Marginal tentacles of the corallimorpharian Rhodactis rhodostoma. 1. Role in competition for space

The corallimorpharian Rhodactis rhodostoma (Ehrenberg, 1934) forms aggregations that dominate patches on some coral reef flats in the Red Sea. The outcomes and mechanisms of competition for space between this corallimorpharian and other sessile organisms are poorly understood. Polyps of R. rhodostoma were observed to overgrow zoanthids, hydrozoan corals, sponges and encrusting macroalgae on a fringing reef at Eilat, northern Red Sea. R. rhodostoma polyps also damaged, and in some cases overgrew, reef-building corals in the families Poritidae, Acroporidae and Pocilloporidae, most of which form branching colonies with small polyps that are subordinate in coral competitive hierarchies. In contrast, most stony corals in the families Faviidae and Mussidae had standoff interactions with R. rhodostoma, in which they prevented the corallimorpharians from damaging them or approaching closer than 1 to 3 cm. The latter corals are ranked at the top of competitive hierarchies for Indo-Pacific corals, and they form massive colonies of large polyps which may develop aggressive organs termed sweeper tentacles. Some soft corals that exude allelopathic chemicals also avoided overgrowth by the corallimorpharians. Tentacles along the oral disk margin of R. rhodostoma polyps were swollen and bulbous during contacts with cnidarians. These bulbous marginal tentacles had significantly thicker ectoderm and a higher proportion of holotrichous nematocysts than did the normally filiform marginal tentacles of R. rhodostoma polyps. It is concluded that, on the reef flat at Eilat, this corallimorpharian damages and overgrows a variety of sessile competitors, including branching stony corals, via the application of specialised marginal tentacles filled with penetrating nematocysts. R. rhodostoma is an intermediate competitor in the aggressive hierarchy among Indo-Pacific Anthozoa, including the reef-building corals. Received: 1 July 1998 / Accepted: 24 March 1999     

Adaptation strategies of the corallimorpharian <Emphasis Type="Italic">Rhodactis rhodostoma</Emphasis> to irradiance and temperature

Corallimorpharians may dominate some habitats on coral reefs and compete with stony corals for access to light, yet little is known concerning their photosynthetic traits. At Eilat in the northern Red Sea, we observed that the abundance of individuals of the corallimorpharian Rhodactis rhodostoma decreased significantly with depth on the reef slope. Field and laboratory experiments revealed that they employ several mechanisms of photoadaptation to high irradiance on the shallow reef flat. Their endosymbiotic microalgae (zooxanthellae) varied significantly in both abundance and chlorophyll content with level of irradiance. Use of a diving pulse amplitude modulated fluorometer revealed that the zooxanthellae of R. rhodostoma effectively disperse excess light energy by expressing significantly higher values of non-photochemical quenching and maximum excitation pressure on photosystem II when experimentally exposed to high light (HL) versus low light (LL). Host corallimorpharian tissues mediated this response by shielding the algal symbionts from high irradiance. The endoderm of host tentacles thickened significantly and microalgal cells were located further from the mesoglea in HL than in LL. The clades of zooxanthellae hosted by the corallimorpharians also varied with depth. In shallow water, all sampled individuals hosted clade C zooxanthellae, while in deep water the majority hosted clade D. The photosynthetic output of individuals of R. rhodostoma was less affected by HL than was that of a stony coral examined. When exposed to both high temperature (HT) and HL, individuals of R. rhodostoma reduced their maximum quantum yield, but not when exposed to HL at low temperature (LT). In contrast, colonies of the scleractinian coral Favia favus reduced their photosynthetic output when exposed to HL in both temperature regimes. After 2 weeks of HT stress, R. rhodostoma polyps appeared to bleach completely but re-established their zooxanthella populations upon return to ambient temperature. We conclude that mechanisms of photoadaptation to high irradiance employed by both the endosymbiotic zooxanthellae and host corallimorpharians may explain in part the abundance of R. rhodostoma on some shallow reef flats. The ability to survive for weeks at HT while bleached also may allow corallimorpharians to repopulate shallow reef areas where scleractinians have been killed by thermal stress. B. Kuguru and G. Winters contributed equally to this work.     

The reef environment and competitive success in the Corallimorpharia

Competitive success within coral reef communities is controlled by various factors. In addition to competitive abilities in direct interactions with a contestant, external influences such as disturbance caused by nutrient input may determine the outcome of antagonistic interactions. We examined the competitive success of corallimorpharians on coral reefs by investigating their distribution patterns within reefs and how well they perform in interference competition with staghorn corals in different environments. Substrate composition and corallimorpharian growth were examined on three reefs in Tanzania under different disturbance regimes using the line-intercept transect and point techniques. A transplant experiment was conducted in which staghorn corals (Acropora formosa) were exposed to the polyps of Rhodactis rhodostoma to establish how competition between corals and corallimorpharians affects their respective distributions. Within reefs corallimorpharians seemed to be more competitive in shallow waters. This could be due to both environmental factors as well as varied competitive abilities depending on surrounding benthos that changed with depth. Reef environment also seemed to influence corallimorpharian growth among reefs as they had the highest densities in the areas with the highest nutrient loads. The transplant experiment revealed that the corallimorpharians had a competitive advantage over the corals, and in comparisons of reefs influenced by different degrees of disturbance, corallimorpharians were most competitive in the area with the highest nutrient content. Hence, stress on coral reefs in the form of raised nutrient loads may favour the competitive success of corallimorpharians.Communicated by P.W. Sammarco, Chauvin     

Mode and timing of reproduction in some common hermatypic corals of Hawaii and Enewetak

Common Hawaiian and Enewetak corals were examined to determine the method and mining of reproduction. Of the 7 Hawaiian species examined for the release of planulae, only 2 have planulated in captivity, Pocillopora damicornis and Cyphastrea ocellina. Both planulate year-round and both are characteristic of reef flats. Four of the 5 species which did not planulate were found to contain eggs, but not planulate, when polyps were examined microscopically. These 5 species do not usually occur on reef flats. Seven of the 12 Enewetak species examined in June, July, August and January planulated; 4 of these were pocilloporids, all of which are common in shallow water. Only 3 of the 8 species of Acropora planulated, and these 3 occur solely in shallow water. A greater proportion of the Pocillopora spp. colonies than Acropora spp. colonies planulated and they released more planulae per head. In previous studies and in this one, coral species which have released planulae are characteristic of shallow-water environments such as reef flats. Most of the 10 species reported on here which failed to planulate in captivity are not commonly found on reef flats. The failure to detect planulation in so many species, particularly those of deeper water, suggests that common hermatypic corals may not all reproduce in the same way, and that mode of reproduction may be related to habitat.     

In situ thermal dynamics of shallow water corals is affected by tidal patterns and irradiance

We studied the diel variation of in situ coral temperature, irradiance and photosynthetic performance of hemispherical colonies of Porites lobata and branching colonies of Porites cylindrica during different bulk water temperature and tidal scenarios on the shallow reef flat of Heron Island, Great Barrier Reef, Australia. Our study presents in situ evidence that coral tissue surface temperatures can exceed that of the surrounding water under environmental conditions typically occurring during low tide in shallow reef or lagoon environments. Such heating may be a regular occurrence on shallow reef flats, triggered by the combined effects of high irradiance and low water flow characteristic of low Spring tides. At these times, solar heating of corals coincides with times of maximum water temperature and high irradiance, where the slow flow and consequent thick boundary layers impede heat exchange between corals and the surrounding water. Despite similar light-absorbing properties, the heating effect was more pronounced for the hemispherical P. lobata than for the branching P. cylindrica. This is consistent with previous laboratory experiments showing the evidence of interspecific variation in coral thermal environment and may result from morphologically influenced variation in convective heat transfer and/or thermal properties of the skeleton. Maximum coral surface warming did not coincide with maximum irradiance, but with maximum water temperature, well into the low-tide period with extremely low water flow in the partially drained reef flat, just prior to flushing by the rising tide. The timing of low tide thus influences the thermal exposure and photophysiological performance of corals, and the timing of tidally driven coral surface warming could potentially have different physiological impacts in the morning or in the afternoon.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Transplantation of juvenile corals: a new approach for enhancing colonization of artificial reefs

 Coral reefs in the northern Gulf of Eilat are exposed to continuous man-made disturbances, resulting in decreased coral coverage and reduced recruitment at the Nature Reserve of Eilat. The construction of artificial reefs on sandy bottoms is a possible option to decrease diving pressure on natural reefs. In the present study we tested this hypothesis by submerging an experimental artificial reef anchored to the bottom at 18 m depth and floated vertically 3 m below water surface. The reef was composed of PVC plates, attached both vertically and horizontally along a wire. Propagules of two coral species, the stony coral Stylophora pistillata and the soft coral Dendronephthya hemprichi, were transplanted to this artificial reef. Planulae of S. pistillata were obtained during the breeding season, seeded in petri dishes in the laboratory and after 2 wk the dishes were transferred to the experimental artificial reef. Automized fragments of D. hemprichi which had previously settled on 10 × 10 cm PVC plates were transplanted onto the experimental artificial reef. The survivorship of the transplanted D. hemprichi colonies was significantly higher on the lower sides of shallower plates. Survivorship of S. pistillata colonies increased with depth when located on the vertical plates, or on the upper sides of the horizontal plates. The highest survivorship of this coral was on the vertical plates and on the upper sides of the horizontal plates, while very low survivorship was recorded on the lower sides. The results indicate that vertical artificial surfaces offer the optimal biotic and abiotic conditions for the survival of the two examined corals. The vertical plates are characterized by low sed imentation rates, low coverage of turf-algae, minimal grazing by sea urchins and absence of the competitor tunicate Didemnum sp. In addition, the vertical orientation of the experimental plates reduces shading and offers the required light intensity for zooxanthellate corals such as S. pistillata. Only a few studies to date have tried to implement artificial reefs in a coral reef environment. The results of the present study indicate the potential of enhancing recruitment of corals by transplantation of juvenile recruits onto appropriate artificial structures. Maximal survivorship of these recruits is dependent upon the structural features of the artificial reef, which should offer optimal conditions. Received: 25 May 1996 / Accepted: 15 July 1996     

Recreational diving impacts and the use of pre-dive briefings as a management strategy on Florida coral reefs

Ecotourism often is promoted as an ecologically sustainable activity, but some ecotourism activities negatively impact coastal ecosystems. Impacts of intensive diving tourism on coral reefs remain poorly understood, especially in the Florida Keys. We determined patterns of recreational dive frequency, diver behaviour, and coral damage on reefs near Key Largo, and assessed how pre-dive briefings and other factors affect these damage rates. Recreational divers contacted live stony corals ~ 18 times per scuba dive; most contacts deposited sediment onto corals, but also caused abrasion to coral tissues and fracture of coral skeletons. Divers who received pre-dive ecological briefings caused significantly less coral damage than those who did not, and divers with cameras and/or gloves caused the most damage. The proportion of damaged corals increased significantly with the estimated rate of recreational diving on each reef, and the percent cover of live corals decreased. We conclude that current rates of recreational diving in Key Largo are unsustainable, resulting in damage to >80 % of coral colonies and reduction of live coral cover to <11 % at heavily-dived sites. We recommend that dive tour operators administer pre-dive ecological briefings to all recreational divers, provide extra briefings to camera and glove users, and employ underwater dive guides who intervene when divers inadvertently damage live stony corals. This study provides a scientific basis to support management of intensive ecotourism on Florida coral reefs.     

Influence of UV radiation on the survival of larvae from broadcast-spawning reef corals

Effects of ambient ultraviolet light on the survivorship of eggs and planulae larvae was investigated for three species of broadcast-spawning reef corals, Acropora palmata, Montastraea annularis, and M. franksi. Eggs and larvae from these corals contain high concentrations of lipids (60–70% by weight) and float in surface waters for 3–4 days following spawning. Larvae originating from colonies living at deeper sites on the reef exhibited significantly lower survivorship than conspecifics originating from parents in shallow water when experimentally exposed for up to 4 days to ambient surface levels of ultraviolet radiation (UVR). Concentrations of the UVR-protective compounds correlated positively with survival and matched concentrations found in parent colonies, implying that higher concentrations of ultraviolet B protective compounds are responsible for greater survival of eggs and larvae from shallow compared to deeper-dwelling parents. Ultraviolet B appears to be responsible for most of the observed differences in larval survivorship with ultraviolet A playing a minor or insignificant role. Data presented here indicate that coral recruits on Caribbean reefs and elsewhere may originate primarily from adult colonies dwelling in shallow water.Communicated by P.W. Sammarco, Chauvin     

Association of <Emphasis Type="Italic">Waminoa</Emphasis> sp. (Acoela) with corals in the Wakatobi Marine Park,South-East Sulawesi,Indonesia

This is the first quantitative study on the prevalence of epizoic Waminoa sp. acoel worms and their association with corals in the Wakatobi Marine National Park (WMNP), South-East Sulawesi, Indonesia. Three replicate transects were laid on the reef crest, flat and slope at six sites in 2006 and eight sites in 2007. Four of the sites were common in both years. In total 69 transects were surveyed in 2006, and 87 transects in 2007. A total of 4.8% of all observed hard corals were associated with acoel worms in 2006 and 2.6% of hard and soft corals in 2007. Acoels were present on 16 and 21 of the coral taxa studied in 2006 and 2007 respectively. The worms were strongly associated with the azooxanthellate coral Tubastrea spp. and were rare or absent on the most abundant coral genera Montipora and Porites. The mean number of corals having acoels was highest on reef slopes, whereas acoels were virtually absent on reef flats. Corals that had a high and a medium cover of worms were more common in 2007 than in 2006. No significant trend in the adaptation of the zooxanthellae of Waminoa sp. to different depths at different sites was revealed. The impact of the worm on the coral is unknown, but high numbers may have a shading effect and a negative impact on the coral’s photophysiology. This acoel merits more study of its life cycle, its photophysiology, and its impact on its host corals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Ecological and biological phenomena influencing coral-species composition on the reef tables at Eilat (Gulf of Aqaba,Red Sea)

Observations performed on coral colonies damaged during a very low tide on the shallow-water reef tables at Eilat (Red Sea), showed that most colonies are able to regenerate if parts of the living tissue remain intact on the skeletons. Brain-like corals, such asFavia favus andPlatygyra lamellina, were found to be more resistant to dessication than the delicate bush-like forms. It was also observed that the exposed skeletons of corals were immediately covered by a successive group of algae with a rich invertebrate epifauna among them. It is postulated that unpredicted extreme low water, occurring once in several years, could be the factor that prevents crowding on the reef tables, thus keeping high the level of coralspecies diversity.Supported, in part, by a grant from the Smithsonian Institution, USA. SFC-70074.This paper was presented at the Intecol Symposium held in Dubrovnik in March, 1972.     

Flesh or bone? Quantifying small-scale coral morphology using with-tissue and without-tissue techniques

The quantification of small-scale coral morphology using tissue-free skeletons often requires colonies, or colony sections, to be removed from the reef. This undesirable destruction can be reduced by using a with-tissue technique based on photographic images of living corals. The results of this study indicate that with-tissue multivariate morphometric data contain similar information to skeleton-derived data and can help identify phenotypic plasticity in the massive coral Favia speciosa. With-tissue data can also be used to discern changes with time in individual polyps exposed to new environments.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Stimulation of fat-body production in the polyps of the coralPocillopora damicornis by the presence of mutualistic crabs of the genusTrapezia

A mutualism exists between the xanthid crabs of the genusTrapezia and their host corals,Pocillopora damicornis. It has previously been established that these obligate coral residents benefit the coral hosts by defending them against echinoderm predators and by increasing the survival of polyps located deep between the coral branches. In turn, the corals apparently benefit the crabs by producing lipid-filled structures on which the trapezid crabs feed; these fat bodies may contain some of the lipid which in previous studies of coral metabolism has been termed excess. It was determined by experiments conducted at the Hawaii Institute of Marine Biology that the presence of crabs in colonies ofP. damicornis stimulates the polyps to produce the lipid-filled fat bodies; removal of crabs causes corals to cease producing fat bodies. A structure very similar to the fat bodies ofP. damicornis has been reported inAcropora durvillei. Both of these coral genera ordinarily possess xanthid-crab mutualists. This association between branching corals and crustaceans may have evolved because corals of these genera provide shelter among their branches and because these shallow-water corals are evidently capable of releasing lipid which is excess to the corals' metabolic needs, but which can be utilized by the crabs.     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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     

Sabaki River sediment load and coral stress: correlation between sediments and condition of the Malindi-Watamu reefs in Kenya (Indian Ocean)

Sediment discharges from rivers have a negative impact on coral reef ecosystems. Indicators of coral decline measured in the present study were: (1) injury to living stony corals; (2) soft coral cover; and (3) bare rocky substrate suitable for colonization by corals. The relationship between these indicators and the distribution of terrigenous sediment was studied for the Malindi-Watamu fringing reef complex along the Kenyan coast off East Africa during 1982 and 1983. Decline of this reef had been repeatedly noted during the preceding decade. The influence of terrigenous sediment from the Sabaki River appears to be strongest in the Watamu area in the south and in the northern-most part of the Malindi reef area. Correlations, between each of the above three coral stress response indicators, on the one hand, and quantitative indicators of sediment loading, on the other hand, were not clear. However, a combined coral stress indicator involving all three factors was shown to have a clear relationship with terrigenous sediment loading and provided a rapid means of field evaluation of the effects of sediment stress on stony corals. Values for the combined coral stress indicator were found to increase in proportion to increasing values of terrigenous sediment loads in both study areas. A higher coral stress indicator value means a high proportion of injured or algae infested corals, and/or a high soft coral cover, and/or a high proportion of rocky substrate suitable for, but unoccupied by, living corals.     

δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N values in reef corals <Emphasis Type="Italic">Porites lutea</Emphasis> and <Emphasis Type="Italic">P. cylindrica</Emphasis> and in their epilithic and endolithic algae

In summer 1998, shallow water corals at Sesoko Island, Japan (26°38′N, 127°52′E) were damaged by bleaching. In August 2003, partially damaged colonies of the massive Porites lutea and the branching P. cylindrica were collected at depths of 1.0–2.5 m. The species composition of epilithic algal communities on dead skeletal surfaces of the colonies (‘red turfs’, ‘green turfs’, ‘red crusts’) and the endolithic algae (living in coral skeletons) growing close to and away from living coral polyps was determined. Carbon and nitrogen stable isotope values of organic matter (δ¹³C and δ¹⁵N) from all six of these biological entities were determined. There were no significant differences in the isotope composition of coral tissues of the two corals, with P. lutea having δ¹³C of −15.3 to −9.6‰ and δ¹⁵N of 4.7–6.1‰ and P. cylindrica having similar values. Polyps in both species living close to an interface with epilithic algae had similar isotope values to polyps distant from such an interface. Despite differences in the relative abundance of the algal species in red turfs and crusts, their δ¹³C and δ¹⁵N values were not significantly different from each other (−18.2 to −13.9, −20.6 to −16.2, 1.1–4.3, and 3.3 to 4.9‰, respectively). The green algal turf had significantly higher δ¹³C values (−14.9 to −9.3‰) than that of red turfs and crusts but similar δ¹⁵N (1.2–4.1‰) to the red algae. The data do not suggest that adjoining associations of epilithic algae and coral polyps exchange carbon- and nitrogen-containing metabolites to a significant extent. The endolithic algae in the coral skeletons had δ¹³C values of −14.8 to −12.3‰ and δ¹⁵N of 4.0–5.4‰. Thus they did not differ significantly from the coral polyps in their carbon and nitrogen isotope values. The similarity in carbon isotope values between the coral polyps and endolithic algae may be attributed to a common source of CO₂ for zooxanthellae and endolithic algae, namely, from respiration by the coral host. While it is difficult to fully interpret similarity in the nitrogen isotope composition of coral tissue and of green endolithic algae and the difference in δ¹⁵N between green epilithic and endolithic algae, the data are consistent with nitrogen-containing metabolites from the scleractinian coral serving as a significant source of nitrogen for the endolithic algae.     

Serial patterns of biodiversity change in corals across shallow reef flats in Ko Phuket,Thailand, due to the effects of local (sedimentation) and regional (climatic) perturbations

A 17-year monitoring programme of reef flats at Ko Phuket, Thailand afforded an opportunity to evaluate both univariate and multivariate measures of environmental stress in an assessment of change on coral reef ecosystems. The sites at Ko Phuket suffered the effects of dredging in 1986-1987 and then anomalously low sea levels in 1997-1998 as a result of climate-related events in the Indian Ocean. Univariate measures of species diversity and taxonomic distinctness ((*) increased across the reef flat, reflecting the greater effects of physical stresses on the inner reef flats (compared with outer reef flats) at all sites, with more congeneric species present on the inner reef and more confamilial species on the outer reef. Multivariate measures showed a clear breakdown in seriation patterns at all sites during the dredging in 1987 and in 1998, as a result of earlier negative sea-level anomalies. Recovery from environmental disturbances was obvious within 12 months in each case. The domination of the reefs by massive coral species, which are physiologically adapted to intertidal living and which display partial rather than total colony mortality, may be a factor contributing to the apparent resilience of the reef flats together with continued recruitment and survival of juvenile corals during adverse environmental conditions. Elevated sea temperatures and extensive bleaching of corals in 1991, 1995, and 1998 had no effect upon coral community measures, with many corals recovering their zooxanthellae numbers within 3-5 months of the bleaching events.