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IntroductionSeagrasses form oxidizing microenvironments around their roots, creating complex and strong redox gradients, thus affecting the rates of microbial carbon mineralization in their surrounding sediments. Since seagrasses are continuously being lost worldwide, a deeper understanding of the changes that occur within different seagrass sediments following the disappearance of the plants is of ecological and global importance.MethodsWe conducted a slurry experiment with sediments that have different characteristics from the northern tip of Gulf of Aqaba; the different sediments included different compartments of the tropical seagrass Halophila stipulacea (old and young leaves, rhizomes, or roots). We measured the changes over time in dissolved inorganic carbon (DIC), alkalinity, ferrous iron (Fe2+), hydrogen sulfide (H2S), sulfate (SO42-), and sulphur isotope ratios in sulfate within water. These measurements were used to calculate the rate of remineralization of each seagrass compartment, allowing us to predict the potential effects of the disappearance of different H. stipulacea compartments on key microbial processes in the surrounding environment.ResultsWe show that H. stipulacea’s rhizomes had the fastest decomposition rates, followed by the young leaves, roots, and old leaves (which also indicates the preservation potential of old leaves).DiscussionHigh concentrations of hydrogen sulfide were detected only in the slurries containing rhizomes and young leaves. High sulfide concentrations can lead to seagrass mortality and cause a positive feedback loop where the loss of seagrass due to sulfide generates further sulfide accumulation. This positive feedback loop can also be further reinforced by the loss of burrowing fauna in the sediment. This emphasizes the importance of understanding the extent of different pathways of seagrass disappearance on the surrounding environment and other geochemical feedbacks.

IntroductionSoil water availability is a key factor in the growth of trees. In arid deserts, tree growth is limited by very dry soil and atmosphere conditions. Acacia tree species are distributed in the most arid deserts of the globe, therefore they are well adapted to heat and long droughts. Understanding why some plants do better than others in some environments is a key question in plant science.MethodsHere we conducted a greenhouse experiment to continuously and simultaneously track the whole-plant water-balance of two desert Acacia species, in order to unravel their physiological responses to low water availability.ResultsWe found that even under volumetric water content (VWC) of 5-9% in the soil, both species maintained 25% of the control plants, with a peak of canopy activity at noon. Moreover, plants exposed to the low water availability treatment continued growing in this period. A. tortilis applied a more opportunistic strategy than A. raddiana, and showed stomatal responses at a lower VWC (9.8% vs. 13.1%, t4= -4.23, p = 0.006), 2.2-fold higher growth, and faster recovery from drought stress.DiscussionAlthough the experiment was done in milder VPD (~3 kPa) compared to the natural conditions in the field (~5 kPa), the different physiological responses to drought between the two species might explain their different topographic distributions. A. tortilis is more abundant in elevated locations with larger fluctuations in water availability while A. raddiana is more abundant in the main channels with higher and less fluctuating water availability. This work shows a unique and non-trivial water-spending strategy in two Acacia species adapted to hyper-arid conditions.

Abstract Background Halophila stipulacea seagrass meadows are an ecologically important and threatened component of the ecosystem in the Gulf of Aqaba. Recent studies have demonstrated correlated geographic patterns for leaf epiphytic community composition and leaf morphology, also coinciding with different levels of water turbidity and nutrient concentrations. Based on these observations, workers have suggested an environmental microbial fingerprint, which may reflect various environmental stress factors seagrasses have experienced, and may add a holobiont level of plasticity to seagrasses, assisting their acclimation to changing environments and through range expansion. However, it is difficult to tease apart environmental effects from host-diversity dependent effects, which have covaried in field studies, although this is required in order to establish that differences in microbial community compositions among sites are driven by environmental conditions rather than by features governed by the host. Results In this study we carried out a mesocosm experiment, in which we studied the effects of warming and nutrient stress on the composition of epiphytic bacterial communities and on some phenological traits. We studied H. stipulacea collected from two different meadows in the Gulf of Aqaba, representing differences in the host and the environment alike. We found that the source site from which seagrasses were collected was the major factor governing seagrass phenology, although heat increased shoot mortality and nutrient loading delayed new shoot emergence. Bacterial diversity, however, mostly depended on the environmental conditions. The most prominent pattern was the increase in Rhodobacteraceae under nutrient stress without heat stress, along with an increase in Microtrichaceae. Together, the two taxa have the potential to maintain nitrate reduction followed by an anammox process, which can together buffer the increase in nutrient concentrations across the leaf surface. Conclusions Our results thus corroborate the existence of environmental microbial fingerprints, which are independent from the host diversity, and support the notion of a holobiont level plasticity, both important to understand and monitor H. stipulacea ecology under the changing climate.

The Gulf of Aqaba (GoA; northern Red Sea) supports extensive seagrass meadows, dominated by the small tropical seagrass species, Halophila stipulacea. Due to its semi-closed structure, in the GoA, regional nutrient loading and global warming are considered the biggest threats to local seagrass meadows, and their combination can potentially amplify their negative impacts. Using a seagrass-dedicated mesocosm, we exposed two seagrass populations with different local “nutrient history” to control (27 °C) and simulated warming (31 °C), with and without nutrients (20 µM DIN). Following four weeks of these treatments (“stress phase”), all plants were returned to control conditions (“recovery phase”) for another three weeks. Results showed that exposure to only thermal stress favoured growth, compared to exposure to only nutrient increase that reduced Fv/Fm and growth but favoured algae proliferation. Exposure to the combined thermal and nutrient stress, negatively affected seagrass performance resulting in high mortality observed after four weeks of combined exposure. The negative effects of combined stressors were stronger in populations with low “nutrient history”. Additionally, we propose two novel fatty acid (FA) biomarkers, one based on FA unsaturation, 16:3n-3/16:2n-6, and the other on FA elongation processes, 18:2n-6/16:2n-6. Fatty acid analyses showed a significant decrease in 16:3n-3/16:2n-6 and 18:3n-3/18:2n-6 with increases in temperature and nutrients; a more drastic decline was found under the interaction of both stressors. Our results point out that C16 PUFAs, that are synthesized within the “prokaryotic pathway”, are more sensitive to thermal and the combined thermal + nutrients stressors than C18 PUFAs, which are synthesized within the “eukaryotic pathway”. In general, following a month of control conditions, a clear recovery of most of the seagrass descriptors was observed, highlighting the great capability of Halophila stipulacea to recover from stress conditions. Our results have important ecological and management implications to the seagrass meadows in the GoA and elsewhere. For seagrasses to survive climate change, managers must put efforts into limiting other stressors such as eutrophication that would potentially reduce the seagrass resilience to climate change.

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.

Background: The evolutionary relationships between plants and their microbiomes are of high importance to the survival of plants in general and even more in extreme conditions. Changes in the plant's microbiome can affect plant development, growth, fitness, and health. Along the arid Arava, southern Israel, acacia trees (Acacia raddiana and Acacia tortilis) are considered keystone species. In this study, we investigated the ecological effects of plant species, microclimate, phenology, and seasonality on the epiphytic and endophytic microbiome of acacia trees. One hundred thirty-nine leaf samples were collected throughout the sampling year and were assessed using 16S rDNA gene amplified with five different primers (targeting different gene regions) and sequenced (150 bp paired-end) on an Illumina MiSeq sequencing platform.Results: Epiphytic bacterial diversity indices (Shannon–Wiener, Chao1, Simpson, and observed number of operational taxonomic units) were found to be nearly double compared to endophyte counterparts. Epiphyte and endophyte communities were significantly different from each other in terms of the composition of the microbial associations. Interestingly, the epiphytic bacterial diversity was similar in the two acacia species, but the canopy sides and sample months exhibited different diversity, whereas the endophytic bacterial communities were different in the two acacia species but similar throughout the year. Abiotic factors, such as air temperature and precipitation, were shown to significantly affect both epiphyte and endophytes communities. Bacterial community compositions showed that Firmicutes dominate A. raddiana, and Proteobacteria dominate A. tortilis; these bacterial communities consisted of only a small number of bacterial families, mainly Bacillaceae and Comamonadaceae in the endophyte for A. raddiana and A. tortilis, respectively, and Geodematophilaceae and Micrococcaceae for epiphyte bacterial communities, respectively. Interestingly, ~60% of the obtained bacterial classifications were unclassified below family level (i.e., “new”).Conclusions: These results shed light on the unique desert phyllosphere microbiome highlighting the importance of multiple genotypic and abiotic factors in shaping the epiphytic and endophytic microbial communities. This study also shows that only a few bacterial families dominate both epiphyte and endophyte communities, highlighting the importance of climate change (precipitation, air temperature, and humidity) in affecting arid land ecosystems where acacia trees are considered keystone species.

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.

Halophila stipulacea is a small tropical seagrass, native to the Red Sea, Persian Gulf, and the Indian Ocean. It invaded the Mediterranean Sea 150 years ago as a Lessepsian migrant, but so far has remained in insulated, small populations across this basin. Surprisingly, in 2002 it was reported in the Caribbean Sea, where within less than two decades it spread to most of the Caribbean Island nations and reaching the South American continent. Unlike its invasion of Mediterranean, in the Caribbean H. stipulacea creates large, continuous populations in many areas. Reports from the Caribbean demonstrated the invasiveness of H. stipulacea by showing that it displaces local Caribbean seagrass species. The motivation for this review comes from the necessity to unify the existing knowledge on several aspects of this species in its native and invasive habitats, identify knowledge gaps and develop a critical strategy to understand its invasive capacity and implement an effective monitoring and conservation plan to mitigate its potential spread outside its native ranges. We systematically reviewed 164 studies related to H. stipulacea to create the “Halophila stipulacea database.” This allowed us to evaluate the current biological, ecological, physiological, biochemical, and molecular knowledge of H. stipulacea in its native and invasive ranges. Here we (i) discuss the possible environmental conditions and plant mechanisms involved in its invasiveness, (ii) assess the impact of H. stipulacea on native seagrasses and ecosystem functions in the invaded regions, (iii) predict the ability of this species to invade European and transoceanic coastal waters, (iv) identify knowledge gaps that should be addressed to better understand the biology and ecology of this species both in its native and non-native habitats, which would improve our ability to predict H. stipulacea's potential to expand into new areas in the future. Considering the predicted climate change scenarios and exponential human pressures on coastal areas, we stress the need for coordinated global monitoring and mapping efforts that will record changes in H. stipulacea and its associated communities over time, across its native, invasive and prospective distributional ranges. This will require the involvement of biologists, ecologists, economists, modelers, managers, and local stakeholders.

Climate change fuels invasions of plant species and displacement of local plants. Little is known about the ecophysiological adaptation of the invasive species, and their ability to cope with the changing conditions in their new habitat. Halophila stipulacea, a tropical seagrass native to the Gulf of Aqaba (GoA; northern Red Sea), became a Lessepsian migrant spreading within the eastern Mediterranean where it could potentially outcompete local species. We analyzed temperature records in the last 35 years and show that water temperature has increased faster in the eastern Mediterranean Sea compared to GoA, suggesting that H. stipulacea’s invasive success is associated with adaptation to thermal warming. Furthermore, we compared the responses of native (Eilat, Israel) and invasive (Limassol, Cyprus) H. stipulacea plants to current (26°C) and predicted thermal maxima (29 and 32°C) in a controlled experimental microcosm. Morphological and photo-physiological results showed negative effects of heat stress on the native plants while un-affected/or even enhanced performance in their invasive counterparts. Gene expression, studied for the 1st time in H. stipulacea, pointed to differences in the molecular responses of two populations to thermal stress. Results predict that sea warming will cause vast reductions in H. stipulacea meadows growing in the GoA while it will facilitate H. stipulacea’s spread within the Mediterranean Sea.

Cumulative human impacts have led to the degradation of marine ecosystems and the decline of biodiversity in the European and contiguous seas. Effective conservation measures are urgently needed to reverse these trends. Conservation must entail societal choices, underpinned by human values and worldviews that differ between the countries bordering these seas. Social, economic and political heterogeneity adds to the challenge of balancing conservation with sustainable use of the seas. Comprehensive macro-regional coordination is needed to ensure effective conservation of marine ecosystems and biodiversity of this region. Under the European Union Horizon 2020 framework programme, the MarCons COST action aims to promote collaborative research to support marine management, conservation planning and policy development. This will be achieved by developing novel methods and tools to close knowledge gaps and advance marine conservation science. This action will provide support for the development of macro-regional and national policies through six key actions: to develop tools to analyse cumulative human impacts; to identify critical scientific and technical gaps in conservation efforts; to improve the resilience of the marine environment to global change and biological invasions; to develop frameworks for integrated conservation planning across terrestrial, freshwater, and marine environments; to coordinate marine conservation policy across national boundaries; and to identify effective governance approaches for marine protected area management. Achieving the objectives of these actions will facilitate the integration of marine conservation policy into macro-regional maritime spatial planning agendas for the European and contiguous seas, thereby offsetting the loss of biodiversity and ecosystem services in this region.

Microbes and seagrass establish symbiotic relationships constituting a functional unit called the holobiont that reacts as a whole to environmental changes. Recent studies have shown that the seagrass microbial associated community varies according to host species, environmental conditions and the host’s health status, suggesting that the microbial communities respond rapidly to environmental disturbances and changes. These changes, dynamics of which are still far from being clear, could represent a sensitive monitoring tool and ecological indicator to detect early stages of seagrass stress. In this review, the state of art on seagrass holobiont is discussed in this perspective, with the aim of disentangling the influence of different factors in shaping it. As an example, we expand on the widely studied Halophila stipulacea’s associated microbial community, highlighting the changing and the constant components of the associated microbes, in different environmental conditions. These studies represent a pivotal contribution to understanding the holobiont’s dynamics and variability pattern, and to the potential development of ecological/ecotoxicological indices. The influences of the host’s physiological and environmental status in changing the seagrass holobiont, alongside the bioinformatic tools for data analysis, are key topics that need to be deepened, in order to use the seagrass-microbial interactions as a source of ecological information.

The tropical seagrass species, Halophila stipulacea, originated from the Indian Ocean and the Red Sea, subsequently invading the Mediterranean and has recently established itself in the Caribbean Sea. Due to its invasive nature, there is growing interest in understanding this species’ capacity to adapt to new conditions. One approach to understanding the natural tolerance of a plant is to compare the tolerant species with a closely related non-tolerant species. We compared the physiological responses of H. stipulacea exposed to different salinities, with that of its nearest freshwater relative, Vallisneria americana. To achieve this goal, H. stipulacea and V. americana plants were grown in dedicated microcosms, and exposed to the following salt regimes: (i) H. stipulacea: control (40 PSU, practical salinity units), hyposalinity (25 PSU) and hypersalinity (60 PSU) for 3 weeks followed by a 4-week recovery phase (back to 40 PSU); (ii) V. americana: control (1 PSU), and hypersalinity (12 PSU) for 3 weeks, followed by a 4-week recovery phase (back to 1 PSU). In H. stipulacea, leaf number and chlorophyll content showed no significant differences between control plants and plants under hypo and hypersalinities, but a significant decrease in leaf area under hypersalinity was observed. In addition, compared with control plants, H. stipulacea plants exposed to hypo and hypersalinity were found to have reduced below-ground biomass and C/N ratios, suggesting changes in the allocation of resources in response to both stresses. There was no significant effect of hypo/hypersalinity on dark-adapted quantum yield of photosystem II (Fv/Fm) suggesting that H. stipulacea photochemistry is resilient to hypo/hypersalinity stress. In contrast to the seagrass, V. americana exposed to hypersalinity displayed significant decreases in above-ground biomass, shoot number, leaf number, blade length and Fv/Fm, followed by significant recoveries of all these parameters upon return of the plants to non-saline control conditions. These data suggest that H. stipulacea shows remarkable tolerance to both hypo and hypersalinity. Resilience to a relatively wide range of salinities may be one of the traits explaining the invasive nature of this species in the Mediterranean and Caribbean Seas.

Genetic diversity and structure of populations at the edge of the species' spatial distribution are important for potential adaptation to environmental changes and consequently, for the long-term survival of the species. Here, we combined classical population genetic methods with newly developed network analyses to gain complementary insights into the genetic structure and diversity of Acacia tortilis, a keystone desert tree, at the northern edge of its global distribution, where the population is under threat from climatic, ecological, and anthropogenic changes. We sampled A. tortilis from 14 sites along the Dead Sea region and the Arava Valley in Israel and in Jordan. In addition, we obtained samples from Egypt and Sudan, the hypothesized origin of the species. Samples from all sites were genotyped using six polymorphic microsatellite loci.Our results indicate a significant genetic structure in A. tortilis along the Arava Valley. This was detected at different hierarchical levels-from the basic unit of the subpopulation, corresponding to groups of trees within ephemeral rivers (wadis), to groups of subpopulations (communities) that are genetically more connected relative to others. The latter structure mostly corresponds to the partition of the major drainage basins in the area. Network analyses, combined with classical methods, allowed for the identification of key A. tortilis subpopulations in this region, characterized by their relatively high level of genetic diversity and centrality in maintaining gene flow in the population. Characterizing such key subpopulations may enable conservation managers to focus their efforts on certain subpopulations that might be particularly important for the population's long-term persistence, thus contributing to species conservation within its peripheral range.

Mass coral bleaching is increasing in frequency and severity, leading to the loss of coral abundance and diversity. However, some corals are less susceptible to bleaching than others and can provide a model for identifying the physiological and biogeochemical traits that underlie coral resilience to thermal stress. Corals from Eilat in the Gulf of Aqaba in the northern Red Sea do not bleach unless seawater temperatures are sustained at +6°C or higher above their average summer maximum. This extreme thermal tolerance qualifies these as super-corals, as most corals bleach when exposed to temperatures that are only +1–2°C above their thermal maximum. Here, we conducted a controlled bleaching experiment (+6°C) for 37 days (equivalent to 32° heating weeks) on three species of corals from Eilat: Stylophora pistillata, Pocillopora damicornis, and Favia favus. To assess the response of the holobiont to thermal stress, the following variables were measured on each coral: endosymbiotic algal cell density, Chlorophyll a, endosymbiotic mitotic cell division, total lipids, protein, carbohydrate, and the stable carbon (δ13C) and oxygen (δ18O) isotopic composition of the skeleton and the δ13C of the animal host tissue and endosymbiotic algae. While all three species appeared visibly bleached, their physiological and biogeochemical responses were species-specific. S. pistillata catabolized lipids but still maintained total energy reserves and biomass. Increases in both skeletal δ13C and δ18O indicates that calcification declined in this species. P. damicornis was the least affected by bleaching. It maintained its total energy reserves and biomass, and isotopic evidence suggests that it maintained calcification and was not dependent on heterotrophy for meeting metabolic demand when bleached. Finally, F. favus catabolized protein and carbohydrates, and suffered losses in total energy reserves and biomass. Nevertheless, isotopic evidence suggest that photosynthesis and calcification were maintained, and that this species has a high baseline heterotrophic capacity. Thus, just like their non-super-coral conspecific counterparts, maintaining energy reserves and biomass, and heterotrophic capacity appear to be traits that underlie the thermal tolerance of these super-corals from Eilat. Given the high thermal tolerance of these super-corals, these populations could provide viable seed stock for repopulating coral losses on other reefs.