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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.

Farming of Eucheuma denticulatum is a major activity in Zanzibar affecting seagrass ecosystems primarily through shading and trampling. The aim of this study was to test the impacts of shading and trampling during seaweed farming on seagrass meadows composed by Halophila stipulacea and Thalassia hemprichii and their associated benthic macroalgae. Areas covered by these species were selected for the building of seaweed farms in three treatments: seaweed farm plots (with shading and trampling effects), trampling plots (with trampling effects only), and control plots (with no shading or trampling effects). Reduction of light within the plots was recorded over 9 weeks. Percentage cover of seagrasses and macroalgae and shoot density of seagrasses were measured over 12 weeks to assess the impact of shading and trampling by seaweed farming activities. Light was significantly reduced in the seaweed farm plots by 75 to 90% by the end of a seaweed growth cycle. H. stipulacea, despite its capacity for rapid growth, was significantly affected by the combination of shading and trampling under the seaweed farm treatment, while the climax seagrass species T. hemprichii was unaffected. Due to the decline in H. stipulacea, benthic macroalgae cover increased in the seaweed farm treatment, suggesting a change in seagrass community dynamics. In contrast, trampling had a negative effect on the benthic macroalgae as an isolated disturbance, which suggests that seagrasses are more resistant to trampling than macroalgae and would likely dominate the benthic macrophyte community under these conditions.

Increasing temperature and CO2 concentration are among the most important factors affecting marine ecosystems under climate change. We investigated the morphological, biochemical, and physiological trait responses of seedlings of the tropical seagrass Enhalus acoroides under experimental conditions. Trait responses were greater under temperature effects than increasing CO2 concentration. Seedlings under rising temperatures showed enhanced leaf growth, lower leaf nutrient content, and stimulated down-regulating mechanisms in terms of photo-physiology. Increasing CO2 concentrations did not show any significant effects independently. There was a significant interaction for some of the trait responses considered, such as leaf number and carbon content in the roots, and trends of higher starch concentrations in the leaves and lower rETRmax under combined enriched CO2 and high temperature, even though none of these interactions were synergistic. Understanding the single and interactive trait responses of seagrass seedlings to increasing temperature and CO2 concentration is of importance to determine the relative responses of early life stages of seagrasses, which may differ from adult plants, in order to form a more holistic view of seagrass ecosystem health under climate change.

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.

Seagrass meadows are declining globally. In Indonesia, 75% loss has been reported in the last 5 years. The decrease of the 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 Enhalus acoroides in the seedling phase, which has not previously been studied. To achieve these aims, a laboratory experiment of combined temperature and nutrient treatments was conducted using recently-germinated seedlings of E. acoroides. The results showed that the seedlings were tolerant to an extended exposure to the current ambient maximum temperature. Under higher temperature treatment, the seedlings were observed to increase in aboveground size traits (e.g., number of leaves, leaf length, biomass and leaf area), as well as in belowground traits, such as root length. The results in this study also showed that the initial seed size matters for morphological responses. On the contrary, nutrient responses of seedlings were practically absent, suggesting they could rely on internal reserves. Interaction between both factors was limited, with the exception of low temperature and high nutrient treatment, in which the AG:BG ratio and leaf elongation rate increased. Fluorescence parameters were not influenced by any of the water treatments. The results in this study suggest that E. acoroides seedlings rely energetically in the reserves within the seedling and that increasing temperature might result in faster seedling development, although no interactions with other organisms were tested. This is of importance when studying the resilience capacity of this species and when restoration attempts are planned, as a faster root development would provide a faster stabilization in the sediment and the survival of the whole plant.

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.

The Euromarine Network funded the initial workshop that led to the development of this manuscript. AM-S was funded by the Leibniz Association as part of the project SEATRAIT and MT was supported by the German Research Foundation (DFG) within the individual Grants Program, project SEAMAC (TE 1046/3-1). AIS was funded by Portuguese national funds through the FCT - Foundation for Science and Technology, I.P., under the project CEECIND/00962/2017 and by FCT/MCTES through the financial support to CESAM (UIDB/50017/2020+UIDP/50017/2020 +LA/P/0094/2020). This study received Portuguese national funds from FCT - Foundation for Science and Technology through projects UIDB/04326/2020, UIDP/04326/2020 and LA/P/0101/2020 of CCMAR. IO was funded by the EU-H2020-MSCA grant n° 752250 (SEAMET). CB was funded by the Åbo Akademi University Foundation Sr and LM was funded by the doctoral network Functional Marine Biodiversity (FunMarBio) at Åbo Akademi University. RS acknowledges the support of the European Union’s Horizon 2020 Research and Innovation Program under the Grant Agreement Number 857251. GP, ED and MR were partially supported by the project Marine Hazard, PON03PE_00203_1 (MUR, Italy); MR was partially supported by a SZN PhD fellowship via the Open University.

Increasing temperature and CO2 concentration are among the most important factors affecting marine ecosystems under climate change. We investigated the morphological, biochemical, and physiological trait responses of seedlings of the tropical seagrass Enhalus acoroides under experimental conditions. Trait responses were greater under temperature effects than increasing CO2 concentration. Seedlings under rising temperatures showed enhanced leaf growth, lower leaf nutrient content, and stimulated down-regulating mechanisms in terms of photo-physiology. Increasing CO2 concentrations did not show any significant effects independently. There was a significant interaction for some of the trait responses considered, such as leaf number and carbon content in the roots, and trends of higher starch concentrations in the leaves and lower rETRmax under combined enriched CO2 and high temperature, even though none of these interactions were synergistic. Understanding the single and interactive trait responses of seagrass seedlings to increasing temperature and CO2 concentration is of importance to determine the relative responses of early life stages of seagrasses, which may differ from adult plants, in order to form a more holistic view of seagrass ecosystem health under climate change.