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.     

Coral mortality following extreme low tides and high solar radiation

Extreme tidal events are one of the most predictable natural disturbances in marine benthic habitats and are important determinants of zonation patterns in intertidal benthic communities. On coral reefs, spring low tides are recurrent disturbances, but are rarely reported to cause mass mortality. However, in years when extremely low tides coincide with high noon irradiances, they have the potential to cause widespread damage. Here, we report on such an event on a fringing coral reef in the central Great Barrier Reef (Australia) in September 2005. Visual surveys of colony mortality and bleaching status of more than 13,000 corals at 14 reef sites indicated that most coral taxa at wave-protected sites were severely affected by the event. Between 40 and 75% of colonies in the major coral taxa (Acropora, Porites, Faviidae, Mussidae and Pocilloporidae) were either bleached or suffered partial mortality. In contrast, corals at wave-exposed sites were largely unaffected (<1% of the corals were bleached), as periodic washing by waves prevented desiccation. Surveys along a 1–9 m depth gradient indicated that high coral mortality was confined to the tidal zone. However, 20–30% of faviid colonies were bleached throughout the depth range, suggesting that the increase in benthic irradiances during extreme low tides caused light stress in deeper water. Analyses of an 8-year dataset of tidal records for the area indicated that the combination of extended periods of aerial exposure and high irradiances occurs during May–September in most years, but that the event in September 2005 was the most severe. We argue that extreme low-tide, high-irradiance events are important structuring forces of intertidal coral reef communities, and can be as damaging as thermal stress events. Importantly, they occur at a time of year when risks from thermal stress, cyclones and monsoon-associated river run-off are minimal.     

Marginal tentacles of the corallimorpharian Rhodactis rhodostoma. 2. Induced development and long-term effects on coral competitors

Polyps of the corallimorpharian Rhodactis rhodostoma (Ehrenberg, 1934) form aggregations that monopolise patches of space on the shallow reef flats of some Red Sea coral reefs. Some of these polyps bear specialised bulbous marginal tentacles (BMTs) where they contact cnidarian competitors. BMTs differ from the normally filiform marginal tentacles (FMTs) of R. rhodostoma, and appear to develop from them. However, their morphogenesis and long-term impacts on spatial competition with reef corals are unknown. We experimentally induced contacts between R. rhodostoma polyps and colonies of the branching stony coral Acropora eurystoma on a shallow coral reef at Eilat, northern Red Sea. During the first 24 d of contact, the A. eurystoma colonies extruded mesenterial filaments that damaged the tissues of the corallimorpharian polyps. After 18 d,>90% of R. rhodostoma individuals had developed BMTs, which resulted in a reversal in the direction of competitive damage. During the subsequent 1.5 years of observation, the corallimorpharians maintained well-developed BMTs, unilaterally damaged the tissues of A. eurystoma, and in some cases moved onto the stony coral skeletons and partially overgrew them. BMTs developed from FMTs in a series of four distinct stages, accompanied by significant changes in their morphology, cnidom, and density of nematocysts. Isolated control polyps did not develop BMTs or show any signs of damage. In contrast, corallimorpharian polyps transplanted into contact with colonies of the massive stony coral Platygyra daedalea began to develop sporadic BMTs, but were unilaterally and severely damaged by the corals, and started to disappear within 21 d, after the corals developed sweeper tentacles. We conclude that long-term outcomes of competition between R. rhodostoma and reef-building corals depend largely on the relative aggressive reach of the competitive mechanisms developed by each species. As a consequence, this corallimorpharian is an intermediate competitor in the aggressive hierarchy among Indo-Pacific reef corals. This study confirms that R. rhodostoma polyps may actively damage and overgrow some stony corals, leading to the formation of an almost continuous blanket of polyps in large patches of some shallow reef flats. Received: 15 July 1998 / Accepted: 24 March 1999     

Possible effects of water pollution on the community structure of Red Sea corals

The community structure and species diversity of hermatypic corals was studied during 1969–1973, in two reef flats in the northern Gulf of Eilat, Red Sea: the reef flat of the nature reserve at Eilat, which is chronically polluted by oil and minerals, and a control reef, located 5 km further south, which is free from oil pollution. In 1969, the nature reserve and the control reef had similar coral community structure. In September, 1970, both reefs suffered approximately 90% mortality of corals, as a result of an unexpected and extremely low tide. In 1973 the control reef was “blooming” with a highly diverse coral community, while almost no signs of coral recolonization have been observed at the nature reserve, and it is significantly lower in diversity. It is suggested that phosphate eutrophication and chronic oil pollution are the major man-made disturbances that interfere with coral colonization of the reef flat at the nature reserve. Although no direct evidence is provided that oil damages hermatypic corals, the data strongly suggest that chronic oil spills prevent normal settlement and/or development of coral larvae. It is possible that chronic oil, pollution results in either one or a combination of the following: (1) damage to the reproductive system of corals; (2) decreased viability of coral larvae; (3) changes in some physical properties of the reef flat which interfere with normal settlement of coral larvae.     

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.     

Aspects écologiques de la zonation intertidale sur les côtes rocheuses des îles Galapagos

Features of the intertidal zonation on the rocky shores of the Galapagos Islands are described and discussed in relation to environmental factors, in particular temperature — which varies with regional hydrology and climatology. Owing to the thermal properties of the coastal waters and to the nature of the substrate (mainly basalt), intertidal organisms are exposed to a wide range of temperatures, which fluctuate with tide, day, and season. As a result, life is mostly restricted to the lowest levels of the shore; the middle- and upper-level populations are sparse. Local variations (biofacies) resulting from water agitation are described. Biota consist of tropical, subtropical, and warm-temperate components. Tropical species dominate in sheltered areas and in middle and upper shores.

---

---

Contribution No. 164, the Charles Darwin Foundation.     

Growth of the staghorn coral Acropora formosa at Houtman Abrolhos, Western Australia

The growth (extension rate, number of radial branches, skeletal mass, branch diameter) of the␣staghorn coral Acropora formosa (Dana, 1846) was examined at four sites on the Beacon Island platform at Houtman Abrolhos, in subtropical Western Australia (28°S). Sites were at depths of 7 to 11 m, with variable exposure to weather and swell conditions. Two sites on the western reef slope were partly exposed to the oceanic swell, and two sites in the lagoon were largely protected from wave action. Linear extension rate between 1994 and 1995 varied significantly between sites, with greater linear extension at the more protected lagoonal sites. However, accumulation of skeletal mass per branch and number of newly initiated radial branches did not vary significantly between the sites. Carbonate was deposited in similar amounts, but either as porous, rapidly extending branches, or as denser branches which extended more slowly. Branch extension rate over 11.5 mo ranged from a mean of 50.3 mm (range=13 to 93 mm) at a reef slope site to a mean of 76.0 mm (range=31 to 115 mm) at a sheltered lagoonal site. Mean extension rates were almost twice that previously reported for this species in Houtman Abrolhos (37 to 43 mm yr⁻¹) from a shallower site where environmental conditions were apparently sub-optimal. Growth was within the range reported for A. formosa from tropical sites, which is consistent with the relatively high calcification and reef-accretion rates recorded for Houtman Abrolhos in geological and metabolic studies. The role of reduced coral growth-rate in limiting coral reef formation at high latitudes remains equivocal. Received: 19 November 1997 / Accepted: 5 May 1998     

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.     

A spatial model of growth and competition strategies in coral communities

A discrete spatial simulation model is developed to investigate the type and intensity of biological and physical factors influencing the structure of coral communities. The model represents reproduction, growth, and interspecific competition by coral colonies in terms of “ownership” of space in a plot of reef habitat. Using data for several eastern Pacific coral species, the model reproduces observed changes in species composition and diversity during coral community development. Model results suggest that during early successional stages, or in areas that are frequently disturbed, larval colonization and rapid growth are more important than dominance achieved by extracoelenteric digestion or by growing over another coral in acquiring and maintaining possession of reef substrate. In mature communities that remain undisturbed, dominance is the best competitive strategy. Although the model was developed to study natural and man-induced changes in the community dynamics of coral reefs, it could be adapted to study other sessile organisms where spatial pattern is an important influence on the frequency and outcome of biological interactions.     

Nitrogen- versus phosphorus-limited growth and sources of nutrients for coral reef macroalgae

Recent investigations of nutrient-limited productivity in coral reef macroalgae have led to the conclusion that phosphorus, rather than nitrogen, is the primary limiting nutrient. In this study, comparison of the dissolved inorganic nitrogen:phosphorus ratio in the water column of Kaneohe Bay, Hawaii, with tissue nitrogen:phosphorus ratios in macroalgae from Kaneohe Bay suggested that nitrogen, rather than phosphorus, generally limits productivity in this system. Results of nutrient-enrichment experiments in a flow-through culture system indicated that inorganic nitrogen limited the growth rates of 8 out of 9 macroalgae species tested. In 6 of the species tested, specific growth rates of thalli cultured in unenriched seawater from the Kaneohe Bay water column were zero or negative after 12 d. These results suggest that, in order to persist in low-nutrient coral reef systems, some macroalgae require high rates of nutrient advection or access to benthic nutrient sources in addition to nutrients in the overlying water column. Nutrient concentrations in water samples collected from the microenvironments inhabited or created by macroalgae were compared to nutrient concentrations in the overlying water column. On protected reef flats, inorganic nitrogen concentrations within dense mats of Gracilaria salicornia and Kappaphycus alvarezii, and inorganic nitrogen and phosphate concentrations in sediment porewater near the rhizophytic algae Caulerpa racemosa and C. sertularioides were significantly higher than in the water column. The sediments associated with these mat-forming and rhizophytic species appear to function as localized nutrient sources, making sustained growth possible despite the oligotrophic water column. In wave-exposed habitats such as the Kaneohe Bay Barrier Reef flat, water motion is higher than at protected sites, sediment nutrient concentrations are low, and zones of high nutrient concentrations do not develop near or beneath macroalgae, including dense Sargassum echinocarpum canopies. Under these conditions, macroalgae evidently depend on rapid advection of low-nutrient water from the water column, rather than benthic nutrient sources, to sustain growth. Received: 1 December 1997 / Accepted: 9 July 1998     

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     

Effects of emersion and temperature upon growth of the tropical brown alga Padina japonica,using a tide-simulation apparatus

A simple tide-simulation apparatus was used to investigate the influence of emersion and temperature on the intertidal growth of Padina japonica Yamada in both the juvenile and adult growth phases. The upper zonation limits are not determined by the sensitivity of any particular growth phase, since all phases show the same emersion tolerance limits. The species grows best when continuously submerged, and growth rates decrease with increasing emersion up to a clear upper zonation boundary. High temperature reduces the emersion tolerance and is therefore an important factor in tropical intertidal zonation.     

Reintroduction of eelgrass (Zostera marina) in the Dutch Wadden Sea; review of research and suggestions for management measures

Eelgrass (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal flats. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

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.     

The 1998 bleaching event and its aftermath on a coral reef in Belize

Widespread thermal anomalies in 1997-1998, due primarily to regional effects of the El Niño-Southern Oscillation and possibly augmented by global warming, caused severe coral bleaching worldwide. Corals in all habitats along the Belizean barrier reef bleached as a result of elevated sea temperatures in the summer and fall of 1998, and in fore-reef habitats of the outer barrier reef and offshore platforms they showed signs of recovery in 1999. In contrast, coral populations on reefs in the central shelf lagoon died off catastrophically. Based on an analysis of reef cores, this was the first bleaching-induced mass coral mortality in the central lagoon in at least the last 3,000 years. Satellite data for the Channel Cay reef complex, the most intensively studied of the lagoonal reefs, revealed a prolonged period of elevated sea-surface temperatures (SSTs) in the late summer and early fall of 1998. From 18 September to 1 October 1998, anomalies around this reef averaged +2.2°C, peaking at 4.0°C above the local HotSpot threshold. In situ temperature records from a nearby site corroborated the observation that the late summer and early fall of 1998 were extraordinarily warm compared to other years. The lettuce coral, Agaricia tenuifolia, which was the dominant occupant of space on reef slopes in the central lagoon, was nearly eradicated at Channel Cay between October 1998 and January 1999. Although the loss of Ag. tenuifolia opened extensive areas of carbonate substrate for colonization, coral cover remained extremely low and coral recruitment was depressed through March 2001. High densities of the sea urchin Echinometra viridis kept the cover of fleshy and filamentous macroalgae to low levels, but the cover of an encrusting sponge, Chondrilla cf. nucula, increased. Further increases in sponge cover will impede the recovery of Ag. tenuifolia and other coral species by decreasing the availability of substrate for recruitment and growth. If coral populations are depressed on a long-term basis, the vertical accretion of skeletal carbonates at Channel Cay will slow or cease over the coming decades, a time during which global-warming scenarios predict accelerated sea-level rise.     

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.     

A numerical model of wave- and current-driven nutrient uptake by coral reef communities

We developed a numerical model capable of simulating the spatial zonation of nutrient uptake in coral reef systems driven by hydrodynamic forcing (both from waves and currents). Relationships between nutrient uptake and bed stress derived from flume and field studies were added to a four-component biogeochemical model embedded within a three-dimensional (3-D) hydrodynamic ocean model coupled to a numerical wave model. The performance of the resulting coupled physical-biogeochemical model was first evaluated in an idealized one-dimensional (1-D) channel for both a pure current and a combined wave-current flow. Waves in the channel were represented by an oscillatory flow with constant amplitude and frequency. The simulated nutrient concentrations were in good agreement with the analytical solution for nutrient depletion along a uniform channel, as well as with existing observations of phosphate uptake across a real reef flat. We then applied this integrated model to investigate more complex two-dimensional (2-D) nutrient dynamics, firstly to an idealized coral reef-lagoon morphology, and secondly to a realistic section of Ningaloo Reef in Western Australia, where nutrients were advected into the domain via alongshore coastal currents. Both the idealized reef and Ningaloo Reef simulations showed similar patterns of maximum uptake rates on the shallow forereef and reef crest, and with nutrient concentration decreasing as water flowed over the reef flat. As a result of the cumulative outflow of nutrient-depleted water exiting the reef channels and then being advected down the coast by alongshore currents, both reef simulations exhibited substantial alongshore variation in nutrient concentrations. The coupled models successfully reproduced the observed spatial-variability in nitrate concentration across the Ningaloo Reef system.     

Analysis of the influence of substrate variables on coral reef fish communities

A simple field technique to obtain a gross estimate of the surface area of a quadrat on a coral reef is described. This measure, termed the substrate rugosity index, was determined, in conjunction with two other substrate variables (vertical relief and coral species richness), in a series of 4 quadrats (10 to 40 m depth) along 4 transects. The mean substrate rugosity and vertical relief of a quadrat were highly correlated. A correlation analysis was made of the substrate variables and several reef fish community parameters (species richness, number of fishes and diversity). Species richness was highly correlated with substrate rugosity. This relationship was tested in two experimental quadrats and the results were generally in accord with those predicted. Stratification of the fish communities by body size revealed that the correlation with substrate rugosity was scale-dependent. The fish community parameters were poorly correlated with percentage substrate cover by corals (ramose and glomerate) and by sand. A significant area effect was determined for two species of sand-dwelling goby.     

Extremes, plasticity, and invariance in vertebrate life history traits: insights from coral reef fishes

Life history theory predicts a range of directional generic responses in life history traits with increasing organism size. Among these are the relationships between size and longevity, mortality, growth rate, timing of maturity, and lifetime reproductive output. Spanning three orders of magnitude in size, coral reef fishes provide an ecologically diverse and species-rich vertebrate assemblage in which to test these generic responses. Here we examined these relationships by quantifying the life cycles of three miniature species of coral reef fish from the genus Eviota (Gobiidae) and compared their life history characteristics with other reef fish species. We found that all three species of Eviota have life spans of < 100 days, suffer high daily mortality rates of 7-8%, exhibit rapid linear growth, and matured at an earlier than expected size. Although lifetime reproductive output was low, consistent with their small body sizes, short generation times of 47-74 days help overcome low individual fecundity and appear to be a critical feature in maintaining Eviota populations. Comparisons with other coral reef fish species showed that Eviota species live on the evolutionary margins of life history possibilities for vertebrate animals. This addition of demographic information on these smallest size classes of coral reef fishes greatly extends our knowledge to encompass the full size spectrum and highlights the potential for coral reef fishes to contribute to vertebrate life history studies.     

Effects of eutrophication on reef-building corals

Seven fringing reef complexes were chosen along the leeward coast (west) of Barbados to study the effects of eutrophication processes upon the scleractinian coral assemblages. The structure of scleractinian coral communities was studied along an eutrophication gradient with a quantitative sampling method (line transect) in terms of species composition, zonation and diversity patterns. On the basis of these data the fringing reefs were divided into three ecological zones: back reef, reef flat, and spur and groove. Statistically discernible and biologically significant differences in scleractinian coral community structure, benthic algal cover and Diadema antillarum Philippi densities were recorded among the seven fringing reefs. High correlations between environmental variables and biotic patterns indicate that the effects of eutrophication processes (nutrient enrichment, sedimentation, turbidity, toxicity and bacterial activity) were directly and/or indirectly affecting the community structure of scleractinian coral assemblages. In general, species diversity was most sensitive in delineating among-reef, and among-zone, differences, which were attributed to intensification of eutrophication processes. Porites astreoides Lamarck, P. porites (Pallas), Siderastrea radians (Pallas), and Agaricia agaricites (Linnaeus) were the most abundant coral species in the polluted southern reefs. The absence and/or low abundance of coral species previously characterized as well adapted to high turbidity and sedimentation [i.e. Montastrea cavernosa Linnaeus, Meandrina meandrites (Linnaeus)] indicate that eutrophication processes may adversely affect these species. It is suggested that sediment rejection abilities, combined with feeding and reproductive strategies, are the primary biological processes of scleractinian corals through which eutrophication processes directly and/or indirectly affect the structure of coral communities.