Opisthobranch molluscs of the Sudanese Red Sea

Eighteen opisthobranch species are described and illustrated from the Sudanese Red Sea, including seven new species, one new subspecies and three species for which this is the first Red Sea record. Eleven bullomorphs, four pleurobranchomorphs, four sacoglossans and four phyllidiid mudibranchs are now known from the Red Sea. There appears to be a high percentage of endemic species in the Red Sea. This is what one might expect, in view of the conditions of near isolation, low temperatures and high salinity that prevailed in this sea 13000 23000 years ago. Of the groups discussed in this paper, one Red Sea species has invaded the Mediterranean Sea, and one Mediterranean species has invaded the Red Sea, since the opening of the Suez Canal in 1867. Many Red Sea species are very colourful, in comparison to Mediterranean ones. This could perhaps be because Red Sea waters are cleaner, thereby favouring the evolution of greater precision in colour markings.     

Opisthobranch molluscs of the Sudanese Red Sea

Eighteen opisthobranch species are described and illustrated from the Sudanese Red Sea, including seven new species, one new subspecies and three species for which this is the first Red Sea record. Eleven bullomorphs, four pleurobranchomorphs, four sacoglossans and four phyllidiid mudibranchs are now known from the Red Sea.
There appears to be a high percentage of endemic species in the Red Sea. This is what one might expect, in view of the conditions of near isolation, low temperatures and high salinity that prevailed in this sea 13000 23000 years ago.
Of the groups discussed in this paper, one Red Sea species has invaded the Mediterranean Sea, and one Mediterranean species has invaded the Red Sea, since the opening of the Suez Canal in 1867.
Many Red Sea species are very colourful, in comparison to Mediterranean ones. This could perhaps be because Red Sea waters are cleaner, thereby favouring the evolution of greater precision in colour markings.     

The status of coral reef ecology research in the Red Sea

The Red Sea has long been recognized as a region of high biodiversity and endemism. Despite this diversity and early history of scientific work, our understanding of the ecology of coral reefs in the Red Sea has lagged behind that of other large coral reef systems. We carried out a quantitative assessment of ISI-listed research published from the Red Sea in eight specific topics (apex predators, connectivity, coral bleaching, coral reproductive biology, herbivory, marine protected areas, non-coral invertebrates and reef-associated bacteria) and compared the amount of research conducted in the Red Sea to that from Australia’s Great Barrier Reef (GBR) and the Caribbean. On average, for these eight topics, the Red Sea had 1/6th the amount of research compared to the GBR and about 1/8th the amount of the Caribbean. Further, more than 50 % of the published research from the Red Sea originated from the Gulf of Aqaba, a small area (<2 % of the area of the Red Sea) in the far northern Red Sea. We summarize the general state of knowledge in these eight topics and highlight the areas of future research priorities for the Red Sea region. Notably, data that could inform science-based management approaches are badly lacking in most Red Sea countries. The Red Sea, as a geologically “young” sea located in one of the warmest regions of the world, has the potential to provide insight into pressing topics such as speciation processes as well as the capacity of reef systems and organisms to adapt to global climate change. As one of the world’s most biodiverse coral reef regions, the Red Sea may yet have a significant role to play in our understanding of coral reef ecology at a global scale.     

A review of elasmobranch research in the Red Sea

Given the global concern about the status of elasmobranch fishes, the paucity of information on elasmobranchs in the Red Sea is worrisome. Management of elasmobranchs in areas other than the Red Sea has been helped by research on population ecology, reproductive biology and resource partitioning, subjects that are virtually absent in the Red Sea elasmobranch literature. This review provides the first comprehensive summary of elasmobranch biology in the Red Sea with the aim of facilitating research in a region that remains remarkably under-studied.     

rDNA analysis of the Red Sea seagrass,Halophila, reveals vicariant evolutionary diversification

The effects of opening the Suez Canal as a connection between the Red Sea and the Mediterranean Sea were reported for a number of marine species. However, the evolutionary origin of the seagrasses in the Red Sea and the linking population genetics of seagrasses between the Arabian Sea, the Gulf of Aden, the Red Sea and the Mediterranean Sea have not yet been investigated in detail. The invasion of Halophila stipulacea Asch. from the Red Sea into the Mediterranean Sea after the opening of the Suez Canal was already recorded. We hypothesize that Halophila ovalis populations in the Red Sea developed through long-term historical processes such as vicariant evolutionary diversification. Seagrass samples were collected along the Egyptian coastline of the Red Sea and analysed by the molecular marker ITS. The sequences were compared with published ITS sequences from seagrasses collected in the whole area of interest. In this study, we reveal the linking population genetics, phylogeography and phylogenetics of two dominant seagrass species, Halophila stipulacea and Halophila ovalis, among species collected in the Red Sea and worldwide. The results indicate that the Red Sea Halophila ovalis populations do not group to Halophila ovalis worldwide, and Halophila major, Halophila ovalis collected worldwide and Halophila ovalis collected at the Red Sea are sister clades. Hence, vicariant evolutionary diversification for Halophila ovalis may occur in the Red Sea.     

Kudoa iwatai (Myxosporea: Multivalvulida) in wild and cultured fish in the Red Sea: redescription and molecular phylogeny

Gilt-head sea bream, Sparus aurata L., the Mediterranean's most important mariculture species, has been cultured for the last 30 yr in Eilat (Israeli Red Sea). Kudoa sp. was the first myxosporean parasite reported from this species. In recent years, an increase in prevalence in both land-based and sea-cage facilities in Eilat has been observed. Infections with the same Kudoa species appeared in cultured European sea bass Dicentrarchus labrax (L.) and grey mullet Mugil cephalus in the same farms, as well as in 10 species of wild Red Sea reef fish, indicating that Kudoa sp. is not fastidious with regard to its host. All affected species displayed 1- to 2-mm (up to 5 mm) whitish, spherical, or oval polysporous plasmodia. The parasite established multiple site infections, most commonly in the muscles and intracranial adipose tissue of the brain and eye periphery. Other sites were subcutaneous adipose tissue, nerve axons, mouth, eye, mesenteries, peritoneum, swim bladder, intestinal musculature, heart, pericardium, kidney, and ovary. On the basis of spore morphology, the parasite was identified as Kudoa iwatai Egusa and Shiomitsu, 1983. Ultrastructural features were comparable to those of previously studied Kudoa species. The 18S rDNA from 7 Red Sea isolates was sequenced and compared with the sequence of the same gene from K. iwatai isolated from cultured red sea bream, Pagrus major, in Japan. The phylogenetic position of K. iwatai within the genus was determined using sequence analysis of all related taxa available in GenBank. The 3 isolates of K. iwatai clustered together on a newly formed, highly supported clade. The Red Sea strain of K. iwatai is apparently native to the region. In the absence of records of this Kudoa sp. from the extensive Mediterranean sea bream and sea bass production industries, introduction with its Mediterranean hosts seems unlikely. Therefore, we conclude that K. iwatai is an Indo-Pacific species that, in the Red Sea, has extended its host range to include the allochthonous gilt-head sea bream, European sea bass, and grey mullet.     

Thermal refugia against coral bleaching throughout the northern Red Sea

Tropical reefs have been impacted by thermal anomalies caused by global warming that induced coral bleaching and mortality events globally. However, there have only been very few recordings of bleaching within the Red Sea despite covering a latitudinal range of 15° and consequently it has been considered a region that is less sensitive to thermal anomalies. We therefore examined historical patterns of sea surface temperature (SST) and associated anomalies (1982–2012) and compared warming trends with a unique compilation of corresponding coral bleaching records from throughout the region. These data indicated that the northern Red Sea has not experienced mass bleaching despite intensive Degree Heating Weeks (DHW) of >15°C‐weeks. Severe bleaching was restricted to the central and southern Red Sea where DHWs have been more frequent, but far less intense (DHWs <4°C‐weeks). A similar pattern was observed during the 2015–2016 El Niño event during which time corals in the northern Red Sea did not bleach despite high thermal stress (i.e. DHWs >8°C‐weeks), and bleaching was restricted to the central and southern Red Sea despite the lower thermal stress (DHWs < 8°C‐weeks). Heat stress assays carried out in the northern (Hurghada) and central (Thuwal) Red Sea on four key reef‐building species confirmed different regional thermal susceptibility, and that central Red Sea corals are more sensitive to thermal anomalies as compared to those from the north. Together, our data demonstrate that corals in the northern Red Sea have a much higher heat tolerance than their prevailing temperature regime would suggest. In contrast, corals from the central Red Sea are close to their thermal limits, which closely match the maximum annual water temperatures. The northern Red Sea harbours reef‐building corals that live well below their bleaching thresholds and thus we propose that the region represents a thermal refuge of global importance.     

Demographic structure and stock status of Lethrinus lentjan in Saudi coastal waters of the Red Sea

Fishes of the family Lethrinidae form a considerable portion of the catch from both the Red Sea and the Arabian Gulf in Saudi Arabia, and the species Lethrinus lentjan (Lacepède, 1802) is one of the most important among these fishes. This study was conducted to evaluate the demographic structure coefficients, survival rate, and stock status of L. lentjan from the Red Sea coast of Saudi Arabia. A total of 593 samples were collected on a monthly basis for a period of one year from the landing site for fishing boats operating in Red Sea waters off Jeddah. The results indicated that 88.87% of the specimens were female; the maximum total length recorded was 43.5 cm with the most frequent length (14.67%) being 23–23.9 cm. The maximum age recorded for both males and females was 6 year-plus, and the 1 year-plus age category represented the majority of the samples (57.67%). The total mortality coefficient, natural mortality coefficient, and fishing mortality coefficient were 1.538, 0.315, and 1.223, respectively; all mortality coefficients in the female fishes were higher than those in the male fishes. The survival rate of males was higher (0.617) than that of females (0.214). The results of the present study indicate that L. lentjan is subjected to overfishing and a new management strategy is necessary to improve the stock status of this fish species.     

Distribution of some planktonic foraminifera species in deep-sea cores from the red sea and their relation to eustatic sea-level changes

Distribution patterns of planktonic foraminifera in four sediment cores from the Red Sea are studied. The most common species are Globigerinoides ruber, G. sacculifer, Globigerinella siphonifera and Orbulina universa. G. ruber and G. sacculifer show opposite trends of distribution in the sediment cores. Abundance of the foraminifera during the glacial periods suggests that the connection of the Red Sea to the Indian ocean was not completely interrupted and the salinity conditions were not extreme.However, higher salinities appear to have existed in the northern Red Sea, where most of the planktonic foraminifera that occur in the southern Red Sea are absent. It is inferred that the salinity in the southern Red Sea during the glacial period was less than 50%, whereas higher salinity might have existed in the north where the influence of the Indian Ocean was minimal.     

Being green in the Red Sea - the photosymbiotic ascidian Diplosoma simile (Ascidiacea: Didemnidae) in the Gulf of Aqaba

The photosymbiotic ascidian Diplosoma simile is characterized by a wide global distribution. The current study provides the first record of D. simile in the Red Sea, with remarks on its spatial distribution and phylogeny. Partial COI and 18S rDNA sequences of D. simile from the Red Sea revealed high similarity to those from the West Pacific, indicating a much broader range than previously recorded. In the Red Sea D. simile is common on natural coral reefs at depths ranging from 1–35 m. Diplosoma simile colonies were found to overgrow and would appear to suffocate several species of live stony corals, though this does not appear to cause mass coral mortality in the Red Sea, unlike that recorded in other locations.     

First record of the paedomorphic fish Schindleria (Gobioidei, Schindleriidae) from the Red Sea

A male specimen of the paedomorphic fish Schindleria sp. was collected by plankton net, in February 2005, from a reef lagoon from Hurghada on the Egyptian coast of the northern Red Sea. This is the first record of the Schindleriidae in the Red Sea.     

A coral reef refuge in the Red Sea

The stability and persistence of coral reefs in the decades to come is uncertain due to global warming and repeated bleaching events that will lead to reduced resilience of these ecological and socio‐economically important ecosystems. Identifying key refugia is potentially important for future conservation actions. We suggest that the Gulf of Aqaba (GoA) (Red Sea) may serve as a reef refugium due to a unique suite of environmental conditions. Our hypothesis is based on experimental detection of an exceptionally high bleaching threshold of northern Red Sea corals and on the potential dispersal of coral planulae larvae through a selective thermal barrier estimated using an ocean model. We propose that millennia of natural selection in the form of a thermal barrier at the southernmost end of the Red Sea have selected coral genotypes that are less susceptible to thermal stress in the northern Red Sea, delaying bleaching events in the GoA by at least a century.     

Impact of late Quaternary environmental changes on deep-sea benthic foraminiferal faunas of the Red Sea

Fluctuations in abundance, diversity and species composition of benthic foraminifera from two sites of the northern and southern Red Sea indicate strong variability of deep-sea ecosystems during the last four glacial to interglacial cycles. In total, five and four different benthic foraminiferal assemblages have been identified in the northern core and southern core, respectively. Comparison with recent faunas from the Red Sea and adjacent oceans allowed the reconstruction of temporal changes in deep-water ventilation, salinity and food availability at the seafloor. Generally, the abundance of infaunal and miliolid taxa increase during glacial intervals indicating increased organic matter fluxes, oxygen decrease and salinity increase in deep waters during these times. These fluctuations are attributed to enhanced oxygen consumption rates and temporarily reduced deep-water formation in the northern Red Sea during glacial intervals. The recorded environmental changes are a reflection of both high- and low-latitude climate changes. The northern Red Sea is mainly influenced by glacio-eustatic sea level fluctuations that control deep-water formation rates and by mid-latitude climate changes of the Mediterranean region that control surface productivity. In contrast, deep-sea ecosystem variability of the southern Red Sea is additionally influenced by low-latitude climate changes attributed to the NE monsoon intensity that drives the inflow of nutrient-rich surface waters from the Gulf of Aden. These results demonstrate the high sensitivity of deep-sea ecosystems of the Red Sea to both global and regional climate changes.     

Marine heatwaves reveal coral reef zones susceptible to bleaching in the Red Sea

As the Earth's temperature continues to rise, coral bleaching events become more frequent. Some of the most affected reef ecosystems are located in poorly monitored waters, and thus, the extent of the damage is unknown. We propose the use of marine heatwaves (MHWs) as a new approach for detecting coral reef zones susceptible to bleaching, using the Red Sea as a model system. Red Sea corals are exceptionally heat‐resistant, yet bleaching events have increased in frequency. By applying a strict definition of MHWs on >30 year satellite‐derived sea surface temperature observations (1985–2015), we provide an atlas of MHW hotspots over the Red Sea coral reef zones, which includes all MHWs that caused major coral bleaching. We found that: (a) if tuned to a specific set of conditions, MHWs identify all areas where coral bleaching has previously been reported; (b) those conditions extended farther and occurred more often than bleaching was reported; and (c) an emergent pattern of extreme warming events is evident in the northern Red Sea (since 1998), a region until now thought to be a thermal refuge for corals. We argue that bleaching in the Red Sea may be vastly underrepresented. Additionally, although northern Red Sea corals exhibit remarkably high thermal resistance, the rapidly rising incidence of MHWs of high intensity indicates this region may not remain a thermal refuge much longer. As our regionally tuned MHW algorithm was capable of isolating all extreme warming events that have led to documented coral bleaching in the Red Sea, we propose that this approach could be used to reveal bleaching‐prone regions in other data‐limited tropical regions. It may thus prove a highly valuable tool for policymakers to optimize the sustainable management of coastal economic zones.     

First record of the blackfin coral goby,Paragobiodon lacunicolus (Kendall and Goldsborough, ), from the Red Sea

Three specimens of the blackfin coral goby, Paragobiodon lacunicolus, were collected in 2010 and 2011 at different locations of the eastern Red Sea along the coast of Saudi Arabia. Reexamination of 14 museum specimens from Eritrea also confirmed the presence of the species for the western Red Sea. This represents the first published report of P. lacunicolus from the Red Sea.     

Cladocera from the Sudan: Red Sea Hills,Jebel Marra and valley of the main Nile

Twenty species of Cladocera are reported from the Nile, where lacustrine species dominate, and from Jebel Marra and the Red Sea Hills, where chydorids dominate. The community found in the Red Sea Hills is more typically desertic than that of Jebel Marra, which appears closely related to the fauna of the West and Central African Sahel.     

Genetic population structure of the endemic fourline wrasse (Larabicus quadrilineatus) suggests limited larval dispersal distances in the Red Sea

The connectivity among marine populations is determined by the dispersal capabilities of adults as well as their eggs and larvae. Dispersal distances and directions have a profound effect on gene flow and genetic differentiation within species. Genetic homogeneity over large areas is a common feature of coral reef fishes and can reflect high dispersal capability resulting in high levels of gene flow. If fish larvae return to their parental reef, gene flow would be restricted and genetic differentiation could occur. Larabicus quadrilineatus (Labridae) is considered as an endemic fish species of the Red Sea and Gulf of Aden. The juveniles of this species are cleaner fish that feed on ectoparasites of other fishes. Here, we investigated the genetic population structure and gene flow in L. quadrilineatus among five locations in the Red Sea to infer connectivity among them. To estimate genetic diversity, we analysed 369 bp of 237 mitochondrial DNA control region sequences. Haplotype and nucleotide diversities were higher in the southern than in the northern Red Sea. Analysis of molecular variance (amova) detected the highest significant genetic variation between northern and central/southern populations (Phi(CT) = 0.01; P < 0.001). Migration analysis revealed a several fold higher northward than southward migration, which could be explained by oceanographic conditions and spawning season. Even though the Phi(ST) value of 0.01 is rather low and implies a long larval dispersal distance, estimates based on the isolation-by-distance model show a very low mean larval dispersal distance (0.44-5.1 km) compared to other studies. In order to enable a sustainable ornamental fishery on the fourline wrasse, the results of this study suggest that populations in the northern and southern Red Sea should be managed separately as two different stocks. The rather low larval dispersal distance of about 5 km needs to be considered in the design of marine protected areas to enable connectivity and self-seeding.     

Absence of post-Miocene Red Sea land bridges: biogeographic implications

In a large number of studies concerned with species movements between Africa and Eurasia, including the migrations of hominids out of Africa, a frequently‐cited dispersal route is across a hypothetical land bridge in the southern Red Sea, which is suggested to have emerged during glacial sea‐level lowstands. This paper, however, unequivocally demonstrates that palaeoceanographic and palaeoecological data are incompatible with the existence of Red Sea land bridges since the Miocene. The case is made by presenting the first quantitative history of water depth above the Red Sea sill for the last 470,000 years, a time period that includes the four most recent glacial–interglacial cycles, and by discussing the predictable consequences of any land bridge formation on the Red Sea sedimentary and microfossil records. The absence of post‐Miocene Red Sea land bridges has extensive implications for biogeographic models in the Afro‐Arabian region. Genetic, morphometric and palaeontological patterns reported in the literature cannot be related to dispersals over a land bridge, or in the case of marine organisms, separation of the Red Sea from the Indian Ocean by a land bridge. If such patterns in terrestrial species are only congruent with a southern Red Sea dispersal route, then they need to be considered in terms of sweepstake rafting, anthropogenic introduction, or in the particular case of the Out‐of‐Africa migration by modern humans, seafaring. The constraints imposed by our palaeoenvironmental record on biogeographic reconstructions within and around the Red Sea will hopefully encourage both the review of previous works and the preference for multidisciplinary approaches in future studies.