Quantitative distribution of herbivorous reef fishes in the Gulf of Aqaba (Red Sea)

The distribution of the main herbivorous fishes (Acanthuridae, Scaridae, Siganidae) was studied across a coral reef of the Jordanian coast in the Gulf of Aqaba (Red Sea). Visual counts were realized by diving along transects (200 m long and 5 m wide), parallel to the shore, at 10 stations located from the lagoon to 40 m deep on the outer reef slope. Herbivorous reef fishes reach their highest abundance on the reef front, where 234 fishes were counted per 1,000 m². Their density decreases on the reef flat, with an average of 150 fish 1,000 m^-2, and is lowest on the outer reef slope (69 fish 1,000 m^-2). Surgeonfishes form 63% of the herbivorous ichthyofauna, parrotfishes 35%, and rabbitfishes 2%. Families and species display different distributions according to biota. The Acanthuridae dominate on the reef flat, whereas the Scaridae are more numerous on the outer reef slope. The evolution of the social structure of the main species was observed: the adults generally school in the lagoon and on the reef flat, but are mainly solitary on the reef slope. The distribution of juvenile individuals is more restricted: they are concentrated on the reef front and on the upper part of the reef slope.This study is part of a cooperation programme between the University of Nice (France) and the University of Jordan, to study the ecology of the coral reefs and the surrounding waters of the Jordanian coast (Gulf of Aqaba, Red Sea)     

Food composition and feeding behavior of shallow-water crinoids at Eilat (Red Sea)

The food of 3 feather-stars, Lamprometra klunzingeri Hartlaub, 1890), Heterometra savignii (J. Müller, 1841) and Capillaster multiradiatus (Linnaeus, 1758), from the shallow water of Eilat (Red Sea) was found to be composed of planktonic and semi-benthalic organisms with a strong dominance of zooplankton. Monthly analyses of stomach contents showed seasonal changes in the participation of various systematic units in the diet. Food-comprising organisms were found to be selected according to size, and those measuring up to 400 formed 90% of the diet.This research has been sponsored partly by the Oceanic Biology Branch, Office of Naval Research, U.S.A. under contract N 62558-4556 and contract F 671052 67C 0043 between the Office of N.R. and the Tel-Aviv University. It represents part of the M.Sc. thesis of the first named author (in the framework of Red Sea investigations headed by Dr. L. Fishelson).     

Seasonal variations in the stable isotopic composition and the skeletal density pattern of the coralPorites lobata (Gulf of Eilat,Red Sea)

The stable carbon and oxygen isotopic compositions of the massive coralPorites lobata collected from the Gulf of Eilat, Red Sea, in May 1987 were measured along the vertical growth axis. The isotopic compositions show annual periodicity. The variations along the isotopic profile, compared with the X-radiography, indicate that the high-density bands are enriched in¹⁸O and deposited during the season of lowest water temperature (winter). In contrast, the low-density bands are depleted in¹⁸O and deposited during the warmest seawater temperatures (summer). The stable carbon and the oxygen isotopic ratios are negatively correlated with a shift in phase. During the season of higher water temperatures and light intensity values (summer), the skeleton is depleted in¹⁸O and enriched in¹³C and vice versa for the winter time. The shift between the carbon and the oxygen isotopic curves reflects the shift between the seasonal light intensities and seasonal temperature variations in the shallow water. The oxygen isotope ratio was used to detect the seasonal variations in seawater temperatures. The coral aragonitic skeleton is depleted in¹⁸O compared to apparent equilibrium with ambient seawater. The disequilibria range from (ca.) -3.10 to -3.50 with an average value of 3.40. The isotope fractionation behavior during skeleton precipitation is discussed in light of the environmental variables.     

Nutrients and biomass distribution in the Gulf of Aqaba (Elat), Red Sea

The distribution of phosphate, nitrate and silicate values obtained from 300 samples and of biomass determined by displacement volume in about 1,900 vertical plankton hauls (65, 200 and 330 m) collected from 8 oceanographic station during 1974–1975 in the Gulf of Aqaba (Elat) are summarized and illustrated by crosssections. Generally, the Gulf is poor in nutrients and the data indicate that it is filled with upper (150 m) Red Sea waters, flowing in over the sill of Tiran. Nutrient content of the upper and deep waters immediately outside the sill are well within the known range of those in the northern Red Sea. Biomass values are relatively low in the Gulf are generally similar to those reported from the northern Red Sea. Both nutrients and biomass values display seasonal and bathymetric variations in the Gulf and outside the sill.     

Feeding of Clausocalanids (Calanoida,Copepoda) on naturally occurring particles in the northern Gulf of Aqaba (Red Sea)

A total of 12 feeding experiments were conducted in the northern Gulf of Aqaba during spring (March/April) and autumn (September/October) 2002 at the Marine Science Station (MSS) in Aqaba. Females of three species of clausocalanids were selected: Clausocalanus farrani, C. furcatus and Ctenocalanus vanus. Natural occurring particle (NOP) larger than 5 μm were investigated as food source. The ambient chlorophyll a concentration at sampling depth (∼70 m) ranged between 0.15 and 1.00 μg chl a l⁻¹ and NOP concentrations ranged between 1.78 and 14.0 × 10³ cells l⁻¹ during the sampling periods. The division of particles into five size classes (5–10, 10–20, 20–50, 50–100 and >100 μm) revealed that most of the particles were found in the size classes below 50 μm (81–98%), while most of the natural occurring carbon (NOC) was concentrated in the size classes larger than 20 μm (70–95%). Ingestion rates were food density dependent rather than size dependent ranging between 0.02 and 1.65 × 10³ NOP ind⁻¹ day⁻¹ and 0.01 and 0.41 μg NOC ind⁻¹ day⁻¹, respectively, equivalent to a body carbon (BC) uptake between 0.4 and 51.8% BC day⁻¹. The share of the size classes to the total ingestion resembled in most cases the size class composition of the natural particle community.     

Migration,habitat use,and competition among mobile corals (Scleractinia: Fungiidae) in the Gulf of Eilat,Red Sea

We examined the impact of seven species of mobile mushroom corals (Fungiidae) on the community structure of sheltered reef slopes in terms of their patterns of migration, habitat use and competition with other benthic organisms. On fringing reefs at Eilat, Red Sea, polyps detached at 1 to 6 cm length, and grew to 11–55 cm length. Attached mushroom corals were oriented vertically in reef cavities. Detached corals migrated downward on the reef slope and onto rubble or soft substratum at the reef base, at 29 to 71 cmyr^–1. Mobility decreased with corallum size and extent of undersurface ornamentation. In aquaria, small corals righted themselves and migrated up to 6 cm d^–1 by nocturnally inflating and pushing their tissues against the substratum. Autonomous coral behavior and storm-generated water motion appeared to account for most fungiid mobility at Eilat. Mushroom corals did not damage each other upon contact, even in multi-species aggregations, but unilaterally damaged non-fungiid scleractinian corals. Their dominance during contact interactions retards overgrowth by larger attached scleractinians, and mobility allows them to colonize soft substrata not accessible to most other reef corals.     

Sea-water seasonal changes at a heavy tourism investment site on the Jordanian northern coast of the Gulf of Aqaba, Red Sea

The Gulf of Aqaba exhibits a strong seasonality due to convective mixing during winter and stratification during summer. The present study provides a detailed appraisal of summer and winter sea-water characteristics at the northern coast of the Gulf of Aqaba, that is witnessing rapid development and increasing changes in its geomorphological characteristics. Sea-water temperature, salinity, nutrients, and chlorophyll a concentrations were measured biweekly at five coastal and four cross-sectional stations during the periods February to April and July to September 2004. Meteorological conditions were continuously recorded at the Marine Science Station. The coastal study sites included four open coastal stations and a marina with one-way exchange with the open water. The effect of convective mixing was clearly apparent on the sea-water characteristics. Natural seasonal characteristics of higher nutrients and chlorophyll a concentrations were recorded during winter at most of the open coastal stations. In the cross-sectional stations, the concentrations of nutrients and chlorophyll a were not different between the surface and the bottom during winter, but the bottom waters had generally higher concentrations during summer. Some deviations from the natural seasonal cycle were recorded at the marina and other coastal stations. Here, higher nutrient and chlorophyll a concentrations were recorded in summer than in winter. These deviations that are most likely due to anthropogenic effects are discussed.     

Community structure and species diversity of hermatypic corals at Eilat,Red Sea

This study investigates the community structure of reefbulding corals in terms of species composition, zonation and diversity patterns, as well as possible factors affecting the observed distributions. The study was carried out by a series of line transects run underwater with SCUBA apparatus from the reef flat to a depth of 30 m. The reefs of Eilat are of the fringing type, with seleractinian corals as the most important hermatypic organisms. The different zones of the reef are analyzed on the basis of topographical characteristics of the reef, as well as from the numerical data on abundance and living coverage, using cluster analysis of all transects surveyed. The present knowledge concerning species diversity is reviewed and analyzed in the context of hermatypic coral data. Three different diversity indices (the species count, Simpson's index and Shannon and Weaver's index) were calculated for estimating the diversity obtained on different zones of the reef. It was found that there is a successive increase in diversity of hermatypic corals from shallow water to a depth of 30 m. Species diversity and living voverage of corals were significantly greater in steeper zones as compared to flatter zones of the reef. A possible explanation for this phenomenon is the accumulation of sediments in the flat zones. It is proposed that the severe and umpredictable nature of the reef flat may account for low abundance and living coverage of corals. It is also proposed that deep-water species which do not invade shallow water are species which have developed high specialization to their local environment. The idea that light intensity is a significant factor in calcium-carbonate deposition by scleractinian corals is supported by field measurements of individual colonies at different depths.This paper is part of a thesis submitted to the State University of New York at Stony Brook in partial fulfillment of the requirements for a Ph. D. degree.     

Sea-water seasonal changes at a heavy tourism investment site on the Jordanian northern coast of the Gulf of Aqaba,Red Sea

The Gulf of Aqaba exhibits a strong seasonality due to convective mixing during winter and stratification during summer. The present study provides a detailed appraisal of summer and winter sea-water characteristics at the northern coast of the Gulf of Aqaba, that is witnessing rapid development and increasing changes in its geomorphological characteristics. Sea-water temperature, salinity, nutrients, and chlorophyll a concentrations were measured biweekly at five coastal and four cross-sectional stations during the periods February to April and July to September 2004. Meteorological conditions were continuously recorded at the Marine Science Station. The coastal study sites included four open coastal stations and a marina with one-way exchange with the open water. The effect of convective mixing was clearly apparent on the sea-water characteristics. Natural seasonal characteristics of higher nutrients and chlorophyll a concentrations were recorded during winter at most of the open coastal stations. In the cross-sectional stations, the concentrations of nutrients and chlorophyll a were not different between the surface and the bottom during winter, but the bottom waters had generally higher concentrations during summer. Some deviations from the natural seasonal cycle were recorded at the marina and other coastal stations. Here, higher nutrient and chlorophyll a concentrations were recorded in summer than in winter. These deviations that are most likely due to anthropogenic effects are discussed.     

Quality standard codes of reference of Jordanian coastal waters of the Gulf of Aqaba, Red Sea

Back in 1992, the Gulf of Aqaba Environmental Action Plan (GAEAP), a collaboration between the Aqaba Region Authority (ARA), Jordan and the World Bank, gave considerable emphasis to the environmental protection of the Gulf of Aqaba [The World Bank. Gulf of Aqaba Environmental Action Plan. Report No. 12244JO (1993).]. The document recommended the establishment of a marine reserve and the long term monitoring of the coastal habitats' environmental quality. The combination of a dedicated follow up, the collaborative efforts of ARA and the Marine Science Station (MSS), and the founding of the Aqaba Special Economic Zone Authority (ASEZA) have turned the recommendations into reality. A comprehensive monitoring program of the Jordanian coastal habitats commenced in 1999. The first three years of the program were financed by a donation from The Global Environmental Facility (GEF). In return, Jordan has committed itself to the maintenance of the monitoring program as an ongoing tool for sustainable coastal management. The monitoring program includes observations on benthic habitat, fish communities, bottom sediments and seawater quality. This paper focuses on the results of seawater-quality monitoring in the first three years. Records of weather conditions, coastal currents, seawater temperature, transparency, salinity, density, pH, alkalinity, dissolved oxygen, ammonia, nitrate, nitrite, phosphate, silicate, particulate matter, chlorophyll a, zooplankton biomass, total coliform, fecal coliform, hydrocarbons and sedimentation rate have been generated monthly since January 1999 at six coastal stations, and one offshore reference station, in the Jordanian waters of the Gulf of Aqaba. The coastal stations are located at sites with different benthic habitats and are occupied by different human activities. Offshore records of density (thermohaline structure), nutrients and chlorophyll a depicted two well-defined seasons; a nutrient-/chlorophyll a-rich, mixed water winter from December to April and a nutrient-/chlorophyll a-poor, stratified water summer from June to October. Short transition seasons appeared in May and November. The mixing and stratification seasons were also clearly depicted in the coastal waters. Statistical analysis of the three-year data collected at the offshore station revealed no significant inter-annual differences in the upper 125 m of the water column with respect to any of the measured parameters. At coastal stations, the water quality at the two northernmost stations was significantly different in comparison to the upper 125 m at the offshore station and to the other coastal stations, with respect to the two key indicator parameters: inorganic nitrogen and chlorophyll a. The three-year findings of the monitoring program are employed to suggest standard codes of reference for the coastal water quality.     

The distribution of molluscan assemblages and their postmortem fate on coral reefs in the Gulf of Aqaba (northern Red Sea)

Molluscan assemblages were studied on fringing reefs (reef flats, Millepora-fringing reefs, fringing reefs with massive corals) and fore-reef hard substrata (coral patches, coral carpets and small patch reefs) in the Gulf of Aqaba at water depths ranging from the intertidal to 26 m. A total of 1,665 molluscan individuals from 51 taxa was counted on 44 transects, which covered 220 m2 at eight diving sites. The most important molluscs in the assemblage were the parasitic gastropod Coralliophila neritoidea, the encrusting gastropod Dendropoma maxima and the coral-associated bivalve Pedum spondyloideum. The dead assemblage, in contrast, was dominated by encrusting bivalves (Ostreoidea, Chamoidea, Spondylidae) and the coral-predating gastropod Drupella cornus. Distinct molluscan assemblages inhabit each of the three fringing reef-habitats and most of the important depth-related community changes occurred within the uppermost 5 m. In contrast, the three deeper fore-reef habitats are characterized by a more uniform molluscan composition. Molluscan assemblages were more dependent on substrata and their coral associations than on water depth. Comparisons with other published studies indicate that reefoidal hard substrata in the northern Red Sea are largely characterized by similar species-abundance patterns. The minor differences to other Red Sea studies probably reflect the northern, isolated position of the Gulf of Aqaba, the lack of certain molluscan habitats, and the differential impact of anthropogenic influences. Strong differences between living and dead assemblages in Aqaba are similar to those observed in other regions and are due to distinct biases in the dead assemblage. Molluscs closely associated with living corals (mostly bivalves and Dendropoma) can easily be overgrown after death and are thus undetectable in visual censuses. Some gastropod taxa are preferentially transported into surrounding soft-substrata postmortem or redistributed by hermit crabs. Such complex relationships between ecology and taphonomy are crucial in evaluating the quality of the molluscan fossil record in coral reef environments. The comparison of our results with literature data documents an increase in coral predators during the last two decades in the northern Red Sea. Due to the greater mollusc biodiversity in the shallower Aqaba reef habitats, damage to this coral reef zone would have the greatest impact on the overall mollusc community.     

Some aspects of recolonization of coral rocks in Eilat (Gulf of Aqaba) by fish populations after poisoning

As a result of an accident, a limited strip of the coast of Eilat (Gulf of Aqaba, Red-Sea) was affected by poison that killed all fishes. The area was observed monthly for the following year, to study the recovery of fish populations. The study also included observations on growth rate of fish and size of populations. Recovery of fish populations was complete 10 to 12 months after the poisoning. It is believed that this is due to the relatively small size of the contaminated area and the survival of most invertebrates that constitute an important part of the biotope of the fish.     

Littoral fauna of the Red Sea: the population of non-scleractinian anthozoans of shallow waters of the Red Sea (Eilat)

Approximately 22 species of sea anemones and 20 species of alcyonarians were found on the reef fringing the shallow water at Eilat (Gulf of Aqaba) and other localities along the Red Sea coast of the Sinai Peninsula. Investigations showed that these 2 groups of non-scleractinian coelenterates from (on different substrates) well identified colonies, part of them inhabiting sandy bottoms, other groups growing on the reef surface. Observations lead to the conclusion that several of the sea anemones and alcyonarians can act, under certain conditions, as factors limiting the development of hermatypic corals. In some areas, several species of the 2 groups of animals are found together with their symbiotic partners, such as fishes and crustaceans.This research has been sponsored partly by a grant from the Smithsonian Institution, No. SFC-7-0074 to L. B. Slobodkin (N. Y.) and L. Fishelson (T.-A.).     

Initial observations of the zooplankton microdistribution on the fringing coral reef at Aqaba (Jordan)

Diurnal and nocturnal zooplankton was sampled simultaneously at the surface and at 35 m depth on the outer slope of the fringing reef off the Jordanian coast in the Guff of Aqaba, Red Sea, over a period of 8 d during July 1982. Complementary surface sampling was carried out above the reef and in the open sea in the morning and afternoon for a period of 5 d. Five groups of organisms were classified according to changes in their vertical distribution in the water column which is analyzed and discussed. Zooplankters of the first group appear to be positively phototactic (Calanus robustior, Mecynocera clausi, Oithona helgolandica, Corycella rostrata). Those of the second group also seem to be positively phototactic, but only as regards average and low light (moonlight) intensities (Nannocalanus minor, Acartia negligens, Sagitta enflata, S. hexaptera, Thalia democratica, Oikopleura fusiformis, O. longicauda). The zooplankton of the third group displays a negative phototaxis (Oithona nana, calyptopis and furcilia euphausiid stages, pelecypods and other mollusc veligers). Species exhibiting uncertain behaviour or caught in low numbers and exhibiting no particular distribution are relegated to the fourth and the fifth groups. The horizontal microdistribution of the zooplankton community is not easily discernible: certain neritic or oceanic affinities can be identified, however, by comparing the WP2 standard net surface samples collected above the reef with those from the open sea. The importance of zooplankton communities is emphasized, and the necessity of elucidating the temporal and spatial migrations and exchanges between reef and planktonic organisms in order to gain more insight into their trophic relationship with the reef system.     

Three party symbiosis: acoelomorph worms,corals and unicellular algal symbionts in Eilat (Red Sea)

Epizoic worms were found to occur on certain coral colonies from reefs off the coast of Eilat (Red Sea). We identified 14 coral species infested by acoelomorph worms at a depth range of 2–50 m. The host corals were all zooxanthellate and included both massive and branching stony corals and a soft coral. Worms from all hosts were identified as belonging to the genus Waminoa and contained two distinct algal symbionts differing in size. The smaller one was identified as Symbiodinium sp. and the larger one is presumed to belong to the genus Amphidinium. Worm-infested colonies of the soft coral, Stereonephthya cundabiluensis, lacked a mucus layer and exhibited distinct cell microvilli, a phenotype not present in colonies lacking Waminoa sp. In most cases, both cnidarian and Acoelomorph hosts displayed high specificity for genetically distinctive Symbiodinium spp. These observations show that the epizoic worms do not acquire their symbionts from the “host” coral.     

Response of the microbial food web to manipulation of nutrients and grazers in the oligotrophic Gulf of Aqaba and northern Red Sea

The control mechanisms within the pelagic microbial food web of the oligotrophic Gulf of Aqaba and the northern Red Sea were investigated in the spring of 1999. Nutrient conditions and potential grazer impact were manipulated in a series of dilution experiments. Ambient nutrient concentrations and autotrophic biomass were very low (0.23–1.21 µmol NO₃ l^–1, 0.06–0.98 µmol NH₄ l^–1, 1.08–1.17 µmol Si l^–1, 0.08–0.12 µmol P l^–1, 0.15–0.36 µg chlorophyll a l^–1). The planktonic community was characterized by low abundances [3.0–5.5×10⁵ heterotrophic bacteria ml^–1, 0.58–7.2×10³ ultraphytoplankton <8 µm ml^–1 (small eukaryotic photoautotrophs and Prochlorococcus sp., excluding Synechococcus sp.), 0.45–4.4×10⁴ Synechococcus sp. ml^–1, 0.32–1.2×10³ heterotrophic nanoflagellates ml^–1, 1.3–3.8×10³ phytoplankton >8 µm l^–1, 0.93–5.4×10² microzooplankton l^–1] and dominated by small forms (0.2–8 µm). Dinoflagellates and oligotrichous ciliates were the most common groups in initial samples among the phytoplankton >8 µm and microzooplankton, respectively. Results show that bottom-up and top-down control mechanisms operated simultaneously. Small organisms were vulnerable to grazing, with maximum grazing rates of 1.1 day^–1 on heterotrophic bacteria and 1.3 day^–1 on ultraphytoplankton. In contrast, algae >8 µm showed stronger signs of nutrient limitation, especially when the final assemblages were dominated by diatoms. Synechococcus sp. were not grazed and only showed moderate to no response to nutrient additions. The high spatial and temporal variation of our results indicates that the composition of the planktonic community determines the prevailing control mechanisms. It further implies that, at this transitional time of the year (onset of summer stratification), the populations fluctuate about an equilibrium between growth and grazing.Communicated by O. Kinne, Oldendorf/Luhe     

Coral recruitment: a spatio-temporal analysis along the coastline of Eilat,northern Red Sea

Recruitment rates of stony corals to artificial substrates were monitored for 2 years at 20 sites along the coast of Eilat, northern Red Sea, to compare with those recorded at other coral reef locations and to assess variation in recruitment at several spatial scales. Coral recruitment was low compared to that observed on the Great Barrier Reef in Australia, but was similar to levels reported from other high-latitude reef locations. Pocilloporids were the most abundant coral recruits in all seasons. Recruitment was twofold higher during the first year than during the second year of study. There was considerable spatial variability, with the largest proportion of variance, apart from the error term, attributable to differences between sites, at a scale of 10² m. Spearmans ranked correlation showed consistency in spatial patterns of recruitment of pocilloporid corals between years, but not of acroporid corals. During spring, when only the brooding pocilloporid coral Stylophora pistillata reproduces at this locality, most coral recruitment occurred at central and southern sites adjacent to well-developed coral reefs. During summer, recruitment patterns varied significantly between years, with wide variation in the recruitment of broadcasting acroporid corals at northern sites located distant from coral reefs. Settlement was low at all sites during autumn and winter. This work is the first detailed analysis of coral recruitment patterns in the Red Sea, and contributes to the understanding of the spatial and temporal scales of variation in this important reef process.Communicated by O. Kinne, Oldendorf/Luhe     

Cytology of an unusual case of hermaphroditic gonads in the tropical sea urchin Tripneustes gratila from Eilat (Red Sea)

One specimen of the tropical sea urchin Tripneustes gratila, distributed along the rocky subtide of Eilat, Israel, was found to be hermaphroditic, producing fertile sperm and eggs. In this case, internal self-fertilization, producing embryos along the gonads, was also observed.