Comparison of morphological and molecular traits for species identification and taxonomic grouping of oncaeid copepods

The pelagic marine copepod family Oncaeidae is highly diversified (over 100 species worldwide) and includes a great number of sibling species, which are difficult to identify morphologically, because of their very small size (0.18–1.2 mm total length as adults). Global investigations of oncaeid biodiversity are severely hampered by insufficient taxonomic knowledge, in particular for species which have first been described from the European Mediterranean Sea (type locality). Many of these species have been reported as key taxa of small-sized copepod communities in very distant oceanic regions. However, due to the taxonomic uncertainties it cannot be excluded that at least some of these allegedly cosmopolitan species in reality represent a complex of distinct, yet closely related, species. To improve the basis for the identification of Oncaeidae of Mediterranean origin, new diagnostic characters in combination with traditional methods were applied in the present study. Copepods were sampled with fine nets of 0.1 mm mesh size down to a maximum depth of 1,000 m on a west-east-transect in the Mediterranean Sea. Oncaeid species and form variants were predefined morphologically and the genetic identity of the morphospecies was analysed by about 650 and 500 bp region of the mitochondrial COI and 12S srRNA gene sequence, respectively (barcoding). A total of 67 individuals from 24 oncaeid species and forms were successfully analysed, including 12 species and one form of Mediterranean origin. For Oncaeidae, the 12S amplification turned out to be more successful (23 species) than the COI amplification (13 species and 1 form). Together, the morphological and molecular results are discussed with respect to three topics: (1) confirmation of a genetic distinction of three Triconia species, which have been interpreted as sibling species by morphological characters, (2) genetic distance of species within the ovalis-complex of oncaeids and (3) the taxonomic status of two form variants of Oncaea mediterranea (Claus).     

The determinants of species richness of a relatively young coral-reef ichthyofauna

Aim Unique topographic features left the Red Sea and its north‐eastern extension into the Gulf of Aqaba practically devoid of coral‐reef‐based organisms during the last glacial maximum. The current ichthyofauna in these two ‘regions’ thus represents the product of relatively recent colonization by species found in the Arabian Sea, which adjoins the Red Sea at its southern tip. We used this system to test why some marine species seemingly fail to extend their geographic range, thereby generating spatial heterogeneity in biodiversity. Location The Arabian Sea, Red Sea, and the Gulf of Aqaba. Methods A list of coral‐reef‐associated fish species, belonging to the 10 most speciose families, was compiled for each region using published sources. The data were analysed (major axis regression, randomization tests) for taxonomic and body‐size‐dependent biases in colonization probabilities. A simple probabilistic model was used to examine the potential contribution of local (within‐region) extinctions to determining species composition in the Red Sea. Results Of the 462 reef‐associated species that inhabit the Arabian Sea, 69% have crossed successfully into the Red Sea; of these, 55% have crossed into the Gulf of Aqaba. A species’ probability of being found in either ‘target’ was independent of presumed innate differences, i.e. ecological correlates of taxonomic affiliation and body size. Similarly, local extinctions were found unlikely to have been of consequence over the past several thousand years. Main conclusions Present‐day differences in the species richness of reef‐associated fish species among the Arabian Sea, Red Sea and Gulf of Aqaba appear to be the product of external, non‐selective constraints on colonization. The random nature of the colonization process is suggestive of ecological redundancy among coral‐reef fish species. Importantly, the study places a time frame on the processes that determine spatial patterns of biodiversity in reef fish.     

Untersuchungen zur Zoogeographie der Euphausiaceen (Crustacea) des Roten Meeres

A joint programme on “Biota of the Red Sea and the Eastern Mediterranean” has been carried out by the Hebrew University of Jerusalem and the Smithsonian Institution of Washington between 1967 and 1972 in order to study the role played by the Suez Canal as a route for faunal exchanges between the Red Sea and Eastern Mediterranean. The euphausiids of this material have been examined and additional samples of zooplankton have been taken in the Gulf of Elat (Aqaba) in 1973 in order to study the vertical distribution of the euphausiids. Four species have been found to occur in the Northern Red Sea and the Gulf of Elat:Euphausia diomedeae, E. sanzoi, Stylocheiron affine andS. abbreviatum. Seven species were found in the Eastern Mediterranean of whichEuphausia brevis is the dominant species. The present studies revealed that an exchange of Euphausiacea through the Suez Canal does not take place. The species of the Red Sea belong to the Indopacific fauna. Despite the particular historic-geographical and ecological situation of the Red Sea, no endemisms or subspecies have evolved there within the Euphausiacea.     

Quantitative Analyse von Korallengemeinschaften des Sanganeb-Atolls (mittleres Rotes Meer). II. Vergleich mit einem Riffareal bei Aqaba (nördliches Rotes Meer) am Nordrande des indopazifischen Riffgürtels

Quantitative studies of coral communities in the central and northern Red Sea were designed for comparison of the community structure in both areas. The central Red Sea provides reef-building Scleractinia and reef-inhabiting Alcyonaria with optimal temperature conditions, whereas the north tip of the Gulf of Aqaba (29°30 N) represents the northernmost outpost of coral reefs in the Indian Ocean. It is generally assumed that coral diversity decreases towards the margins of the global reef-belt. In the Red Sea, generic diversity of hermatypic Scleractinia slightly decreases from the central to the northern part (51 : 48 genera); but cnidarian species abundance (species number per 25 m² area) was found to increase from 62 to 98 species and the Shannon-Wiener diversity index increased from 2.58 to 3.67 with regard to colony number. The mean colony size was 189 cm² at Sanganeb-Atoll, but only 52 cm² at Aqaba. The mean numbers of colonies were inversely related: 662 per 25 m² at Sanganeb-Atoll and 2028 at Aqaba. Uninhabited parts of the studied areas amounted to 47 % at Sanganeb-Atoll and to 56 % at Aqaba. The community structure of the studied areas indicates that occasional perturbations prevent the progress of the community towards a low-diversity equilibrium state. Since severe hydrodynamic damage is extremely rare in 10 m depth, major disturbances may occur by sedimentation, by the interference of grazers (e. g.Diadema setosum) and due to overgrowth by space-competitors (mainly soft corals). These events are to be regarded as throwbacks in the process of monopolization of the area by well adapted species. Recovery from such perturbations (i.e. recolonization of dead areas) obviously takes place at different velocities in the northern and central Red Sea, for the mean water temperature at Aqaba is 5 °C lower than in the central Red Sea. Hence the process of taking over a given space by a few species proceeds further in the central Red Sea than at its northern end. The increase in diversity per area towards high latitudes is comparable to that with depth. It is concluded from the great number of species at Aqaba that these reefs mark the northernmost outpost of the Indian Ocean only geographically but not ecophysiologically; they would occur at even higher latitudes, if the Gulf of Aqaba extended farther north.     

Plagiopsetta biocellata,a new species of righteye flounder from the Gulf of Aqaba,northern Red Sea (Teleostei: Samaridae)

Ichthyological Research - The samarid flatfish Plagiopsetta biocellata sp. nov. is described from Egypt, Gulf of Aqaba, southern Red Sea, based on a specimen collected in the coral-reef lagoon at a...     

Evolution in Action-Peacock-Feather Like Supraocular Tentacles of the Lionfish, Pterois volitans – the Distribution of a New Signal

Synopsis Observations on populations of the lionfish, Pterois volitans (Scorpaenidae), in the Red Sea revealed that during the last four decades a novel phenotype of this species has started to evolve, which instead of the typical ray-like supraocular tentacles bears a pair of feather-like tentacles, one or both of which bear a sharply defined eye-mark, previously noted in this fish in the eastern Indian ocean. During the ongoing studies of the last 25 years it has become clear that this ornamentation, possibly first observed in the Red Sea at the southern tip of Sinai, had gradually extended both westward to the Egyptian shores, and eastward, along the 180 km of the Gulf of Aqaba. In the last decade such ornamented specimens have begun to appear and multiply at the northern tip of the Gulf, within the large population of ‘regular’ lionfish bearing ray-like tentacles. The evolutionary basis for this development, and the possible usefulness of this novel signal are discussed.     

Taxonomy of Oncaeidae (Copepoda,Poecilostomatoida) from the Red Sea. III. Morphology and phylogenetic position of Oncaea subtilis Giesbrecht, 1892

Both sexes of Oncaea subtilis Giesbrecht, 1892, a small oncaeid species commonly occurring in temperate and tropical regions, are redescribed on the basis of material from the Red Sea. It is placed in a new monotypic genus, Monothula, on the basis of the loss of the outer spine on the third segment of the endopod of legs 2–4, and the presence of a single dorsal egg-sac, which is attached medially to the genital double-somite. The latter character is unique among oncaeids. The geographical distribution of M. subtilis comb. nov. is reviewed, and additional taxonomic data based on material from the eastern Mediterranean, the Arabian Sea and the eastern Indian Ocean are presented. The phylogenetic relationships of M. subtilis within the Oncaeidae are discussed.     

Genetic population structure of the lionfish Pterois miles (Scorpaenidae, Pteroinae) in the Gulf of Aqaba and northern Red Sea

The aim of this study is to reveal gene flow between populations of the coral reef dwelling lionfish Pterois miles in the Gulf of Aqaba and northern Red Sea. Due to the fjord-like hydrography and topology of the Gulf of Aqaba, isolation of populations might be possible. Analysis of 5' mitochondrial control region sequences from 94 P. miles specimens detected 32 polymorphic sites, yielding 38 haplotypes. Sequence divergence among different haplotypes ranged from 0.6% to 9.9% and genetic diversity was high (h=0.85, pi=1.9%). AMOVA indicates panmixia between the Gulf of Aqaba and northern Red Sea, but analysis of migration pattern shows an almost unidirectional migration originating from the Red Sea.     

Spatial and temporal distribution of mesozooplankton in the Gulf of Aqaba and the northern Red Sea in February/March 1999

The distribution pattern, taxonomic composition and communitystructure of mesozooplankton was studied along a transect with10 positions between the Gulf of Aqaba and the northern RedSea. Five positions were resampled two or three times duringa cruise of RV ‘Meteor’ in February/March 1999.In spite of clear differences in the density stratificationbetween the Gulf of Aqaba and the northern Red Sea, the mesozooplanktoncomposition was very similar: Copepods were by far the mostabundant taxon, contributing 76–95% to the total community.The remainder was composed largely of ostracods, chaetognaths,appendicularians and molluscs. The mesozooplankton of the deeplymixed stations was homogeneously distributed, at all other stationsthe bulk of the mesozooplankton (>70%) was concentrated inthe mixed surface layer with peaks of calanoids, cyclopoidsand appendicularians in the vicinity of the chlorophyll a (Chla) maximum layer. Ostracods and poecilostomatoids dominatedthe layers below. Standing stocks within the total water column(550–1200 m) varied between 93 and 431 x 10³ individualsm^–2 for copepods and 5–76 x 10³ individuals m^–2for other mesozooplankton with highest numbers in the northernGulf of Aqaba, where vertical mixing was deep (400–500m) and Chl a and mesozooplankton distributions homogeneous throughoutthe water column. Towards the south, the mixed depth decreasedfrom 300 m in the central Gulf of Aqaba to 50 m in the Red Sea.Cluster analysis separated three distinct groups of stations,compounding the observed differences between the northern Gulfof Aqaba (Position I) and the other positions. The analysisalso revealed temporal differences between the February andMarch sections of the cruise, indicating the winter–springtransition. The stations sampled in March are characterisedby a higher total abundance and by a higher percentage of appendiculariansand ostracods than the stations sampled in February     

Marine reserves along the Sinai Peninsula (northern Red Sea)

The coral world of the northern Red Sea, situated along the shallow waters of the Sinai Peninsula, has been intensively investigated by Israeli scientists during the last decade. Many aspects of these investigations revealed the presence of unique ecosystems which developed mainly after the last postglacial period less than 10,000 years ago! At this time the cooling of the ambient waters swept out most of the neopleistocenic coral formations, and all the richness and diversity of species observed today evolved after those profound events. At present, typical ecoformations, in which the corals comprise the main part, are found in the northern part of the Gulf of Aqaba, around the southern tip (Tiran Straits) of the Sinai Peninsula, and in the shallow parts of the Gulf of Suez. In 1967 teams of our scientists investigated the marine habitats, and a number of marine parks and sanctuaries, protected by law, have been established. Thus destruction of unique faunal elements is being prevented. Today, with the withdrawal of Israel and the return of Egypt to its territories, such parks should receive much attention and remain protected habitats for the benefit of nature and mankind.     

First record of Pristipomoides multidens (Day, 1871) from the Gulf of Aqaba,with a note on the species of Pristipomoides in the Red Sea (Actinopterygii: Lutjanidae)

The snapper Pristipomoides multidens is reported for the first time from the Gulf of Aqaba. A comparison of mitochondrial cytochrome oxidase subunit 1 (COI) with available data revealed a high similarity with a sample from Mauritius and a lower similarity with samples from the China Sea. The status and distribution of the Red Sea species of Pristipomoides are summarised.     

Experimental removal of piscivorous groupers of the genus Cephalopholis (Serranidae) from coral habitats in the Gulf of Aqaba (Red-Sea)

Synopsis Elimination of adult groupers (Cephalopholis spp.) from various reef formations in the Gulf of Aqaba (Red Sea) resulted in replenishment by other predatory fishes of different taxa and only partly (21.6%) by species of the same genus. Three years later, the initial population composition had not been restored. Recolonization by juvenile groupers presumably settling from the plankton was only 11.3%. Censuses of the typical piscine prey of groupers, before and 36 months after the removal of Cephalopholis, showed virtually no change in abundace and species composition.     

Copper enrichment reduces thermal tolerance of the highly resistant Red Sea coral Stylophora pistillata

Coral Reefs - Corals in the Gulf of Aqaba (GoA) in the northern Red Sea show high thermal tolerance. The GoA has therefore been suggested as a coral reef refuge from climate change. However, as a...     

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.     

Flexibility in Red Sea Tridacna maxima‐Symbiodiniaceae associations supports environmental niche adaptation

Giant clams (Tridacninae) are important members of Indo‐Pacific coral reefs and among the few bivalve groups that live in symbiosis with unicellular algae (Symbiodiniaceae). Despite the importance of these endosymbiotic dinoflagellates for clam ecology, the diversity and specificity of these associations remain relatively poorly studied, especially in the Red Sea. Here, we used the internal transcribed spacer 2 (ITS2) rDNA gene region to investigate Symbiodiniaceae communities associated with Red Sea Tridacna maxima clams. We sampled five sites spanning 1,300 km (10° of latitude, from the Gulf of Aqaba, 29°N, to the Farasan Banks, 18°N) along the Red Sea''s North‐South environmental gradient. We detected a diverse and structured assembly of host‐associated algae with communities demonstrating region and site‐specificity. Specimens from the Gulf of Aqaba harbored three genera of Symbiodiniaceae, Cladocopium, Durusdinium, and Symbiodinium, while at all other sites clams associated exclusively with algae from the Symbiodinium genus. Of these exclusively Symbiodinium‐associating sites, the more northern (27° and 22°) and more southern sites (20° and 18°) formed two separate groupings despite site‐specific algal genotypes being resolved at each site. These groupings were congruent with the genetic break seen across multiple marine taxa in the Red Sea at approximately 19°, and along with our documented site‐specificity of algal communities, contrasted the panmictic distribution of the T. maxima host. As such, our findings indicate flexibility in T. maxima‐Symbiodiniaceae associations that may explain its relatively high environmental plasticity and offers a mechanism for environmental niche adaptation.     

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.     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

11-yr of coral community dynamics in reefs around Dahab (Gulf of Aqaba,Red Sea): the collapse of urchins and rise of macroalgae and cyanobacterial mats

Coral Reefs - Corals from the Gulf of Aqaba (northern Red Sea) are resilient to high temperatures and therefore this region is regarded as globally important for reef conservation. However,...     

Antithamnion makroklonion sp. nov. (Rhodophyta, Ceramiales) from the Gulf of Aqaba (Red Sea)

Antithamnion makroklonion sp. nov. is described from Elat, Gulf of Aqaba (Gulf of Elat) in the Red Sea, where vegetative, spermatangial, and tetrasporangial specimens were found growing in the upper sublittoral zone. The new alga belongs to a group of Antithamnion species characterized by distichous-alternate ramification of branches, sessile tetrasporangia, and decussately arranged laterals along the bearing axis. It shows a unique combination of distinctive features, including the proximal development of gland cells on elongate branchlets that overtop the parent branch. In addition to the characteristic position, and in contrast to congeners, gland cells of A. makroklonion often have a large and prominent cytoplasmic band and one or two vacuoles. Morphological features of A. makroklonion and eight related species are tabulated and the characteristics of the new species are discussed.