In agreement with global patterns of climate change and increasing temperatures in the tropical oceans, the Northern Red Sea (NRS) has been warming over the last few decades. Using 18 years of remotely-sensed chlorophyll-a data (Chl-a, an index of phytoplankton biomass), we investigate the potential impacts of climate warming on phytoplankton abundance and phenology in the Northern Red Sea by exploring the mechanistic links with the regional physical environment. The results of the analysis reveal that, in accordance with other tropical ecosystems, phytoplankton biomass in the NRS will decrease in response to warmer climate scenarios. This is attributed to lower heat fluxes (heat loss to the atmosphere) during the bloom period, and enhanced vertical stratification, which prevents vertical mixing of nutrients into the euphotic layer. In addition, we show that during warmer conditions (when heat fluxes are weakened), the winter bloom initiates significantly later (by up to 10 weeks) and its duration is considerably reduced. The biological implications of alterations to phytoplankton phenology may include increased larval mortality of pelagic species, reduced recruitment, fisheries impacts and changes to community structure.

Aquatic primary productivity mainly depends on pelagic phytoplankton. The globally abundant marine picocyanobacteria Prochlorococcus comprises a significant fraction of the photosynthetic biomass in most tropical, oligotrophic oceans. The Red Sea is an enclosed narrow body of water characterized by continuous solar irradiance, and negligible annual rainfall, in addition to elevated temperatures and salinity levels, which mimics a global warming scenario. Analysis of 16S rRNA sequences of bacterioplankton communities indicated the predominance of a high-light adapted ecotype (HL II) of Prochlorococcus at the surface of the Northern and Central Red Sea. To this end, we analyzed the distribution of Prochlorococcus at multiple depths within and below the euphotic zone in different regions of the Red Sea, using clone libraries of the 16S–23S rRNA internal transcribed spacer (ITS) region. Results indicated a high diversity of Prochlorococcus ecotypes at the 100 m depth in the water column and an unusual dominance of HL II-related sequences in deeper waters of the Red Sea. To further investigate the microdiversity of Prochlorococcus over a wider biogeographical scope, we used a 454-pyrosequencing approach to analyze rpoC1 gene pyrotags. Samples were collected from the surface of the water column to up to 500 m at 45 stations that span the Red Sea’s main basin from 4 north to south. Phylogenetic analysis of abundant rpoC1 OTUs revealed genotypes of recently discovered strains that belong to the high-light and lowlight clades. In addition, we used a rapid community-profiling tool (GraftM) and quantitatively analyzed rpoC1 gene abundance from 45 metagenomes to assess the Prochlorococcus community structure across vertical and horizontal physicochemical gradients. Results revealed the clustering of samples according to their depth and a strong influence on ecotypic distribution by temperature and oxygen levels. In efforts to better understand how the cells survive the unusual features of the Red Sea, a Prochlorococcus strain of the HL II adapted clade from the euphotic zone was cultured, enabling morphological analyses and growth rates measurements for the strain. In addition, we successfully sequenced and annotated the genome of the strain, which was then used for genomic comparison with other ecotypes. Interestingly, the set of unique genes identified in the draft genome included genes encoding proteins involved in salt tolerance mechanisms. The expression level and pattern of these genes in the Red Sea water column was explored through metatranscriptomic mapping and revealed their occurrence throughout, independent of the diel cycle. This led to the hypothesis that Prochlorococcus populations in the highly saline Red Sea are able to biosynthesize additional compatible solutes via several pathways to counterbalance the effects of salt stress. The results presented in this dissertation provide the first glimpse on how the environmental parameters of the Red Sea can affect the evolution, diversity and distribution patterns of Prochlorococcus ecotypes.

Satellite sea surface height (SSH) data over 1992-2012 are analyzed to study the spatial and temporal variability of sea level in the Red Sea. Empirical orthogonal functions (EOF) analysis suggests the remarkable seasonality of SSH in the Red Sea, and a significant correlation is found between SSH variation and seasonal wind cycle. A winding-angle based eddy identification algorithm is employed to derive the mesoscale eddy information from SSH data. Totally more than 5500 eddies are detected, belonging to 2583 eddy tracks. Statistics suggest that eddies generate over the entire Red Sea, with two regions in the central basin of high eddy frequency. 76% of the detected eddies have a radius ranging from 40km to 100km, of which both intensity and absolute vorticity decrease with eddy radius. The average eddy lifespan is about 5 weeks, and eddies with longer lifespan tend to have larger radius but less intensity. Different deformation rate exists between anticyclonic eddies (AEs) and cyclonic eddies (CEs), those eddies with higher intensity appear to be less deformed and more circular. Inspection of the 84 long-lived eddies suggests the AEs tend to move a little more northward than CEs. AE generation during summer is obviously lower than that during other seasons, while CE generation is higher during spring and summer. Other features of AEs and CEs are similar with both vorticity and intensity reaching the summer peaks in August and winter peaks in January. Inter-annual variability reveals that the eddies in the Red Sea are isolated from the global event. The eddy property tendencies are different from the south and north basin, both of which exhibit a two-year cycle. Showing a correlation coefficient of -0.91, Brunt–Väisälä frequency is negatively correlated with eddy kinetic energy (EKE), which results from AE activities in the high eddy frequency region. Climatological vertical velocity shear variation is identical with EKE except in the autumn, suggesting the vertical shear could convert the energy from baroclinic instability into eddy activity. Finally, numerical simulation results from the MIT general circulation model (MITgcm) are validated with previous studies and observations. The vertical structure of the simulated flux through Bab el Mandeb is successfully reproduced. Further validation with the 2010 cruise suggests that the thermocline occurs at ~200m, but the model vertical salinity gradient is lower than the observations. The model surface eddy variability is also examined, suggesting good agreement with satellite observations.

Eukaryotic microalgae from the Red Sea were isolated, characterized and identified with the purpose of building a culture collection that will serve future research activities in the area of industrial microbiology. Seven sampling locations were surveyed using an in-house designed isolation protocol. Microalgae enrichment was carried out in vitro using the streak plate method and fluorescence activated cell sorting approaches. Colonial and cellular microscopy, growth media preference assays, as well as temperature, pH and salinity tolerance tests were carried out to describe the isolates. DNA extraction, PCR amplification, template sequencing and in silico analyses were carried out to identify the isolates and arrange them in a proper phylogenetic description. In total, 129 isolates were obtained. From these, only 39 were selected for characterization given their increased ability of accumulating large amounts of biomass in solid and liquid media in relatively short periods of time. All of these have a green color, are unicellular, non-motile, photosynthetic organisms and have a cell size ranging from 5 to 8 µm. More than half of them showed growth preference in Walne media, followed by F/2, MN and BG-11 SW. Maximum temperature tolerance of all organisms was around 38 ºC, while optimum growth was observed close to 25 ºC. pH preference was diverse and three groups were identified: acidic (6), intermediate (8 - 9) and alkaline (> 10) growing isolates. Salinity tests showed an overall growth preference at 25 PSU, approximately 10 units lower than that found at the sampling stations. Most isolates showed diminished growth at high salinity and high pH, except for OS3S1b which grew well in both cases, and could be an interesting strain to study further. Twenty four isolates were related to Ulvophyceae sp. MBIC10591 by BLAST approaches with a maximum identity of 96 - 97%. A maximum likelihood phylogenetic tree was created for these isolates, relative to the BLAST hits and to some model eukaryotic microalgae for positioning reference. It was shown that the 24 OS isolates are related to each other with a confidence value of 84%. Differential responses of all high biomass producing isolates towards abiotic stresses might suggest that each represents a distinct, novel, unclassified marine organism.

The overall goal of this study was to elucidate the relationship between coral nutrition and the observed prey preferences exhibited by corallivorous butterflyfishes. Fifteen species of coral (thirteen hard, two soft) and stomach/hindgut contents from six species of butterflyfish were analyzed in this study, all collected from the central Saudi Arabian Red Sea. All samples were analyzed for lipid, total-nitrogen (proxy for protein), and ash (proxy for minerals and when combined with lipid data, allows for calculation of carbohydrate). Unfortunately, substantial errors were encountered in the experimental lipid data, precluding the use of this data set. Using the value of (protein/ash) as a proxy for potential nutritional quality, it was determined that Pocillopora cf. verrucosa and P. damicornis have the highest nutritional quality, while Acropora hyacinthus and Stylophora pistillata have intermediate nutritional quality, and all remaining 11 species have low nutritional quality. This suggests that the high nutritional quality of Pocillopora damicornis and Acropora hyacinthus may be the cause of the well documented predator preferences for these two species. Fish gut content samples were, on average, twice as rich in protein and half as rich in minerals as the coral tissue samples, suggesting either selective consumption of especially rich parts of the coral colony, or consumption of other food sources (facultative corallivores). In all six butterflyfish species, stomach content samples were consistently richer in protein and poorer in mineral content than the hindgut content samples; this suggests significant and measureable uptake of protein in the butterflyfish digestion process.

The Red Sea has long been recognized as a global hotspot of marine biodiversity. Ongoing overfishing, however, is threatening this unique ecosystem, recently leading to the identification of the Red Sea as one of three major hotspots of extinction risk for sharks and rays worldwide. Elasmobranch catches in Saudi Arabian Red Sea waters are unregulated, often misidentified and unrecorded, resulting in a lack of species-specific landings information, which would be vital for the formulation of effective management strategies. Here we employed an integrated approach of fisheries dependent and independent survey methods combined with molecular tools to provide biological, ecological and fisheries data to aid in the assessment of the status of elasmobranch populations in the Red Sea. Over the course of two years, we conducted market surveys at the biggest Saudi Arabian fish market in Jeddah. Market landings were dominated by, mostly immature individuals - implying both recruitment and growth overfishing. Additionally, we employed baited remote underwater video (BRUVS) and longline surveys along almost the entire length of the Red Sea coast of Saudi Arabia as well as at selected reef systems in Sudan. The comparison of catch per unit effort (CPUE) data for Saudi Arabian Red Sea BRUVS and longline surveys to published data originating from non-Red Sea ocean systems revealed CPUE values several orders of magnitude lower for both survey methods in the Red Sea compared to other locations around the world. Finally, we infered the regional population structure of four commercially important shark species between the Red Sea and the Western Indian Ocean.We genotyped nearly 2000 individuals at the mitochondrial control region as well as a total of 20 microsatellite loci. Genetic homogeneity could not be rejected for any of the four species across the spatial comparison. Based on high levels of region-wide exploitation, we suggest that, for management purposes, the population structure of all four species should be considered as single stock in the three marginal seas surrounding Arabia. Overall, our combined results indicate a severe depletion of sharks in Saudi Arabian Red Sea waters, likely caused by drastic overfishing of elasmobranch populations.

Observing populations at different spatial scales gives greater insight into the specific processes driving genetic differentiation and population structure. Here we determined population connectivity across multiple spatial scales in the Red Sea to determine the population structures of two reef building corals Stylophora pistillata and Pocillopora verrucosa. The Red sea is a 2,250 km long body of water with extremely variable latitudinal environmental gradients. Mitochondrial and microsatellite markers were used to determine distinct lineages and to look for genetic differentiation among sampling sites. No distinctive population structure across the latitudinal gradient was discovered within this study suggesting a phenotypic plasticity of both these species to various environments. Stylophora pistillata displayed a heterogeneous distribution of three distinct genetic populations on both a fine and large scale. Fst, Gst, and Dest were all significant (p-value<0.05) and showed moderate genetic differentiation between all sampling sites. However this seems to be byproduct of the heterogeneous distribution, as no distinct genetic population breaks were found. Stylophora pistillata showed greater population structure on a fine scale suggesting genetic selection based on fine scale environmental variations. However, further environmental and oceanographic data is needed to make more inferences on this structure at small spatial scales. This study highlights the deficits of knowledge of both the Red Sea and coral plasticity in regards to local environmental conditions.

Phytoplankton is at the basis of the marine food chain and therefore play a fundamental role in the ocean ecosystem. However, the large-scale phytoplankton dynamics of the Red Sea are not well understood yet, mainly due to the lack of historical in situ measurements. As a result, our knowledge in this area relies mostly on remotely-sensed observations and large-scale numerical marine ecosystem models. Models are very useful to identify the mechanisms driving the variations in chlorophyll concentration and have practical applications for fisheries operation and harmful algae blooms monitoring. Modelling approaches can be divided between physics- driven (dynamical) approaches, and data-driven (statistical) approaches. Dynamical models are based on a set of differential equations representing the transfer of energy and matter between different subsets of the biota, whereas statistical models identify relationships between variables based on statistical relations within the available data. The goal of this thesis is to develop, implement and test novel dynamical and statistical modelling approaches for studying and forecasting the variability of chlorophyll concentration in the Red Sea. These new models are evaluated in term of their ability to efficiently forecast and explain the regional chlorophyll variability. We also propose innovative synergistic strategies to combine data- and physics-driven approaches to further enhance chlorophyll forecasting capabilities and efficiency.

Coral reefs are among the most diverse marine ecosystems, but they are threatened by climate change. The foundation of reef ecosystems is the coral holobiont or metaorganism that consists of the coral animal host, photosynthetic microalgae, bacteria, and viruses (among other organisms). While microalgae provide the energy for corals to build the massive three-dimensional skeletons, bacteria support functions related to metabolism, immunity, and environmental adaptation. Conversely, the function of viruses is less well understood. Although viruses were previously associated with coral disease and bleaching, we are missing an overall understanding of the diversity and identity of viruses associated with corals, in particular for understudied areas such as the Red Sea. Here we characterized coral-associated viral community composition using a large metagenomic and metatransciptomic dataset covering > 1 billion sequences across > 100 coral samples collected from 14 different coral species in the central Red Sea. The viral sequence portion shows that coral species significantly differ from each other, but the most abundant viral families were consistently present. Notably, we found a pervasive abundance of Baculoviridae in metagenomes. In contrast, Polydnaviridae were the most abundant viruses in metatranscriptomes, highlighting that the combined approach of metagenomics and metatranscriptomics is informative with regard to deciphering viral diversity and activity. Our study provides a first comprehensive description of viruses associated with Red Sea corals. In line with previous studies, we confirm the presence of Baculoviridae, Polydnaviridae, Phycodnaviridae, Mimiviridae, and Herpesviridae, which may be considered viral families that are globally and commonly associated with corals. The reason for the pervasive abundance of Baculoviridae in Red Sea corals at present remains unknown, but it is tempting to speculate that the association is related to the uniquely warm and salty environment of the Red Sea.

A diversity of human-assisted approaches to rehabilitate and boost coral health have been suggested and investigated throughout the past years. Vitamins and trace-metal supplementation is a well-known strategy in human medicine and aquaculture, but vitamin addition is not currently actively tested for coral growth and recovery. These molecules are essential cofactors that have been correlated with coral thermal resistance and upregulated in corals treated with beneficial microorganisms (i.e., probiotics). To assess the effects of B12, B6 and zinc supplementation on coral health, we conducted a 2-month experiment in an open-closed-loop system mesocosm joined to a peristaltic pump continuously dosing the vitamins and/or zinc to individual 250 L tanks. Fragments of five different colonies of Acropora hemprichii were randomly distributed into the respective treatment tanks (B12, B6, zinc, multi-treatment and control). After 21 days, the corals were exposed to a pulse (1 day) of thermal stress, followed by three weeks of recovery. Substantial mortality (55%) in the control treatment was observed during the stress and recovery, with B12, B6, zinc and multi treatments exhibiting significantly less mortality (<20%). Coral health data combined with analysis of microbiome and metabolomic approaches suggest that both vitamins and zinc have a positive effect on coral health recovery. This study is the first to provide evidence that complex B-vitamins accompanied by zinc supplementation, can be a valuable tool for coral reef rehabilitation, and paves the way to further understanding specific mechanisms by which these nutrients promote coral health will be needed.