Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

Nutrient impacts on epifaunal density and species composition in a subtropical seagrass bed

The capacity of epifauna to control algal proliferation following nutrient input depends on responses of both grazers and upper trophic level consumers to enrichment. We examined the responses of Thalassia testudinum (turtle grass) epifaunal assemblages to nutrient enrichment at two sites in Florida Bay with varying levels of phosphorus limitation. We compared epifaunal density, biomass, and species diversity in 2 m² plots that had either ambient nutrient concentrations or had been enriched with nitrogen and phosphorus for 6 months. At the severely P-limited site, total epifaunal density and biomass were two times higher in enriched than in unenriched plots. Caridean shrimp, grazing isopods, and gammarid amphipods accounted for much of the increase in density; brachyuran crabs, primary predatory fish, and detritivorous sea cucumbers accounted for most of the increase in biomass. At the less P-limited site, total epifaunal density and biomass were not affected by nutrient addition, although there were more caridean shrimp and higher brachyuran crab and pink shrimp biomass in enriched plots. At both sites, some variation in epifaunal density and biomass was explained by features of the macrophyte canopy, such as T. testudinum and Halodule wrightii percent cover, suggesting that enrichment may change the refuge value of the macrophyte canopy for epifauna. Additional variation in epifaunal density and biomass was explained by epiphyte pigment concentrations, suggesting that enrichment may change the microalgal food resources that support grazing epifauna. Increased epifaunal density in enriched plots suggests that grazers may be able to control epiphytic algal proliferation following moderate nutrient input to Florida Bay. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Plant-herbivore interactions in seagrass meadows

During the past two decades we have gained much insight into the factors that regulate the productivity of seagrass dominated ecosystems, especially those at low latitudes. Here, we review and reassess the importance of plant-herbivore interactions in seagrass meadows, focusing on recent studies that have examined: 1) grazing on live seagrass leaves; 2) consumption of epiphytic algae growing on seagrass leaves; and 3) consumption of planktonic algae from the waters surrounding seagrass meadows. The major conclusion is that, in contrast to what has been reported in much of the literature on food webs in seagrass meadows, a diverse grazing pathway continues to represent an important conduit for the transfer of energy from the primary producers to higher order consumers. This remains true, although in many areas consumption of seagrasses is reduced in an historical context, owing to the overharvesting of many large species of herbivorous waterfowl, turtles and mammals.We also summarize our view of the important gaps in understanding the broadly defined topic of herbivory in seagrass-dominated ecosystems. We suggest that future studies should focus on: understanding the foraging strategies of seagrass herbivores; quantifying the impact of herbivory on seagrass demography, including effects on sexual reproduction, the fate of flowers, and the production of fruits and seeds; and documenting the commonness of compensatory responses to grazing. In addition, the role of chemical defenses in seagrass species remains inadequately investigated. Studies of the roles of nutritional content (as measured by C/N/P ratios) and chemical defenses are also fertile grounds for future studies of epiphytes and their grazers, as are additional experiments to quantify the relative roles of top-down and bottom-up factors as they determine algal growth and abundance. There is also a need to expand the geographical scope of studies of epiphyte-grazer interactions from cold temperate to sub-tropical and tropical waters. Suspension feeders also need to be studied more broadly, with additional experiments required to quantify their effects on water clarity and their ability to fertilize pore waters, and whether benefits from these activities balances the costs of shading and competition for space that can result from both epifaunal and infaunal suspension feeders.     

Global seagrass distribution and diversity: A bioregional model

Seagrasses, marine flowering plants, are widely distributed along temperate and tropical coastlines of the world. Seagrasses have key ecological roles in coastal ecosystems and can form extensive meadows supporting high biodiversity. The global species diversity of seagrasses is low (< 60 species), but species can have ranges that extend for thousands of kilometers of coastline. Seagrass bioregions are defined here, based on species assemblages, species distributional ranges, and tropical and temperate influences. Six global bioregions are presented: four temperate and two tropical. The temperate bioregions include the Temperate North Atlantic, the Temperate North Pacific, the Mediterranean, and the Temperate Southern Oceans. The Temperate North Atlantic has low seagrass diversity, the major species being Zostera marina, typically occurring in estuaries and lagoons. The Temperate North Pacific has high seagrass diversity with Zostera spp. in estuaries and lagoons as well as Phyllospadix spp. in the surf zone. The Mediterranean region has clear water with vast meadows of moderate diversity of both temperate and tropical seagrasses, dominated by deep-growing Posidonia oceanica. The Temperate Southern Oceans bioregion includes the temperate southern coastlines of Australia, Africa and South America. Extensive meadows of low-to-high diversity temperate seagrasses are found in this bioregion, dominated by various species of Posidonia and Zostera. The tropical bioregions are the Tropical Atlantic and the Tropical Indo-Pacific, both supporting mega-herbivore grazers, including sea turtles and sirenia. The Tropical Atlantic bioregion has clear water with a high diversity of seagrasses on reefs and shallow banks, dominated by Thalassia testudinum. The vast Tropical Indo-Pacific has the highest seagrass diversity in the world, with as many as 14 species growing together on reef flats although seagrasses also occur in very deep waters. The global distribution of seagrass genera is remarkably consistent north and south of the equator; the northern and southern hemispheres share ten seagrass genera and only have one unique genus each. Some genera are much more speciose than others, with the genus Halophila having the most seagrass species. There are roughly the same number of temperate and tropical seagrass genera as well as species. The most widely distributed seagrass is Ruppia maritima, which occurs in tropical and temperate zones in a wide variety of habitats. Seagrass bioregions at the scale of ocean basins are identified based on species distributions which are supported by genetic patterns of diversity. Seagrass bioregions provide a useful framework for interpreting ecological, physiological and genetic results collected in specific locations or from particular species.     

Detection of inconspicuous epiphytic algae supporting food webs in seagrass meadows

Summary Detritus from common seagrasses and other marine angiosperms may often be a less important basis for estuarine food webs than previously believed. In NW Gulf of Mexico seagrass meadows, epiphytic algae have high productivities, palatability, and a more important trophic role than common large plants have. Interdisciplinary field experiments show (1) intensive night-time ingestion of epiphytes by various invertebrate detritivores, (2) very high productivity of epiphytic algae on seagrasses, and (3) assimilation of epiphytes rather than seagrasses, as measured by ¹³C comparisons. These combined data show that many naturally concentrated and potentially competing invertebrates in Gulf of Mexico seagrass meadows feed largely on the algal overgrowth on seagrass blades, even when such algae appear to be sparse. Primary productivity of these epiphytic algae can equal that of the seagrasses, per blade or per unit biomass. Animal ¹³C values tracked epiphytic values rather than seagrass values when comparisons were made over six sites. These measurements reinforce the view that epiphytic algae can be the primary basis of the food web in seagrass meadows.Contribution No. 608 of The University of Texas Marine Science Institute     

Non‐consumptive effects of avian predators on fish behavior and cascading indirect interactions in seagrasses

Top–down impacts of avian predators are often overlooked in marine environments despite evidence from other systems that birds significantly impact animal distribution and behavior; instead, birds are typically recognized for the impacts of their nutrient rich guano. This is especially true in shallow seagrass meadows where restoration methods utilize bird perches or stakes to attract birds as a passive fertilizer delivery system that promotes the regrowth of damaged seagrasses. However, this method also increases the local density of avian piscivores that may have multiple unexplored non‐consumptive effects on fish behavior and indirect impacts to seagrass communities. We utilized laboratory and field experiments to investigate whether visual cues of avian predators impacted the behavior of the dominant demersal fish in seagrass habitats, the pinfish Lagodon rhomboides, and promoted cascading interactions on seagrass‐associated fauna and epiphytes. In laboratory mesocosms, pinfish displayed species specific responses to models of avian predators, with herons inducing the greatest avoidance behaviors. Avoidance patterns were confirmed in field seagrass meadows where heron models significantly reduced the number of fish caught in traps. In a long term field experiment, we investigated whether avian predators caused indirect non‐consumptive effects on seagrass communities by monitoring fish abundances, invertebrate epiphyte grazers, and the seagrass epiphytes in response to heron models, bird exclusions, and bird stakes. On average, more fish were recovered under bird exclusions and fewer fish under heron models. However, we found no evidence of cascading effects on invertebrate grazers or epiphytes. Bird stake treatments only displayed a simple nutrient effect where higher bird abundances resulted in higher epiphyte biomass. Our results indicate that although birds and their visual cues can affect fish and epiphyte abundance through non‐consumptive effects and nutrient enrichment, these impacts do not propagate beyond one trophic level, most likely because of dampening by omnivory and larger scale processes.     

Distribution of seagrasses along the Indian coast

Seagrass environments, from the main coast of India, Lakshadweep and Andaman Islands, were surveyed for seagrass and marine algal composition. Extensive seagrass meadows and the maximum number of species (seven genera and 12 species) occurred along the Tamil Nadu coast. Seagrasses were observed from intertidal to subtidal regions down to 8 m depth. Thalassia hemprichii (Ehrenberg) Aschers. and Cymodocea serrulata (R. Brown) Aschers. & Magnus were the dominant seagrasses in the subtidal zones. Halophila beccarii Aschers. was restricted to the intertidal mudflats in association with mangroves. The rich growth of seagrasses along the Tamil Nadu coast and Lakshadweep can be attributed mainly to high salinity, clarity of the water and sandy substratum. One hundred species of marine algae were recorded from the seagrass environments of India.     

Varying foraging strategies of Labridae in seagrass habitats: Herbivory in temperate seagrass meadows?

The diets of five species of Labridae in south-western Australia were examined to determine whether: (1) grazing of seagrass and epiphytic algae is a prominent feature of the food web within the deeper seagrass meadows of this temperate region; (2) levels of grazing differ among different seagrass systems; and diets differ among these closely-related species. Fish were collected seasonally from three seagrass habitats mainly comprising either Posidonia sinuosa, Posidonia coriacea or Amphibolis griffithii between the summer of 1996/97 and spring of 1997. Consumption of considerable amounts of algae and seagrass by Odax acroptilus and seagrass by Haletta semifasciata indicates that macrophyte grazing by fish is a component of the trophic dynamics of south-western Australian seagrass meadows. O. acroptilus and H. semifasciata were both omnivorous, feeding on a range of epifauna, infauna and flora, whereas Siphonognathus radiatus, Neoodax balteatus and Notolabrus parilus were carnivorous, feeding predominantly on motile epifauna, such as molluscs and crustaceans. The level of macrophyte grazing is likely to be underestimated in temperate offshore meadows of P. sinuosa and A. griffithii where omnivorous labrids, monacanthids and terapontids are abundant. Stable isotope data for O. acroptilus from the study region suggest that animal prey is more important to tissue maintenance than macrophyte material. Macrophytes may be grazed to acquire attached animal prey or for fulfilling energy requirements. Based on the distribution of prey, it appears that species in A. griffithii meadows forage within and below the seagrass canopy, whilst species in P. sinuosa meadows are likely to forage towards the basal area of this seagrass.     

Reti trofiche e flussi di energia nei sistemi a fanerogame marine

Abstract

Food webs and energy flow in seagrass ecosystems. A review on the pathways in the food webs of seagrass ecosystems, both tropical and temperate, with a particular emphasis to Mediterranean Posidonia oceanica meadows is given. Three main pathways of energy transfer from primary producers (host plant and algal epiphytes) were identified: i) the plant itself through photosynthetic tissue; ii) the leaf detritus which in some species forms a litter compartment; iii) the algal epiphytes of leaf blades. The detritus and epiphyte ways are the most common, but they can be differently important according to the season and the spatial patterns of the meadows.     

Temporal changes in the abundance, leaf growth and photosynthesis of three co-occurring Philippine seagrasses

The analysis of the temporal changes in shoot density, areal leaf biomass, leaf growth and parameters of the photosynthesis–irradiance relationship of three tropical seagrass species (Enhalus acoroides, Thalassia hemprichii and Cymodocea rotundata), co-existing in a shallow subtidal meadow in Cape Bolinao, Philippines, shows that species-specific traits are significant sources of temporal variability, and indicates that these seagrass species respond differently to a common environmental forcing. Species-specific differences are much less important as source of variability of the temporal change in chlorophyll concentration of seagrass leaves. The results indicate that the temporal changes in photosynthetic performance of these seagrasses were driven by environmental forcing and their specific responses to it mostly, but the temporal change in their abundance and leaf growth was also controlled by other factors. The significant contribution of species-specific factors in the temporal changes of biomass, growth and photosynthetic performance of co-occurring seagrass species in Cape Bolinao should contribute to the maintenance of the multispecific, highly productive meadows characteristic of pristine coastal ecosystems in Southeast (SE) Asia.     

Contribution of genetics and genomics to seagrass biology and conservation

Genetic diversity is one of three forms of biodiversity recognized by the IUCN as deserving conservation along with species and ecosystems. Seagrasses provide all three levels in one. This review addresses the latest advances in our understanding of seagrass population genetics and genomics within the wider context of ecology and conservation. Case studies are used from the most widely studied, northern hemisphere species Zostera marina, Z. noltii, Posidonia oceanica and Cymodocea nodosa.

We begin with an analysis of the factors that have shaped population structure across a range of spatial and temporal scales including basin-level phylogeography, landscape-scale connectivity studies, and finally, local-scale analyses at the meadow level—including the effects of diversity, clonality and mating system. Genetic diversity and clonal architecture of seagrass meadows differ within and among species at virtually all scales studied. Recent experimental studies that have manipulated seagrass genetic biodiversity indicate that genotypic diversity matters in an immediate ecological context, and enhances population growth, resistance and resilience to perturbation, with positive effects on abundance and diversity of the larger community. In terms of the longer term, evolutionary consequences of genetic/genotypic diversity in seagrass beds, our knowledge remains meagre. It is here that the new tools of ecogenomics will assist in unravelling the genetic basis for adaptation to both biotic and abiotic change. Gene expression studies will further assist in the assessment of physiological performance which may provide an early warning system under complex disturbance regimes that seagrasses are at or near their tolerance thresholds.

At the most fundamental level, ecological interactions of seagrasses with their environment depends on the genetic architecture and response diversity underlying critical traits. Hence, given the rapid progress in data acquisition and analysis, we predict an increasing role of genetic and genomic tools for seagrass ecology and conservation.     

Trophic Transfers from Seagrass Meadows Subsidize Diverse Marine and Terrestrial Consumers

In many coastal locations, seagrass meadows are part of a greater seascape that includes both marine and terrestrial elements, each linked to the other via the foraging patterns of consumers (both predators and herbivores), and the passive drift of seagrass propagules, leaves, roots and rhizomes, and seagrass-associated macroalgal detritus. With seagrasses declining in many regions, the linkages between seagrass meadows and other habitats are being altered and diminished. Thus, it is timely to summarize what is known about the prevalence and magnitude of cross-habitat exchanges of seagrass-derived energy and materials, and to increase awareness of the importance of seagrasses to adjacent and even distant habitats. To do so we examined the literature on the extent and importance of exchanges of biomass between seagrass meadows and other habitats, both in the form of exported seagrass biomass as well as transfers of animal biomass via migration. Data were most abundant for Caribbean coral reefs and Australian beaches, and organisms for which there were quantitative estimates included Caribbean fishes and North American migratory waterfowl. Overall, data from the studies we reviewed clearly showed that seagrass ecosystems provide a large subsidy to both near and distant locations through the export of particulate organic matter and living plant and animal biomass. The consequences of continuing seagrass decline thus extend far beyond the areas where seagrasses grow.     

Seagrass bed patchiness: effects on epifaunal communities in San Diego Bay, USA

Seagrass habitat structure influences epifaunal density, diversity, community composition and survival, but covariation of structural elements at multiple scales (e.g., shoot density or biomass per unit area, patch size, and patch configuration) can confound studies attempting to correlate habitat structure with ecological patterns and processes. In this study, we standardized simulated seagrass shoot density and bed area among artificial seagrass beds in San Diego Bay, California, USA to evaluate the singular effect of seagrass bed configuration (“patchiness”) on the density and diversity of seagrass epifauna. Artificial seagrass beds all were 1 m², but were composed of a single large patch (“continuous” treatment), four smaller patches (“patchy” treatment), or 16 very small patches (“very patchy” treatment). We allowed epifauna to colonize beds for 1 month, and then sampled beds monthly over the next 3 months. Effects of seagrass bed patchiness on total epifaunal density and species-specific densities were highly variable among sampling dates, and there was no general trend for the effects of fragmentation on epifaunal densities to be positive or negative. Epifaunal diversity (measured as Simpson's index of diversity) was highest in very patchy or patchy beds on two out of the three sampling dates. Very patchy beds exhibited the highest dissimilarity in community composition in the first two sampling periods (August and September), but patchy beds exhibited the highest dissimilarity in the third sampling period (October). Our results indicate that seagrass patch configuration affects patterns of epifaunal density, diversity, and community composition in the absence of covarying bed area or structural complexity, and that patchy seagrass beds may be no less valuable as a habitat than are continuous seagrass beds. The spatial pattern employed when harvesting or planting seagrass may influence epifaunal habitat use and should be a key consideration in restoration plans.     

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.     

Scale-dependant control of motile epifaunal community structure along a coral reef fishing gradient

Large-scale fishing is mostly conducted using towed gears that reduce the biomass and diversity of benthic invertebrates. However, it is impossible to differentiate between the physical disturbance effect of towed gears from the effect of fish predator removal upon benthic invertebrate communities. Here we explore the impact of fish removal alone on the community structure of small motile coral reef invertebrates (epifauna) along a subsistence fishing intensity gradient in the Lau group, Fiji. We deployed settlement plates at three areas in each of six fishing grounds and examined the density and class richness of the motile epifaunal communities and the associated algal communities in relation to the structure of fish and benthic communities. Motile epifaunal density was unrelated to fishing intensity. However, at smaller inter-area scale (0.5-10 km) motile epifaunal density was negatively related to plate algal biomass, whereas at the larger inter-fishing-ground scale (4-180 km) motile epifaunal density was positively related to the rugosity (substrate complexity) of the surrounding benthos. The class richness and diversity (Margalef's d) of motile epifaunal communities were negatively related to fishing intensity, but unrelated to grazing intensity, rugosity or algal biomass at either scale. Benthic community structure varied significantly with fishing intensity; hard-coral cover was lower and turf-algal cover was higher at high fishing pressure. The variation in benthic community structure was associated with variation in fish community structure, which in turn varied with fishing intensity. Motile epifaunal community structure upon plates was linked to the structure of the surrounding benthic community, but was not directly linked to the plate algal community. We suggest the decline in richness of the motile epifauna community along the fishing gradient is attributable to either to exploiter-mediated coexistence or the reduction in ‘habitat quality’ of the surrounding benthos. At the large spatial scale substrate complexity is the key determinant of motile epifaunal density, suggesting predation by fishes plays an important structuring role at this scale. Assuming that rugosity is inversely related to predation risk then this study represents the first evidence for spatial-dependence on the top-down (predation) vs. bottom-up (algal biomass) control of community structure. We argue fisheries exploitation, in the absence of a physical disturbance can negatively influence motile epifaunal community structure at large spatial scales.     

What lies beneath: Why knowledge of belowground biomass dynamics is crucial to effective seagrass management

Conservation of seagrasses meadows is important, because these habitats are ecologically important and under threat. Monitoring and modelling are essential tools for assessing seagrass condition and potential threats, however there are many seagrass indicators to choose from, and differentiating between natural variability and declining conditions poses a serious challenge. Tropical seagrass meadows in the Indo-Pacific, in contrast to most temperate meadows, are characterized by a multi-species composition and a year-round growth. Differences in characteristics between species growing within one meadow could induce uncertainty in the assessment of the dynamics of these meadows if variation in productivity and related biomass turnover timescales are not taken into consideration. We present data on biomass distribution, production and turnover timescales of above- and belowground tissues for three key tropical seagrass species (Thalassia hemprichii, Cymodocea rotundata and Halodule uninervis) in two mixed-species meadows in the Spermonde Archipelago, Indonesia. Seagrass leaf turnover time scales were comparable for the three studied seagrass species and varied between 25 and 30 days. Variation in leaf and rhizome turnover timescales were small (or insignificant) between the two meadows. In contrast, rhizome turnover time scales were around ten times longer than leaf turnover timescales, and large differences in rhizome turnover time scales (200–500 days) were observed between the species. The late-successional species T. hemprichii had much slower rhizome turnover compared to the two early successional species. Furthermore, since rhizome biomass has a much longer turnover time compared to leaf biomass, changes in rhizome biomass reflect effects on seagrass meadows on a much longer timescale compared to changes in leaf biomass for these tropical meadows. We conclude that belowground biomass dynamics are an important proxy to assess long-term effects of environmental stressors on seagrass ecosystems and should be included in tropical seagrass management programmes.     

Seagrass response to CO2 contingent on epiphytic algae: indirect effects can overwhelm direct effects

Increased availability of dissolved CO₂ in the ocean can enhance the productivity and growth of marine plants such as seagrasses and algae, but realised benefits may be contingent on additional conditions (e.g. light) that modify biotic interactions between these plant groups. The combined effects of future CO₂ and differing light on the growth of seagrass and their algal epiphytes were tested by maintaining juvenile seagrasses Amphibolis antarctica under three different CO₂ concentrations representing ambient, moderate future and high future forecasts (i.e. 390, 650 vs. 900 µl l^?1) and two light levels representing low and high PAR (i.e. 43 vs. 167 µmol m^?2 s^?1). Aboveground and belowground biomass, leaf growth, epiphyte cover, tissue chemistry and photosynthetic parameters of seagrasses were measured. At low light, there was a neutral to positive effect of elevated CO₂ on seagrass biomass and growth; at high light, this effect of CO₂ switched toward negative, as growth and biomass decreased at the highest CO₂ level. These opposing responses to CO₂ appeared to be closely linked to the overgrowth of seagrass by filamentous algal epiphytes when high light and CO₂ were combined. Importantly, all seagrass plants maintained positive leaf growth throughout the experiment, indicating that growth was inhibited by some experimental conditions but not arrested entirely. Therefore, while greater light or elevated CO₂ provided direct physiological benefits for seagrasses, such benefits were likely negated by overgrowth of epiphytic algae when greater light and CO₂ were combined. This result demonstrates how indirect ecological effects from epiphytes can modify independent physiological predictions for seagrass associated with global change.     

Evaluation of aboveground and belowground biomass recovery in physically disturbed seagrass beds

Several studies addressed aboveground biomass recovery in tropical and subtropical seagrass systems following physical disturbance. However, there are few studies documenting belowground biomass recovery despite the important functional and ecological role of roots and rhizomes for seagrass ecosystems. In this study, we compared the recovery of biomass (g dry weight m(-2)) as well as the biomass recovery rates in ten severely disturbed multi-species seagrass meadows, after the sediments were excavated and the seagrasses removed. Three sites were located in the tropics (Puerto Rico) and seven in the subtropics (Florida Keys), and all were originally dominated by Thalassia testudinum. Total aboveground biomass reached reference values at four out of ten sites studied, two in the Florida Keys and two in Puerto Rico. Total belowground biomass was lower at the disturbed locations compared to the references at all sites, apart from two sites in the Florida Keys where the compensatory effect of opportunistic species (Syringodium filiforme and Halodule wrightii) was observed. The results revealed large variation among sites in aboveground and belowground biomass for all species, with higher aboveground recovery than belowground for T. testudinum. Recovery rates for T. testudinum were highly variable across sites, but a general trend of faster aboveground than belowground recovery was observed. Equal rates between aboveground and belowground biomass were found for opportunistic species at several sites in the Florida Keys. These results indicate the importance of belowground biomass when assessing seagrass recovery and suggest that the appropriate metric to assess seagrass recovery should address belowground biomass as well as aboveground biomass in order to evaluate the full recovery of ecological services and functions performed by seagrasses. We point out regional differences in species composition and species shifts following severe disturbance events and discuss ecological implications of gap dynamics in multi-species seagrass meadows.     

Seagrass meadows mixed with calcareous algae have higher plant productivity and sedimentary blue carbon storage

Seagrass meadows capture and store large amounts of carbon in the sediment beneath, thereby serving as efficient sinks of atmospheric CO₂. Carbon sequestration levels may however differ greatly among meadows depending on, among other factors, the plant community composition. Tropical seagrass meadows are often intermixed with macroalgae, many of which are calcareous, which may compete with seagrass for nutrients, light, and space. While the photosynthetic CO₂ uptake by both seagrasses and calcareous algae may increase the overall calcification in the system (by increasing the calcium carbonate saturation state, Ω), the calcification process of calcareous algae may lead to a release of CO₂, thereby affecting both productivity and calcification, and eventually also the meadows’ carbon storage. This study estimated how plant productivity, CaCO₃ production, and sediment carbon levels were affected by plant community composition (seagrass and calcareous algae) in a tropical seagrass‐dominated embayment (Zanzibar, Tanzania). Overall, the patterns of variability in productivity differed between the plant types, with net areal biomass productivity being highest in meadows containing both seagrass and calcareous algae. Low and moderate densities of calcareous algae enhanced seagrass biomass growth, while the presence of seagrass reduced the productivity of calcareous algae but increased their CaCO₃ content. Sedimentary carbon levels were highest when seagrasses were mixed with low or moderate cover of calcareous algae. The findings show that plant community composition can be an important driver for ecosystem productivity and blue carbon sequestration.