotundata) that showed dramatic shoot mortality at the same exposure regimes. For all species, 4 h represented the upper limit of exposure, in both experimental and distributional studies. However, despite their wider tolerance of exposure to air, H. beccarii and H. uninervis did not dominate the entire meadow. This could be a result either of their poor tolerance to other environmental factors or their lower competitive abilities among other mechanisms. This suggests that in tropical multi-specific meadows, strong environmental filters could override clear intertidal zonation to create patchy matrices based on species tolerances.

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Seagrass meadows are declining at a global scale, threatening their capacity as blue carbon sinks. Restoration of seagrasses (via seagrass seeds or plant transplantation) may recover their carbon sequestration capacity. Previous studies have predominantly focused on sediment organic carbon (SOC), while variations in sediment carbon compositions remain poorly understood, limiting our comprehension of the influence of seagrass restoration on sediment carbon stability. Here, we researched the differences in surface (0-3 cm) sediment carbon compositions in response to tropical seagrass transplantation among species (Thalassia hemprichii and Enhalus acoroides); specifically, differences in labile, recalcitrant and refractory SOC, as well as sediment inorganic carbon (SIC) compositions variations under transplanted T. hemprichii and E. acoroides communities. It was found that seagrass transplantation enhanced suspended particle organic matter, and epiphyte and macroalgae input to surface sediment, which recovered the surface SOC concentration and stock rapidly to natural levels (increased ∼1.6-fold) within two years following transplantation. The elevated contribution of epiphyte and macroalgae significantly increased the surface labile sediment organic matter (SOM), but not the recalcitrant and refractory SOM composition after short-term transplantation. Meanwhile, surface SIC was significantly elevated, which might be mainly ascribed to allochthonous carbonate particle trapped under transplanted area with implications for carbon sequestration. The higher canopy and longer leaf seagrass species, E. acoroides, had elevated SOC, SIC and was more labile composition, compared to T. hemprichii transplant. Overall, this research suggests that tropical seagrass transplantation can increase the surface SOC, SIC concentration by increasing the labile organic matter and allochthonous carbonate particle input, respectively, with varying significantly among seagrass species.

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Large grazers (megaherbivores) have a profound impact on ecosystem functioning. However, how ecosystem multifunctionality is affected by changes in megaherbivore populations remains poorly understood. Understanding the total impact on ecosystem multifunctionality requires an integrative ecosystem approach, which is especially challenging to obtain in marine systems. We assessed the effects of experimentally simulated grazing intensity scenarios on ecosystem functions and multifunctionality in a tropical Caribbean seagrass ecosystem. As a model, we selected a key marine megaherbivore, the green turtle, whose ecological role is rapidly unfolding in numerous foraging areas where populations are recovering through conservation after centuries of decline, with an increase in recorded overgrazing episodes. To quantify the effects, we employed a novel integrated index of seagrass ecosystem multifunctionality based upon multiple, well-recognized measures of seagrass ecosystem functions that reflect ecosystem services. Experiments revealed that intermediate turtle grazing resulted in the highest rates of nutrient cycling and carbon storage, while sediment stabilization, decomposition rates, epifauna richness, and fish biomass are highest in the absence of turtle grazing. In contrast, intense grazing resulted in disproportionally large effects on ecosystem functions and a collapse of multifunctionality. These results imply that (i) the return of a megaherbivore can exert strong effects on coastal ecosystem functions and multifunctionality, (ii) conservation efforts that are skewed toward megaherbivores, but ignore their key drivers like predators or habitat, will likely result in overgrazing-induced loss of multifunctionality, and (iii) the multifunctionality index shows great potential as a quantitative tool to assess ecosystem performance. Considerable and rapid alterations in megaherbivore abundance (both through extinction and conservation) cause an imbalance in ecosystem functioning and substantially alter or even compromise ecosystem services that help to negate global change effects. An integrative ecosystem approach in environmental management is urgently required to protect and enhance ecosystem multifunctionality.

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Tropical seagrass meadows are formed by an array of seagrass species that share the same space. Species sharing the same plot are competing for resources, namely light and inorganic nutrients, which results in the capacity of some species to preempt space from others. However, the drivers behind seagrass species competition are not completely understood. In this work, we studied the competitive interactions among tropical seagrass species of Unguja Island (Zanzibar, Tanzania) using a trait-based approach. We quantified the abundance of eight seagrass species under different trophic states, and selected nine traits related to light and inorganic nutrient preemption to characterize the functional strategy of the species (leaf maximum length and width, leaves per shoot, leaf mass area, vertical rhizome length, shoots per meter of ramet, rhizome diameter, roots per meter of ramet, and root maximum length). From the seagrass abundance we calculated the probability of space preemption between pairs of seagrass species and for each individual seagrass species under the different trophic states. Species had different probabilities of space preemption, with the climax species Thalassodendron ciliatum, Enhalus acoroides, Thalassia hemprichii, and the opportunistic Cymodocea serrulata having the highest probability of preemption, while the pioneer and opportunistic species Halophila ovalis, Syringodium isoetifolium, Halodule uninervis, and Cymodocea rotundata had the lowest. Traits determining the functional strategy showed that there was a size gradient across species. For two co-occurring seagrass species, probability of preemption was the highest for the larger species, it increased as the size difference between species increased and was unaffected by the trophic state. Competitive interactions among seagrass species were asymmetrical, i.e., negative effects were not reciprocal, and the driver behind space preemption was determined by plant size. Seagrass space preemption is a consequence of resource competition, and the probability of a species to exert preemption can be calculated using a trait-based approach.

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Seagrass habitats at the Cocos (Keeling) Islands (CKI), a remote atoll in the Indian Ocean, have suffered a catastrophic decline over the last decade. Seagrass monitoring (1996-2020) in relation to dredging and coastal development works (2009 to 2011) provide a historical baseline, and document the decline of mixed tropical seagrass Thalassia hemprichii and macroalgal (predominantly Caulerpa spp.) beds over a decadal scale time series. Attribution of loss to coastal development is confounded by lagoon-wide die-off events in 2007, 2009 and 2012 and high air and water temperatures from 2009 to 2016, with evidence of broad scale changes, visible in satellite imagery between 2006 and 2018. We conclude that up to 80% of seagrass habitats in the CKI lagoon (~1200 ha) have been lost due to multiple stressors including episodic die-off events related to high temperatures and calm conditions, and loss due to sediment disturbance and increased turbidity. Grazing pressure from the resident green sea turtles (Chelonia mydas) may have also exacerbated the loss of seagrass, which in turn poses a dire threat to their ongoing health and survival. This study highlights the fragility of tropical seagrass habitats and the cascading effect of system imbalance as a result of anthropogenic pressures and climate drivers. Although small in comparison to global estimates, the loss of seagrass habitats at CKI could change the entire ecosystem of a remote atoll. Due to the significance of the Thalassia beds for coastal stability, as food for an isolated population of green sea turtles and as a fish nursery, rehabilitation efforts are warranted.

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Two novel diterpenes Enhoidin A (1) and Enhoidin B (2) featuring an unusual gibberellane skeleton were isolated from the stems and leaves of Enhalus acoroides. Their structures were elucidated on the basis of spectroscopic analysis including 1D and 2D NMR techniques and HR-ESI-MS. This is the first time that this type of lactone ring between C-18 and C-20 has been found among gibberellanes from the tropical seagrasses. Evaluation of the all compounds for cytotoxicity against four human cancer cell lines (MCF-7, HCT-116, HepG-2 and HeLa), and showed moderate cytotoxic activities.

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Seagrass longevity up to 47 years in well-restored, well-sited seagrass restorations are demonstrated from 253 trials at 83 regional sites in tropical and subtropical portions of three oceans (Atlantic, Pacific, Indian Oceans). These trials include over 3.04 million planted units into 306.3 ha. Approximately 12% of the total global tropical restored seagrass by Van Katwijk, Thorhaug et al. (2016) calculations from 1786 trials are included. Almost all projects herein reviewed persisted since date of planting except several cases with harsh anthropogenic impact or forceful natural events in first post-planting months. The oldest tropical/subtropical restoration continually observed is 47 yrs, many are 35 yrs. An array of observed and/or measured restored services accompanied these. This review may provide informational background for government resource managers, legislators, scientists, and citizens concerning tropical/subtropical seagrass longevity. This data from these trials may substantiate future seagrass restoration investments. Public outreach, national & regional government training,and outreach occurred, needing continuation.

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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.

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The effect of repeated midday temperature stress on the photosynthetic performance and biomass production of seagrass was studied in a mesocosm setup with four common tropical species, including Thalassia hemprichii, Cymodocea serrulata, Enhalus acoroides, and Thalassodendron ciliatum. To mimic natural conditions during low tides, the plants were exposed to temperature spikes of different maximal temperatures, that is, ambient (29-33°C), 34, 36, 40, and 45°C, during three midday hours for seven consecutive days. At temperatures of up to 36°C, all species could maintain full photosynthetic rates (measured as the electron transport rate, ETR) throughout the experiment without displaying any obvious photosynthetic stress responses (measured as declining maximal quantum yield, Fv/Fm). All species except T. ciliatum could also withstand 40°C, and only at 45°C did all species display significantly lower photosynthetic rates and declining Fv/Fm. Biomass estimation, however, revealed a different pattern, where significant losses of both above- and belowground seagrass biomass occurred in all species at both 40 and 45°C (except for C. serrulata in the 40°C treatment). Biomass losses were clearly higher in the shoots than in the belowground root-rhizome complex. The findings indicate that, although tropical seagrasses presently can cope with high midday temperature stress, a few degrees increase in maximum daily temperature could cause significant losses in seagrass biomass and productivity.

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Existing mitigations to address deterioration in water clarity associated with human activities are based on responses from single seagrass species but may not be appropriate for diverse seagrass assemblages common to tropical waters. We present findings from a light experiment designed to determine the effects of magnitude and duration of low light on a mixed tropical seagrass assemblage. Mixed assemblages of three commonly co-occurring Indo-West Pacific seagrasses, Cymodocea serrulata, Halodule uninervis and Halophila ovalis were grown in climate-controlled tanks, where replicate pots were subjected to a gradient in light availability (0.9-21.6 mols PAR m-2 day-1) for 12 weeks. Increased shading resulted in declines in growth and changes in cellular and photosynthesis responses for all species, although time-scale and magnitude of response were species-specific. Applying management criteria (e.g. thresholds) relevant to one species may under- or over-estimate potential for impact on other species and the meadow as a whole.

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Rising sea water temperature will play a significant role in responses of the world's seagrass meadows to climate change. In this study, we investigated seasonal and latitudinal variation (spanning more than 1,500 km) in seagrass productivity, and the optimum temperatures at which maximum photosynthesis and net productivity (for the leaf and the whole plant) occurs, for three seagrass species (Cymodocea serrulata, Halodule uninervis, and Zostera muelleri). To obtain whole plant net production, photosynthesis, and respiration rates of leaves and the root/rhizome complex were measured using oxygen-sensitive optodes in closed incubation chambers at temperatures ranging from 15 to 43°C. The temperature-dependence of photosynthesis and respiration was fitted to empirical models to obtain maximum metabolic rates and thermal optima. The thermal optimum (Topt) for gross photosynthesis of Z. muelleri, which is more commonly distributed in sub-tropical to temperate regions, was 31°C. The Topt for photosynthesis of the tropical species, H. uninervis and C. serrulata, was considerably higher (35°C on average). This suggests that seagrass species are adapted to water temperature within their distributional range; however, when comparing among latitudes and seasons, thermal optima within a species showed limited acclimation to ambient water temperature (Topt varied by 1°C in C. serrulata and 2°C in H. uninervis, and the variation did not follow changes in ambient water temperature). The Topt for gross photosynthesis were higher than Topt calculated from plant net productivity, which includes above- and below-ground respiration for Z. muelleri (24°C) and H. uninervis (33°C), but remained unchanged at 35°C in C. serrulata. Both estimated plant net productivity and Topt are sensitive to the proportion of below-ground biomass, highlighting the need for consideration of below- to above-ground biomass ratios when applying thermal optima to other meadows. The thermal optimum for plant net productivity was lower than ambient summer water temperature in Z. muelleri, indicating likely contemporary heat stress. In contrast, thermal optima of H. uninervis and C. serrulata exceeded ambient water temperature. This study found limited capacity to acclimate: thus the thermal optima can forewarn of both the present and future vulnerability to ocean warming during periods of elevated water temperature.

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Seasonal dynamics were studied by monthly monitoring of biological and environmental variables in permanent quadrats in two contrasting intertidal seagrass beds in South Sulawesi, Indonesia, from February 1991 to January 1992. Datasets were analysed with canonical correlation analysis for correlations between environmental and biological variables. Considerable variation in biomass, production and plant tissue nutrient contents in a monospecific seagrass bed of Enhalus acoroides, growing on a coastal terrigenous mudbank (Gusung Tallang), was assumed to be related to riverine influences of the nearby Tallo River. The variation in seagrass variables at this site could, however, not be significantly correlated to seasonal patterns in rainfall, salinity, tides, nutrient availability, water motion or turbidity. A seasonal cycle in biomass, production and nutrient contents in a mixed seagrass bed of Thalassia hemprichii and E. acoroides, growing on carbonate sand on the reef flat of an offshore coral island (Barang Lompo), was found to be largely determined by tidal exposure and water motion. Exposure of the intertidal seagrass bed during hours of low water during spring tides showed a gradual shift from exposure during the night (January-June) to exposure during daylight (July-December). Daylight exposure resulted in a significant loss of above-ground plant biomass through desiccation and 'burning' of leaves. The observed seasonal dynamics of the seagrass bed on reef sediment contrast with reports from the Caribbean, where the effect of tidal exposure on comparable shallow-water seagrass communities is relatively insignificant due to a small tidal amplitude.