Tropical bloom-forming mesoalgae Cladophoropsis sp. and Laurencia sp.-responses to ammonium enrichment and a simulated heatwave.

Algal blooms are increasing worldwide, driven by elevated nutrient inputs. However, it is still unknown how tropical benthic algae will respond to heatwaves, which are expected to be more frequent under global warming. In the present study, a multifactorial experiment was carried out to investigate the potential synergistic effects of increased ammonium inputs (25 µM, control at 2.5 µM) and a heatwave (31°C, control at 25°C) on the growth and physiology (e.g., ammonium uptake, nutrient assimilation, photosynthetic performance, and pigment concentrations) of two bloom-forming algal species, Cladophoropsis sp. and Laurencia sp. Both algae positively responded to elevated ammonium concentrations with higher growth and chlorophyll a and lutein concentrations. Increased temperature was generally a less important driver, interacting with elevated ammonium by decreasing the algaes' %N content and N:P ratios. Interestingly, this stress response was not captured by the photosynthetic yield (Fv/Fm) nor by the carbon assimilation (%C), which increased for both algae at higher temperatures. The negative effects of higher temperature were, however, buffered by nutrient inputs, showing an antagonistic response in the combined treatment for the concentration of VAZ (violaxanthin, antheraxanthin, zeaxanthin) and thalli growth. Ammonium uptake was initially higher for Cladophoropsis sp. and increased for Laurencia sp. over experimental time, showing an acclimation capacity even in a short time interval. This experiment shows that both algae benefited from increased ammonium pulses and were able to overcome the otherwise detrimental stress of increasingly emerging temperature anomalies, which provide them a strong competitive advantage and might support their further expansions in tropical marine systems.

A trait-based framework for seagrass ecology: Trends and prospects.

In the last three decades, quantitative approaches that rely on organism traits instead of taxonomy have advanced different fields of ecological research through establishing the mechanistic links between environmental drivers, functional traits, and ecosystem functions. A research subfield where trait-based approaches have been frequently used but poorly synthesized is the ecology of seagrasses; marine angiosperms that colonized the ocean 100M YA and today make up productive yet threatened coastal ecosystems globally. Here, we compiled a comprehensive trait-based response-effect framework (TBF) which builds on previous concepts and ideas, including the use of traits for the study of community assembly processes, from dispersal and response to abiotic and biotic factors, to ecosystem function and service provision. We then apply this framework to the global seagrass literature, using a systematic review to identify the strengths, gaps, and opportunities of the field. Seagrass trait research has mostly focused on the effect of environmental drivers on traits, i.e., "environmental filtering" (72%), whereas links between traits and functions are less common (26.9%). Despite the richness of trait-based data available, concepts related to TBFs are rare in the seagrass literature (15% of studies), including the relative importance of neutral and niche assembly processes, or the influence of trait dominance or complementarity in ecosystem function provision. These knowledge gaps indicate ample potential for further research, highlighting the need to understand the links between the unique traits of seagrasses and the ecosystem services they provide.

Limited trait responses of a tropical seagrass to the combination of increasing pCO2 and warming.

Understanding species-specific trait responses under future global change scenarios is of importance for conservation efforts and to make informed decisions within management projects. The combined and single effects of seawater acidification and warmer average temperature were investigated by means of the trait responses of Cymodocea serrulata, a tropical seagrass, under experimental conditions. After a 35 d exposure period, biochemical, morphological, and photo-physiological trait responses were measured. Overall, biochemical traits mildly responded under the individual exposure to high temperature and increasing pCO2 values. The response of C. serrulata was limited to a decrease in %C and an increase in the sucrose content in the rhizome under the high temperature treatment, 32 °C. This suggests that this temperature was lower than the maximum tolerance limit for this species. Increasing pCO2 levels increased %C in the rhizome, and also showed a significant increase in leaf δ13C values. The effects of all treatments were sublethal; however, small changes in their traits could affect the ecosystem services they provide. In particular, changes in tissue carbon concentrations may affect carbon storage capacity, one key ecosystem service. The simultaneous study of different types of trait responses contributes to establish a holistic framework of seagrass ecosystem health under climate change.

Current advances in seagrass research: A review from Viet Nam.

Seagrass meadows provide valuable ecosystem services but are fragile and threatened ecosystems all over the world. This review highlights the current advances in seagrass research from Viet Nam. One goal is to support decision makers in developing science-based conservation strategies. In recent years, several techniques were applied to estimate the size of seagrass meadows. Independent from the method used, there is an alarming decline in the seagrass area in almost all parts of Viet Nam. Since 1990, a decline of 46.5% or 13,549 ha was found. Only in a few protected and difficult-to-reach areas was an increase observed. Conditions at those sites could be investigated in more detail to make suggestions for conservation and recovery of seagrass meadows. Due to their lifestyle and morphology, seagrasses take up compounds from their environment easily. Phytoremediation processes of Thalassia hemprichii and Enhalus acoroides are described exemplarily. High accumulation of heavy metals dependent on their concentration in the environment in different organs can be observed. On the one hand, seagrasses play a role in phytoremediation processes in polluted areas; on the other hand, they might suffer at high concentrations, and pollution will contribute to their overall decline. Compared with the neighboring countries, the total C org stock from seagrass beds in Viet Nam was much lower than in the Philippines and Indonesia but higher than that of Malaysia and Myanmar. Due to an exceptionally long latitudinal coastline of 3,260 km covering cool to warm water environments, the seagrass species composition in Viet Nam shows a high diversity and a high plasticity within species boundaries. This leads to challenges in taxonomic issues, especially with the Halophila genus, which can be better deduced from genetic diversity/population structures of members of Hydrocharitaceae. Finally, the current seagrass conservation and management efforts in Viet Nam are presented and discussed. Only decisions based on the interdisciplinary cooperation of scientists from all disciplines mentioned will finally lead to conserve this valuable ecosystem for mankind and biodiversity.

Local Victory: Assessing Interspecific Competition in Seagrass From a Trait-Based Perspective.

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.

Nutrient History Affects the Response and Resilience of the Tropical Seagrass Halophila stipulacea to Further Enrichment in Its Native Habitat.

Eutrophication is one of the main threats to seagrass meadows, but there is limited knowledge on the interactive effects of nutrients under a changing climate, particularly for tropical seagrass species. This study aimed to detect the onset of stress in the tropical seagrass, Halophila stipulacea, by investigating the effect of in situ nutrient addition during an unusually warm summer over a 6-month period. We measured a suite of different morphological and biochemical community metrics and individual plant traits from two different sites with contrasting levels of eutrophication history before and after in situ fertilization in the Gulf of Aqaba. Nutrient stress combined with summer temperatures that surpassed the threshold for optimal growth negatively affected seagrass plants from South Beach (SB), an oligotrophic marine protected area, while H. stipulacea populations from North Beach (NB), a eutrophic and anthropogenically impacted area, benefited from the additional nutrient input. Lower aboveground (AG) and belowground (BG) biomass, reduced Leaf Area Index (LAI), smaller internodal distances, high sexual reproductive effort and the increasing occurrence of apical shoots in seagrasses from SB sites indicated that the plants were under stress and not growing under optimal conditions. Moreover, AG and BG biomass and internodal distances decreased further with the addition of fertilizer in SB sites. Results presented here highlight the fact that H. stipulacea is one of the most tolerant and plastic seagrass species. Our study further demonstrates that the effects of fertilization differ significantly between meadows that are growing exposed to different levels of anthropogenic pressures. Thus, the meadow's "history" affects it resilience and response to further stress. Our results suggest that monitoring efforts on H. stipulacea populations in its native range should focus especially on carbohydrate reserves in leaves and rhizomes, LAI, internodal length and percentage of apical shoots as suitable warning indicators for nutrient stress in this seagrass species to minimize future impacts on these valuable ecosystems.

Unusually Warm Summer Temperatures Exacerbate Population and Plant Level Response of Posidonia oceanica to Anthropogenic Nutrient Stress.

Posidonia oceanica is a key foundation species in the Mediterranean providing valuable ecosystem services. However, this species is particularly vulnerable towards high coastal nutrient inputs and the rising frequency of intense summer heat waves, but their combined effect in situ has received little attention so far. Here, we investigated the effects of in situ nutrient addition during an unusually warm summer over a 4-month period, comparing different morphological, physiological and biochemical population metrics of seagrass meadows growing in protected areas (Ischia) with meadows already exposed to significant anthropogenic pressure (Baia - Gulf of Pozzuoli). Our study highlights that the effects of warmer than usual summer temperatures on the population level of seagrass meadows can be exacerbated if the plants are already exposed to higher anthropogenic pressures. Morphological and population level indicators mainly changed over time, possibly impacted by season and the warmer temperatures, and displayed more pronounced reductions in seagrasses from impacted sites. The additional nutrient supply had even more deleterious effects, as shown by a decrease in approximately 67% in cover in fertilized plots at high impacted sites and 33% at low impacted sites. Moreover, while rhizome starch concentration showed a seasonal increase in plants from low impacted sites it displayed a trend of a 27% decrease in fertilized plots of the high impacted sites. Epiphyte biomass was approximately four-fold higher on leaves of plants growing in impacted sites and even doubled with the additional nutrient input. Predicting and anticipating stress in P. oceanica is of crucial importance for conservation and management efforts, given the limited colonizing and reproductive ability and extremely slow growth of this ecosystem engineer. Our results suggest that monitoring efforts should focus especially on leaf area index (LAI), carbohydrate concentrations in the rhizomes, and epiphyte cover on leaves as indicators of the onset of stress in Posidonia oceanica, which can be used by decision makers to take appropriate measures before damage to the ecosystem becomes irreversible, minimize future human interference and strengthen the resilience of these important ecosystems.

Projected Rapid Habitat Expansion of Tropical Seagrass Species in the Mediterranean Sea as Climate Change Progresses.

During the last 150 years, the tropical seagrass species Halophila stipulacea has established itself in the southern and eastern parts of the Mediterranean Sea. More recently (2018), Halophila decipiens was observed for the first time in the eastern Mediterranean, and was described as the second non-native seagrass species in the Mediterranean Sea. We implemented a species distribution model (SDM) approach to (1) hindcast the habitat suitability of H. stipulacea over the last 100 years in the Mediterranean basin, and (2) to model the increase in the potential habitat suitability of H. stipulacea and H. decipiens during the current century under two very different climate scenarios, RCP 2.6 (lowest carbon emission scenario) and RCP 8.5 (highest carbon emission scenario). In addition, a principal component analysis (PCA) and k-means cluster based on temperature and salinity drivers were applied to visualize the distance and relatedness between the native and invasive H. stipulacea and H. decipiens populations. Results from this PCA suggest that the H. stipulacea populations of the Mediterranean and Red Sea are likely to be similar. In contrast, H. decipiens from the Mediterranean is more related to the Atlantic populations rather than to the Red Sea populations. The hindcast model suggests that the expansion of H. stipulacea was related to the increases in seawater temperatures in the Mediterranean over the last 100 years. The SDMs predict that more suitable habitat will become available for both tropical species during this century. The habitat suitability for H. stipulacea will keep expanding westward and northward as the Mediterranean continues to become saltier and warmer. In comparison, the SDMs built for H. decipiens forecast a restricted habitat suitability in the south-eastern Mediterranean Sea at the present environmental conditions and predicts a progressive expansion with a potential increase in habitat suitability along 85% of the Mediterranean coastline. The predicted rapid expansion of non-native seagrass species could alter the Mediterranean's seagrass community and may entail massive impacts on associated ecosystem functions and services, impacts that have severe socio-economic consequences.

Species-Specific Trait Responses of Three Tropical Seagrasses to Multiple Stressors: The Case of Increasing Temperature and Nutrient Enrichment.

Seagrass meadows are declining globally. The decrease of seagrass area is influenced by the simultaneous occurrence of many factors at the local and global scale, including nutrient enrichment and climate change. This study aims to find out how increasing temperature and nutrient enrichment affect the morphological, biochemical and physiological responses of three coexisting tropical species, Thalassia hemprichii, Cymodocea serrulata and Halophila stipulacea. To achieve these aims, a 1-month experiment under laboratory conditions combining two temperature (maximum ambient temperature and current average temperature) and two nutrient (high and low N and P concentrations) treatments was conducted. The results showed that the seagrasses were differentially affected by all treatments depending on their life-history strategies. Under higher temperature treatments, C. serrulata showed photo-acclimation strategies, while T. hemprichii showed decreased photo-physiological performance. In contrast, T. hemprichii was resistant to nutrient over-enrichment, showing enhanced nutrient content and physiological changes, but C. serrulata suffered BG nutrient loss. The limited response of H. stipulacea to nutrient enrichment or high temperature suggests that this seagrass is a tolerant species that may have a dormancy state with lower photosynthetic performance and smaller-size individuals. Interaction between both factors was limited and generally showed antagonistic effects only on morphological and biochemical traits, but not on physiological traits. These results highlight the different effects and strategies co-inhabiting seagrasses have in response to environmental changes, showing winners and losers of a climate change scenario that may eventually cause biodiversity loss. Trait responses to these stressors could potentially make the seagrasses weaker to cope with following events, due to BG biomass or nutrient loss. This is of importance as biodiversity loss in tropical seagrass ecosystems could change the overall effectiveness of ecosystem functions and services provided by the seagrass meadows.

Different strategies of nitrogen acquisition in two tropical seagrasses under nitrogen enrichment.

Tropical marine seagrasses live in environments with low nutrient concentrations. However, as land development intensifies along tropical coastlines, the marine environment in which these organisms grow is becoming more nutrient-rich. Nitrogen (N) uptake, assimilation, translocation and storage under a diversity of N sources in enriched conditions were investigated in two tropical seagrass species, Cymodocea serrulata and Thalassia hemprichii, from an oligotrophic marine environment. Both seagrasses were able to take up different inorganic and organic N sources through their above- and belowground tissues when enriched with high N concentrations. The uptake rates of T. hemprichii were generally higher than C. serrulata in leaves and rhizome, whereas root uptake was systematically higher in C. serrulata. Acropetal and basipetal translocation was observed in both species. Reduction and assimilation of N, measured in terms of their nitrate reductase and glutamine synthetase activity, were correlated with nitrate and ammonium uptake rates, respectively. Cymodocea serrulata showed a tendency to immediately use the available N, whereas T. hemprichii allocated more N in assimilation and storage investment. The responses of these seagrasses to N-enrichment demonstrate their ability to adapt to over-enrichment by varying N sources in the first step of the eutrophication process.

Bacterial Community Composition and Potential Driving Factors in Different Reef Habitats of the Spermonde Archipelago, Indonesia.

Coastal eutrophication is a key driver of shifts in bacterial communities on coral reefs. With fringing and patch reefs at varying distances from the coast the Spermonde Archipelago in southern Sulawesi, Indonesia offers ideal conditions to study the effects of coastal eutrophication along a spatially defined gradient. The present study investigated bacterial community composition of three coral reef habitats: the water column, sediments, and mucus of the hard coral genus Fungia, along that cross-shelf environmental and water quality gradient. The main research questions were: (1) How do water quality and bacterial community composition change along a coastal shelf gradient? (2) Which water quality parameters influence bacterial community composition? (3) Is there a difference in bacterial community composition among the investigated habitats? For this purpose, a range of key water parameters were measured at eight stations in distances from 2 to 55 km from urban Makassar. This was supplemented by sampling of bacterial communities of important microbial habitats using 454 pyrosequencing. Findings revealed that the population center Makassar had a strong effect on the concentrations of Chlorophyll a, suspended particulate matter (SPM), and transparent exopolymer particles (TEP), which were all significantly elevated at the inshore compared the other seven sites. Shifts in the bacterial communities were specific to each sampled habitat. Two OTUs, belonging to the genera Escherichia/Shigella (Gammaproteobacteria) and Ralstonia (Betaproteobacteria), respectively, both dominated the bacterial community composition of the both size fractions of the water column and coral mucus. The sampled reef sediments were more diverse, and no single OTUs was dominant. There was no gradual shift in bacterial classes or OTUs within the sampled habitats. In addition, we observed very distinct communities between the investigated habitats. Our data show strong changes in the bacterial community composition at the inshore site for water column and sediment samples. Alarmingly, there was generally a high prevalence of potentially pathogenic bacteria across the entire gradient.

Effect of Inorganic and Organic Carbon Enrichments (DIC and DOC) on the Photosynthesis and Calcification Rates of Two Calcifying Green Algae from a Caribbean Reef Lagoon.

Coral reefs worldwide are affected by increasing dissolved inorganic carbon (DIC) and organic carbon (DOC) concentrations due to ocean acidification (OA) and coastal eutrophication. These two stressors can occur simultaneously, particularly in near-shore reef environments with increasing anthropogenic pressure. However, experimental studies on how elevated DIC and DOC interact are scarce and fundamental to understanding potential synergistic effects and foreseeing future changes in coral reef function. Using an open mesocosm experiment, the present study investigated the impact of elevated DIC (pHNBS: 8.2 and 7.8; pCO2: 377 and 1076 µatm) and DOC (added as 833 µmol L-1 of glucose) on calcification and photosynthesis rates of two common calcifying green algae, Halimeda incrassata and Udotea flabellum, in a shallow reef environment. Our results revealed that under elevated DIC, algal photosynthesis decreased similarly for both species, but calcification was more affected in H. incrassata, which also showed carbonate dissolution rates. Elevated DOC reduced photosynthesis and calcification rates in H. incrassata, while in U. flabellum photosynthesis was unaffected and thalus calcification was severely impaired. The combined treatment showed an antagonistic effect of elevated DIC and DOC on the photosynthesis and calcification rates of H. incrassata, and an additive effect in U. flabellum. We conclude that the dominant sand dweller H. incrassata is more negatively affected by both DIC and DOC enrichments, but that their impact could be mitigated when they occur simultaneously. In contrast, U. flabellum can be less affected in coastal eutrophic waters by elevated DIC, but its contribution to reef carbonate sediment production could be further reduced. Accordingly, while the capacity of environmental eutrophication to exacerbate the impact of OA on algal-derived carbonate sand production seems to be species-specific, significant reductions can be expected under future OA scenarios, with important consequences for beach erosion and coastal sediment dynamics.

Non-Random Variability in Functional Composition of Coral Reef Fish Communities along an Environmental Gradient.

Changes in the coral reef complex can affect predator-prey relationships, resource availability and niche utilisation in the associated fish community, which may be reflected in decreased stability of the functional traits present in a community. This is because particular traits may be favoured by a changing environment, or by habitat degradation. Furthermore, other traits can be selected against because degradation can relax the association between fishes and benthic habitat. We characterised six important ecological traits for fish species occurring at seven sites across a disturbed coral reef archipelago in Indonesia, where reefs have been exposed to eutrophication and destructive fishing practices for decades. Functional diversity was assessed using two complementary indices (FRic and RaoQ) and correlated to important environmental factors (live coral cover and rugosity, representing local reef health, and distance from shore, representing a cross-shelf environmental gradient). Indices were examined for both a change in their mean, as well as temporal (short-term; hours) and spatial (cross-shelf) variability, to assess whether fish-habitat association became relaxed along with habitat degradation. Furthermore, variability in individual traits was examined to identify the traits that are most affected by habitat change. Increases in the general reef health indicators, live coral cover and rugosity (correlated with distance from the mainland), were associated with decreases in the variability of functional diversity and with community-level changes in the abundance of several traits (notably home range size, maximum length, microalgae, detritus and small invertebrate feeding and reproductive turnover). A decrease in coral cover increased variability of RaoQ while rugosity and distance both inversely affected variability of FRic; however, averages for these indices did not reveal patterns associated with the environment. These results suggest that increased degradation of coral reefs is associated with increased variability in fish community functional composition resulting from selective impacts on specific traits, thereby affecting the functional response of these communities to increasing perturbations.

The physiological response of two green calcifying algae from the Great Barrier Reef towards high dissolved inorganic and organic carbon (DIC and DOC) availability.

Increasing dissolved inorganic carbon (DIC) concentrations associated with ocean acidification can affect marine calcifiers, but local factors, such as high dissolved organic carbon (DOC) concentrations through sewage and algal blooms, may interact with this global factor. For calcifying green algae of the genus Halimeda, a key tropical carbonate producer that often occurs in coral reefs, no studies on these interactions have been reported. These data are however urgently needed to understand future carbonate production. Thus, we investigated the independent and combined effects of DIC (pCO2 402 µatm/ pHtot 8.0 and 996 µatm/ pHtot 7.7) and DOC (added as glucose in 0 and 294 µmol L-1) on growth, calcification and photosynthesis of H. macroloba and H. opuntia from the Great Barrier Reef in an incubation experiment over 16 days. High DIC concentrations significantly reduced dark calcification of H. opuntia by 130 % and led to net dissolution, but did not affect H. macroloba. High DOC concentrations significantly reduced daily oxygen production of H. opuntia and H. macroloba by 78 % and 43 %, respectively, and significantly reduced dark calcification of H. opuntia by 70%. Combined high DIC and DOC did not show any interactive effects for both algae, but revealed additive effects for H. opuntia where the combination of both factors reduced dark calcification by 162 % compared to controls. Such species-specific differences in treatment responses indicate H. opuntia is more susceptible to a combination of high DIC and DOC than H. macroloba. From an ecological perspective, results further suggest a reduction of primary production for Halimeda-dominated benthic reef communities under high DOC concentrations and additional decreases of carbonate accretion under elevated DIC concentrations, where H. opuntia dominates the benthic community. This may reduce biogenic carbonate sedimentation rates and hence the buffering capacity against further ocean acidification.

CO2 and inorganic nutrient enrichment affect the performance of a calcifying green alga and its noncalcifying epiphyte.

Ocean acidification studies in the past decade have greatly improved our knowledge of how calcifying organisms respond to increased surface ocean CO2 levels. It has become evident that, for many organisms, nutrient availability is an important factor that influences their physiological responses and competitive interactions with other species. Therefore, we tested how simulated ocean acidification and eutrophication (nitrate and phosphate enrichment) interact to affect the physiology and ecology of a calcifying chlorophyte macroalga (Halimeda opuntia (L.) J.V. Lamouroux) and its common noncalcifying epiphyte (Dictyota sp.) in a 4-week fully crossed multifactorial experiment. Inorganic nutrient enrichment (+NP) had a strong influence on all responses measured with the exception of net calcification. Elevated CO2 alone significantly decreased electron transport rates of the photosynthetic apparatus and resulted in phosphorus limitation in both species, but had no effect on oxygen production or respiration. The combination of CO2 and +NP significantly increased electron transport rates in both species. While +NP alone stimulated H. opuntia growth rates, Dictyota growth was significantly stimulated by nutrient enrichment only at elevated CO2, which led to the highest biomass ratios of Dictyota to Halimeda. Our results suggest that inorganic nutrient enrichment alone stimulates several aspects of H. opuntia physiology, but nutrient enrichment at a CO2 concentration predicted for the end of the century benefits Dictyota sp. and hinders its calcifying basibiont H. opuntia.

The future of the northeast Atlantic benthic flora in a high CO2 world.

Seaweed and seagrass communities in the northeast Atlantic have been profoundly impacted by humans, and the rate of change is accelerating rapidly due to runaway CO2 emissions and mounting pressures on coastlines associated with human population growth and increased consumption of finite resources. Here, we predict how rapid warming and acidification are likely to affect benthic flora and coastal ecosystems of the northeast Atlantic in this century, based on global evidence from the literature as interpreted by the collective knowledge of the authorship. We predict that warming will kill off kelp forests in the south and that ocean acidification will remove maerl habitat in the north. Seagrasses will proliferate, and associated epiphytes switch from calcified algae to diatoms and filamentous species. Invasive species will thrive in niches liberated by loss of native species and spread via exponential development of artificial marine structures. Combined impacts of seawater warming, ocean acidification, and increased storminess may replace structurally diverse seaweed canopies, with associated calcified and noncalcified flora, with simple habitats dominated by noncalcified, turf-forming seaweeds.

Nutrient availability affects the response of the calcifying chlorophyte Halimeda opuntia (L.) J.V. Lamouroux to low pH.

Atmospheric carbon dioxide emissions cause a decrease in the pH and aragonite saturation state of surface ocean water. As a result, calcifying organisms are expected to suffer under future ocean conditions, but their physiological responses may depend on their nutrient status. Because many coral reefs experience high inorganic nutrient loads or seasonal changes in nutrient availability, reef organisms in localized areas will have to cope with elevated carbon dioxide and changes in inorganic nutrients. Halimeda opuntia is a dominant calcifying primary producer on coral reefs that contributes to coral reef accretion. Therefore, we investigated the carbon and nutrient balance of H. opuntia exposed to elevated carbon dioxide and inorganic nutrients. We measured tissue nitrogen, phosphorus and carbon content as well as the activity of enzymes involved in inorganic carbon uptake and nitrogen assimilation (external carbonic anhydrase and nitrate reductase, respectively). Inorganic carbon content was lower in algae exposed to high CO2, but calcification rates were not significantly affected by CO2 or inorganic nutrients. Organic carbon was positively correlated to external carbonic anhydrase activity, while inorganic carbon showed the opposite correlation. Carbon dioxide had a significant effect on tissue nitrogen and organic carbon content, while inorganic nutrients affected tissue phosphorus and N:P ratios. Nitrate reductase activity was highest in algae grown under elevated CO2 and inorganic nutrient conditions and lowest when phosphate was limiting. In general, we found that enzymatic responses were strongly influenced by nutrient availability, indicating its important role in dictating the local responses of the calcifying primary producer H. opuntia to ocean acidification.

Differences in urbanization and degree of marine influence are reflected in delta13C and delta15N of producers and consumers in seagrass habitats of Puerto Rico.

Couplings between land use and marine food webs in tropical systems are poorly understood. We compared land-sea coupling in seven sites around Puerto Rico, differing in the degree of precipitation and urbanization, by measuring delta(13)C and delta(15)N in producers and consumers. delta(15)N values were influenced by human activity: the food web from sites near urbanized centers was on average 1 per thousand heavier in delta(15)N compared to undeveloped sites. This is most likely due to wastewater inputs from septic systems relatively near the shoreline. Changes in delta(13)C were best explained by differences in the degree of marine influence. Where terrestrial inputs from a major river dominated, delta(13)C values were lighter, whereas sites further from land and in locations exposed to oceanic currents had heavier delta(13)C values, characteristic of a marine source of dissolved organic carbon. We found no significant effect of precipitation on connectivity in spite of a twofold difference in annual average rainfall between the north and south coast. The results suggest there is some connectivity between land and sea in Puerto Rico, despite high rates of evaporation relative to precipitation.