Georg A. Heiss

FACTORS CONTROLLING HOLOCENE REEF GROWTH - AN INTERDISCIPLINARY APPROACH

SummaryThis interim report deals with investigations on key factors controlling reef growth by zoophysiologists, ecologists, paleontologists and geologists. The different levels of emphasis are the coral animal and the reef community. The main study area is the Red Sea which reaches over 20°C latitude up to the northernmost margin of the global coral reef belt. Supplementary results on microborer ecology are provided from the Bahamas.The desert enclosed Red Sea, not influenced by land runoff and only minimally by anthropogenic (urban and touristic) nutrient inputs, is predestined for a study on the principal influence of light on calcification within bathymetrical and latitudinal gradients. Hence, on the level of the zooxanthellate scleractinian animal phototrophic and heterotrophic energy supply and its bearing on calcification are being measured in different coral species—in particular inPorites sp., one of the most important reef builders.The growth of 15 zooxanthellate scleractinians in the Gulf of Aqaba correlates with the annual light cycle. This correlation is observable down to 40 m depth. Other growth promoting factors seem to have less influence on coral extension. The availability of organically enriched sediments in shallow water probably yields nutritional value, in particular for filter feeding species, thus restricting their distribution to those areas. Zooxanthellae, when isolated fromMycedium elephantotus, are different in their dependence on depth in maximum rates of photosynthesis and photosynthetic efficiency (-slope). Increasing concentrations of pigments as a function of depth could be determined. Maximum rates of photosynthesis of zooxanthellae in vivo, collected at corresponding depth, have been 4 times higher. Structural and physiological adaptations improving heterotrophic and phototrophic energy intake are highlighted. Porites sp. was the subject of annual growth studies at locations extending from Aqaba in the North over the northern and southern Egyptian coast and islands, Sanganeb Atoll and Wingate reef offshore Sudan to the Gulf of Tadjoura in the Gulf of Aden (Djibouti). Mean growth rates in the shallow water zone increase with decreasing latitude and are highest at the southernmost studied reefs in the Gulf of Tadjoura. However, the observed latitutdinal growth reduction is restricted to the upper ca. 15 m of the water column. The upper limit of growth potential decreases with depth parallel to the decrease of light availability. Highest growth rates are recorded in shallow depth (10–2.9 mm yr−1). This zone reaches at Aqaba (29°30′N) to a depth of ca. 10 m. At the southern Egyptian reefs (24°30′N) this zone extends to ca. 15 m water depth. This effect is probably a result of the stronger reduction of winter light levels and water temperature in the northern regions. Compared to other oceans the decrease of growth with increasing latitude of Red SeaPorites corals is far less, and growth rates at Aqaba are the highest observed at these latttudes.On the level of the community of reef inhabitants four principal topics are addressed:The first one is the dynamics of the proportions of hermatypic and ahermatypic organisms and open space. The occurrence of stony and soft corals and the sharing of empty space in different reef sections at Aqaba and on Sanganeb Atoll were quantified. Soft corals, mainlySinularia- and xeniid species, occupy decreasing shares with depth. Among theXenia species a bathymetrical zonation pattern was detected.The next issue is the growth impeding role of soft corals and gastropod parasites and predators on scleractinians. Experimental and field observations showed xeniid soft corals to be opportunistic i.e. occupying rapidly open space rather than to attacking and outcompeting stony corals. An increasingly specialized behaviour was detected among corallivorous gastropods of the family Coralliophilidae to exploit their coral hosts. Whereas these snails are more or less sessile and depend for a long time on the surrounding host polyps the mobileDrupella cornus (Thaididae) forms feeding aggregations which denude mainly branching corals on shallow reef parts.Furthermore, the role counteracting reef growth of macro- and microbioeroders is investigated.Diadema setosum is a major destructive agent on reefs at Aqaba (not in the central Red Sea). The grazing sea urchins do not only keep potential colonization area free but also erode carbonate material (e. g. 1468 g/m2/year, 10 m depth). Demographic and bathymetric patterns in the sea urchin population are analyzed including their bearing on bioerosion of the reef. Investigations on microboring organisms in carbonate material have started in the Red Sea; initial results, however, are only available from similar studies near Lee Stocking Island, Bahamas.Three major environments have been identified based on the distribution of the different microborers. These are1)the intertidal environment dominated by boring cyanobacteria.,2)reef sites from 2 to 30 m water depth dominated by a diverse assemblage of boring cyanobacteria and chlorophytes, and3)the deep reef slope from 100 to 300 m dominated by boring green algae and heterotrophs. The boring chlorophyte genusPhaeophila appears rapidly and dominates at sites from 2 to 30 m, but it leaves vacated borings and is replaced byOstreobium quekettii after 1 year. Different substrate types show very different rates of colonization by microborers. The greatest excavation rates (100 g/m2/3 months) occur in fine-grained limestone, while the slowest rates (0.5 g/m2/3 months) occur in calcite crystals. Molluscan shell material shows intermediate rates of excavation. Light conditions appear very important in determining the growth rate and distribution of different microborers between the sites, however, the interaction of light with other factors, such as substrate, time period of exposure, and water quality conditions may be involved.

Increased seasonality in the Gulf of Aqaba, Red Sea, recorded in the oxygen isotope record of aPorites lutea coral

At Aqaba in the Northern Gulf of Aqaba, Red Sea, the stable oxygen isotope record of aPorites lutea coral colony was studied. From a vertical and a horizontal core two parts from the base of the colony (10 years of coral growth), and two parts from the living surface (12 years of coral growth) were analysed with a monthly sampling resolution. The results show that seasonality of sea surface temperatures (SST) in the northern Gulf of Aqaba has increased since the early 19th century, mainly due to higher summer temperatures. δ18O-ratios indicate that SST increased by at least 1.3°C. Horizontal and vertical cores display different variations in the modern parts, probably caused by the very shallow water depth of the youngest part of the vertical core.KurzfassungBei Aqaba im nördlichen Golf von Aqaba, Rotes Meer, wurde die Zusammensetzung der stabilen Sauerstoffisotope in einer Kolonie der SteinkorallePorites lutea untersucht. Aus einem vertikalen und einem horizontalen Bohrkern wurden jeweils ein Abschnitt von der Basis der Kolonie (10 Jahre Korallenwachstum), sowie je ein Abschnitt mit 12 Jahren Korallenwachstum bis zur lebenden Oberfläche mit einer monatlichen Beprobungsauflösung analysiert. Die Ergebnisse zeigen, daß die Saisonalität der Meerwassertemperatur im nördlichen Golf von Aqaba seit dem frühen 19. Jh. deutlich verstärkt ist, was vor allem auf erhöhte Sommertemperaturen zurückzuführen ist. Die δ18O-Werte lassen daraufschließen, daß die Temperatur um mindestens 1.3°C gestiegen ist. Die jungen Abschnitte des horizontalen und vertikalen Kerns zeigen unterschiedliche Variationen sowohl in der langfristigen Erwärmung als auch in der verstärkten Saisonalität, die auf die sehr geringe Wassertiefe des jüngsten Teils im vertikalen Kern zurückgeführt werden.

Carbonate production by scleractinian corals at Aqaba, Gulf of Aqaba, Red Sea

SummaryIn a fringing reef at Aqaba at the northern end of the Gulf of Aqaba (29°26′N) growth rates, density, and the calcification rate ofPorites were investigated in order to establish calculations of gross carbonate production for the reefs in this area. Colony accretion ofPorites decreases with depth as a function of decreasing growth rates. The calcification rate ofPorites is highest in shallow water (0–5 m depth) with 0.9 g·cm−2·yr−1 and falls down to 0.5 g·cm−2·yr−1 below 30 m. Scleractinian coral gross production is calculated from potential productivity and coral coverage. It is mainly dependent on living coral cover and to a lesser extent on potential productivity. Total carbonate production on the reef ranged from 0 to 2.7 kg/m2 per year, with a reef-wide average of 1.6 kg/m2 perycar. Maximum gross carbonate production by corals at Aqaba occurs at the reef crest and in the middle fore-reef from 10 to 15 m water depth. Production is low in sandy reef parts. Below 30 m depth values still reach ca. 50% of shallow water values. Mean potential production of colonies and gross carbonate production of the whole reef community at Aqaba is lower than in tropical reefs. However, carbonate production is higher than in reef areas at the same latitude in the Pacific, indicating a northward shift of reef production in the Red Sea.