Spatio-temporal variability in fish recruitment to a coral reef (Moorea,French Polynesia)

Spatial and temporal patterns of recruitment of juvenile coral reef fishes were studied on the reefs of the high island of Moorea (Society Archipelago, French Polynesia) during the wet season (October 1988 to April 1989). Two size-classes of fishes (new-recruits and juveniles) were censused by visual counts within 24 quadrats along a transect across the lagoon. Correspondence analysis was used to calculate the hypothetical movements of the two size classes. Spatial variability was far more important than temporal variability; six main spatial communities were revealed, which could result from differences among habitats. Temporal variability in recuitment occurred only at nearshore stations; stations on the outer fringing reef and inner barrier reef displayed stable recruitment patterns.     

Effects of disturbance on coral communities: bleaching in Moorea,French Polynesia

This study examines patterns of susceptibility and short-term recovery of corals from bleaching. A mass coral bleaching event began in March, 1991 on reefs in Moorea, French Polynesia and affected corals on the shallow barrier reef and to >20 m depth on the outer forereef slope. There were significant differences in the effect of the bleaching among common coral genera, with Acropora, Montastrea, Montipora, and Pocillopora more affected than Porites, Pavona, leptastrea or Millepora. Individual colonies of the common species of Acropora and Pocillopora were marked and their fate assessed on a subsequent survey in August, 1991 to determine rates of recovery and mortality. Ninety-six percent of Acropora spp. showed some degree of bleaching compared to 76% of Pocillopora spp. From March to August mortality of bleached colonies of Pocillopora was 17%, 38% recovered completely, and many suffered some partial mortality of the tissue. In contrast, 63% of the Acropora spp. died, and about 10% recovered completely. Generally, those colonies with less than 50% of the colony area affected by the bleaching recovered at a higher rate than did those with more severe bleaching. Changes in community composition four months after the event began included a significant decrease only in crustose algae and an increase in cover of filamentous algae, much of which occupied plate-like and branching corals that had died in the bleaching event. Total coral cover and cover of susceptible coral genera had declined, but not significantly, after the event.     

Recent disturbances augment community shifts in coral assemblages in Moorea, French Polynesia

Coral reefs are often subject to disturbances that can cause enduring changes in community structure and abundance of coral reef organisms. In Moorea, French Polynesia, frequent disturbances between 1979 and 2003 caused marked shifts in taxonomic composition of coral assemblages. This study explores recent changes in live cover and taxonomic structure of coral communities on the north coast of Moorea, French Polynesia, to assess whether coral assemblages are recovering (returning to a previous Acropora-dominated state) or continuing to move towards an alternative community structure. Coral cover declined by 29.7% between July 2003 and March 2009, mostly due to loss of Acropora and Montipora spp. Coral mortality varied among habitats, with highest levels of coral loss on the outer reef slope (7–20 m depth). In contrast, there was limited change in coral cover within the lagoon, and coral cover actually increased on the reef crest. Observed changes in coral cover and composition correspond closely with the known feeding preferences and observed spatial patterns of Acanthaster planci L., though observed coral loss also coincided with at least one episode of coral bleaching, as well as persistent populations of the corallivorous starfish Culcita novaeguineae Muller & Troschel. While climate change poses an important and significant threat to the future structure and dynamics coral reef communities, outbreaks of A. planci remain a significant cause of coral loss in Moorea. More importantly, these recent disturbances have followed long-term shifts in the structure of coral assemblages, and the relative abundance of both Pocillopora and Porites continue to increase due to disproportionate losses of Acropora and Montipora. Moreover, Pocillopora and Porites dominate assemblages of juvenile corals, suggesting that there is limited potential for a return to an Acropora-dominated state, last recorded in 1979.     

Spatiotemporal distribution of nocturnal coral reef fish juveniles in Moorea Island,French Polynesia

This study aimed to investigate the spatial structure of nocturnal fish communities at settlement on coral reefs in Moorea Island lagoon, French Polynesia; and the temporal consistency of habitat selection between winter (April to June 2001) and summer (November 2001). The Moorea lagoon was divided into 12 habitat zones (i.e., coral reef zones), which were distinct in terms of depth, wave exposure, and substratum composition. Nocturnal visual censuses among the 12 habitats found that the recently settled juveniles of 25 species recorded were dispatched among three communities spatially distributed according to the distance from the reef crest (reef crest, barrier reef, and fringing reef communities). This spatial communities structure of nocturnal juveniles was consistent in both winter and summer and would be explained primarily by a current gradient in Moorea lagoon (current speed was high near the reef crest and decreased towards the beach) and by the topographic characteristics of reef zones. Among the 25 species, 13 were recorded in both winter and summer. A comparison of the spatial distribution between summer and winter for 13 species that occurred during both seasons found that only 4 differed between the two seasons. For these species, habitat selection would be organized primarily by some stochastic processes such as inter- and intraspecific competition, predation, and food availability. Overall, the present study allowed us to highlight that most nocturnal coral reef fish juveniles at Moorea Island exhibited striking patterns in their distribution and current and topographic characteristics of reef zones might exert considerable influence on the distribution of fishes.     

High spatial variability in coral bleaching around Moorea (French Polynesia): patterns across locations and water depths

Mass coral bleaching events are one of the main threats to coral reefs. A severe bleaching event impacted Moorea, French Polynesia, between March and July 2002, causing 55+/-14% of colonies to suffer bleaching around the island. However, bleaching varied significantly across coral genera, locations, and as a function of water depth, with a bleaching level as high as 72% at some stations. Corals in deeper water bleached at a higher rate than those in shallow water, and the north coast was more impacted than the west coast. The relatively small scale of variability in bleaching responses probably resulted from the interaction between extrinsic factors, including hydrodynamic condition, and intrinsic factors, such as differential adaptation of the coral/algal association.     

Zonation of macrobenthic communities along two bays in an insular coral reef ecosystem (Moorea, French Polynesia)

Zonation of macrobenthic communities (corals, algae, molluscs, sponges and echinoderms) on fringing reefs was investigated in two bays in a coral reef ecosystem (Moorea, French Polynesia). Species richness, abundance and coral cover, and species richness of macroalgae increased from the bayhead to the bay entrance. For molluscs and sponges, no general trend was observed. Species richness and abundance of echinoderms increased from the head to the entrance of Opunohu Bay, but this trend was less pronounced in Cook Bay. The gradients observed for corals, macroalgae and echinoderms were correlated with one or several of the following abiotic factors: salinity, turbidity, concentration of silicates in surface waters, and concentrations of organic carbon, carbohydrates and amino acids in the sediments. These factors are associated with terrestrial run-off via river discharge that occurs at the bayheads, where the major river is located. The high degree of confinement in the bayheads allowed the establishment of only a few tolerant macrobenthic species. Although the low diversity and abundance of corals and echinoderms seem to be a characteristic of Polynesian bays, a high diversity of corals can be found in the vicinity of the bayheads in coral reefs of the western Pacific.     

Direct measurements of carbon and carbonate export from a coral reef ecosystem (Moorea Island , French Polynesia)

The export of carbon and carbonate from coral reefs was investigated through a multidisciplinary investigation of the hydrological, geochemical, sedimentological and biological features of Tiahura reef on the northwestern coast of Moorea Island (French Polynesia). The hydrology of the fore-reef is characterised by prevailing longshore western currents and a strong thermocline. As revealed by turbidity structures (benthic and intermediate nepheloid layers) and by the amount of particles collected by near-bottom sediment traps, horizontal and downslope advections of particles dominate over offshore vertical transport. The exported material is rich in carbonate (ca. 80%) and poor in organic matter (ca. 4%). Sedimentation rates at 430 m depth, i.e. definitive export, reached 209.6 mg m^-2 d^-1 (dry weight). Estimates of carbon and carbonates export for Tiahura reef also reported here represent respectively 47% and 21% of the organic and inorganic carbon produced within the reef.     

Distributions of coral reef macroalgae in a back reef habitat in Moorea,French Polynesia

On tropical reefs where macroalgae are subjected to continuous herbivore pressure, spatial refuges typically are identified as large-scale, landscape interfaces that limit foraging behavior. However, algal distributions and community assemblages may also rely on the availability of smaller scale spatial refuges within the reef. The results of this study demonstrate that the patterns of macroalgal distribution across the back reef of Moorea, French Polynesia, are maintained by herbivores interacting with the small-scale structural complexities of the coral reef landscape. Although the majority of space available for colonization is composed of exposed surfaces, macroalgae rarely are found in the open. Instead, macroalgal occurrence is highest in the protected narrow crevices and hole microhabitats provided by massive Porites spp. coral heads. These distributions are determined initially by post-settlement mortality of young algal recruits in exposed habitats. Rates of consumption for two of the most common macroalgal species found in refuges across the back reef, Halimeda minima and Amansia rhodantha, indicate that algal recruits in exposed habitats are limited by herbivory. While algal abundance and community structure are highly dependent upon herbivore grazing, the availability of small-scale spatial refuges ultimately shapes the distinct community patterns and distributional boundaries of coral reef macroalgae in the back reefs of Moorea.     

Spatio-temporal heterogeneity in coral recruitment around Moorea, French Polynesia: Implications for population maintenance

The spatio-temporal variability of scleractinian coral recruitment was investigated from December 2000 to December 2003 around Moorea Island (French Polynesia). Nine stations, each with 20 terracotta tiles, were placed on the outer reef slope, at 3 sites (Vaipahu, Tiahura, Haapiti), and at 3 depths (6, 12 and 18 m). The relative contribution of the different families of recruits (Pocilloporidae: 60.4%, Poritidae: 18.8%, Acroporidae: 11.2%) and the very low recruitment rates (maximum: 35 recruits per tile, higher mean densities of 10.8 recruits per tile, average 40.77 recruits m^− 2 year^− 1) recorded at Moorea are similar to recruitment patterns recorded on sub-tropical reefs. Over the duration of the study period, we detected a marked seasonal variability in recruitment rates, with the peak recruitment for all families recorded in December-March periods, corresponding to periods of warmest SSTs. However, recruits of Acroporidae were also relatively abundant in the September-December period in some years, which coincides with the known spawning periods of some Acropora species. Total recruitment rate decreased after the first year of the survey, and was probably the result of the bleaching event that occurred in early 2002, which may have reduced fecundity of some coral populations. A lower proportion of recruits were found on the upper surface of the tiles (14.5%), compared to the lower surfaces (57.1%) and sides (28.4%), which is likely the result of intense grazing by herbivorous fish and urchins. We detected a patchy distribution at the station scale, and a significant variation in recruitment patterns among depths and sites. Pocilloporidae recruited more abundantly at 6 and 18 m, whereas Poritidae were generally more abundant at 12 m depth. In contrast, Acroporidae showed no clear depth pattern during the study period. Recruitment was lowest at the most exposed site (Haapiti), especially for Acroporidae, and probably reflects lower settlement rates and/or higher early post-settlement mortality caused by frequent high swells and their associated strong currents. These distinctive characteristics in recruitment patterns of the 3 dominant families of recruits underline the important role of life history strategies in understanding the spatial patterns, dynamics and maintenance processes of coral populations. The relatively low coral recruitment rates recorded from this study indicate that recovery from severe or frequent perturbations will be slow.     

Regional and local variability in recovery of shallow coral communities: Moorea,French Polynesia and central Great Barrier Reef

Coral communities at Moorea, French Polynesia, and on the Great Barrier Reef (GBR), Australia, were severely depleted by disturbances early in the 1980s. Corals were killed by the predatory starfish Acanthaster planci, by cyclones, and/or by depressed sea level. This study compares benthic community structure and coral population structures on three disturbed reefs (Vaipahu-Moorea; Rib and John Brewer Reefs-GBR) and one undisturbed reef (Davies Reef-GBR) in 1987–89. Moorea barrier reefs had been invaded by tall macrophytes Turbinaria ornata and Sargassum sp., whereas the damaged GBR reefs were colonised by a diverse mixture of short macrophytes, turfs and coralline algae. The disturbed areas had broadly similar patterns of living and dead standing coral, and similar progress in recolonisation, which suggests their structure may converge towards that of undisturbed Davies Reef. Corals occupying denuded areas at Vaipahu, Rib and John Brewer were small (median diameter 5 cm in each case) and sparse (means 4–8 m^-2) compared to longer established corals at Davies Reef (median diameter 9 cm; mean 18 m^-2). At Moorea, damselfish and sea urchins interacted with corals in ways not observed in the GBR reefs. Territories of the damselfish Stegastes nigricans covered much of Moorea's shallow reef top. They had significantly higher diversity and density of post-disturbance corals than areas outside of territories, suggesting that the damselfish exerts some influences on coral community dynamics. Sea urchins on Moorea (Diadema setosum Echinometra mathaei, Echinotrix calamaris) were causing widespread destruction of dead standing coral skeletons. Overall, it appears that the future direction and speed of change in the communities will be explicable more in terms of local than regional processes.     

Sexual reproduction of Acropora reef corals at Moorea, French Polynesia

Little information is available on reproductive processes among corals in isolated central Pacific reef regions, including French Polynesia. This study examined the timing and mode of sexual reproduction for Acropora reef corals at Moorea. Spawning was observed and/or inferred in 110 Acropora colonies, representing 12 species, following full moon periods in September through November 2002. Gamete release was observed and inferred in four species of Acropora between 9 and 13 nights after the full moon (nAFM) in September 2002. Twelve Acropora spp. spawned gametes between 5 and 10 nAFM in October 2002, with six species spawning 7 nAFM and four species spawning 9 nAFM. In November 2002, spawning of egg and sperm bundles was observed and inferred in 27 colonies of Acropora austera, 6 nAFM. These are the first detailed records of spawning by Acropora corals in French Polynesia.     

Genetic evidence of closed life-cycles for some coral reef fishes within Taiaro Lagoon (Tuamotu Archipelago, French Polynesia)

 Taiaro Lagoon has no permanent or regular connection with the ocean (ingress is restricted to episodes of high sea-level and/or strong wave action) raising the question of how fish populations with normally dispersive larvae are maintained inside this lagoon. We compared the genetic population structures of two coral reef fishes, Acanthurus triostegus and Chaetodon ulietensis, present on both sides of the atoll rim to determine whether there is evidence of reproductive isolation. Genetic surveys showed that the lagoonal and oceanic populations were statistically different at five loci (AAT ^* -3, GDA ^*, HPD ^*, MDH ^* and SDH ^*) in A. triostegus and three loci (PGI-2^*, IDH ^* and PGD ^*) in C. ulietensis, producing high F _st values of 0.055 and 0.021, respectively. Our genetic and demographic data on these species suggest that both may be completing their life-cycles inside the lagoon, which leads us to question the common assumption that coral reef fishes require oceanic conditions for larval development. Accepted: 28 August 1997     

Identical digeneans in coral reef fishes from French Polynesia and the Great Barrier Reef (Australia) demonstrated by morphology and molecules

Three coral reef fish species, Zanclus cornutus, Chaetodon vagabundus and Naso lituratus, were collected in French Polynesia and on the Great Barrier Reef, Queensland. These fish species were each infected by one morphologically similar digenean species in both localities; Schistorchis zancli Hanson, 1953 was found in Zanclus cornutus, Preptetos laguncula Bray and Cribb, 1996 in Naso lituratus and Neohypocreadium dorsoporum Machida and Uchida, 1987 in Chaetodon vagabundus. In addition, on the Great Barrier Reef P. laguncula was also found in Naso unicornis and N. dorsoporum in Chaetodon ephippium and Chaetodon flavirostris. Morphometric differences between the species from the two sites were only slight. Sequences from the second internal transcribed spacer of the ribosomal DNA of each worm revealed total homology or negligible divergence between samples from hosts caught in French Polynesia and on the Great Barrier Reef. These results show that across more than 6000 km these digeneans are similar in morphology and genotype. Some species of fishes and molluscs are considered to have distributions that encompass the entire tropical Indo-West Pacific. These findings suggest that at least some of their parasites have similarly broad distributions.     

Changes in Bleaching Susceptibility among Corals Subject to Ocean Warming and Recurrent Bleaching in Moorea,French Polynesia

Background

Climate-induced coral bleaching poses a major threat to coral reef ecosystems, mostly because of the sensitivities of key habitat-forming corals to increasing temperature. However, susceptibility to bleaching varies greatly among coral genera and there are likely to be major changes in the relative abundance of different corals, even if the wholesale loss of corals does not occur for several decades. Here we document variation in bleaching susceptibility among key genera of reef-building corals in Moorea, French Polynesia, and compare bleaching incidence during mass-bleaching events documented in 1991, 1994, 2002 and 2007.

Methodology/Principal Findings

This study compared the proportion of colonies that bleached for four major genera of reef-building corals (Acropora, Montipora, Pocillopora and Porites), during each of four well-documented bleaching events from 1991 to 2007. Acropora and Montipora consistently bleached in far greater proportions (up to 98%) than Pocillopora and Porites. However, there was an apparent and sustained decline in the proportion of colonies that bleached during successive bleaching events, especially for Acropora and Montipora. In 2007, only 77% of Acropora colonies bleached compared with 98% in 1991. Temporal variation in the proportion of coral colonies bleached may be attributable to differences in environmental conditions among years. Alternately, the sustained declines in bleaching incidence among highly susceptible corals may be indicative of acclimation or adaptation.

Conclusions/Significance

Coral genera that are highly susceptible to coral bleaching, and especially Acropora and Montipora, exhibit temporal declines in their susceptibility to thermal anomalies at Moorea, French Polynesia. One possible explanation for these findings is that gradual removal of highly susceptible genotypes (through selective mortality of individuals, populations, and/or species) is producing a coral assemblage that is more resistant to sustained and ongoing ocean warming.     

Temporal variations of macroborers in massive Porites lobata on Moorea,French Polynesia

Massive colonies of Porites lobata on the barrier reef of Tiahura, Moorea, can be divided into four categories: living colonies, colonies consisting of 50% live coral and 50% dead skeleton, 100% dead coral and colonies which have been reduced to a basal plate. Replicate samples of each of these colony types were collected in the same vicinity of the barrier reef during October 1987. The macroborers were extracted, identified, counted and their volumes determined by displacement. Kruskal-Wallis tests showed that three different boring communities occur within these four categories of Porites colonies. Live colonies are characterised by only 3 species, the bivalve, Lithophaga laevigata; the vermetid Dendropoma maximun; and the non boring serpulid polychaete Spirobranchus. The completely dead colonies contain up to 17 boring species, with five to six individuals per 100 cm. Sipunculans are the dominant bioeroders with the most abundant species being Aspidosiphon elegans, sp A and sp B. Colonies of Porites which have been reduced to basal plates contain up to 18 boring species of which the bivalve Lithophaga hanleyana and the sipunculan Aspidosiphon sp. B are the most abundant.The cumulative volume of CaCO₃ lost by boring activity increases from 0.1 cm³ per 100 cm³ in a completely dead Porites colony to 1.4 cm³ per 100 cm³ in the residual basal plates of Porites. These can be extrapolated to minimum losses of 14.2 kg m^-3. We suggest that rates of boring increase with the time which has elapsed since the death of the colony and the dominant agents of boring also change with increasing age of the coral structure. There are significant additional losses of 5.25 kg m^-3 CaCO₃ caused by grazing echinoids and scarids.     

The significance of exposed coral conglomerates from French Polynesia (Pacific Ocean) as indicators of recent relative sea-level changes

In many Pacific islands, reef tracts possess exposed coral conglomerates ranging in age from the present to about 6,000 years B.P. These reach a maximum elevation of more than 1 metre above the present mean tide level. The outcrops occur mainly as subhorizontal rampart-platforms of poorly-sorted, cemented, skeletal detritus deposited by storm processes. With the aim of clarifying their significance as indicators of Recent sea level changes, we have attempted to determine the environments of lithification of these outcrops. To this end, rock specimens were collected from seaward vertical profiles at various levels of outcrops on French Polynesian coral and volcanic islands. For each outcrop, two sequences of marine cementation could be defined on the basis of petrological criteria: (1) a lower sequence exhibiting at least three generations of cements. The earlier generations are characterized by high amounts of high-magnesian calcite pelleted micrite normally completely occluding interparticle pore spaces. The later generation is characterized by the generalized occurrence of irregular rims of truncated aragonite fibers and pendant microstalactites. Residual interparticle porosity is relatively low; (2) an upper sequence mainly exhibiting blunted argonite needles and microstalactites; residual porosity is relatively high. The diagenetic limit between these two sequences defines the position of a former marine water table that is regarded as having been closely related to a former mean low tide level. All this seems to indicate that the lower sequence has spent time in two distinctly different diagenetic environments (phreatic overlain by vadose), whereas the upper sequence has always been within one diagenetic zone (vadose). Thus some relative fall in water table level has occurred after the deposition and early lithification of the lower deposits. Such conglomerates are believed to be useful indicators of former sea stands, particularly interesting in the case of islands devoid of other markers of Recent emergence.     

Spatial Patterns in the Distribution,Diversity and Abundance of Benthic Foraminifera around Moorea (Society Archipelago,French Polynesia)

Coral reefs are now subject to global threats and influences from numerous anthropogenic sources. Foraminifera, a group of unicellular shelled organisms, are excellent indicators of water quality and reef health. Thus we studied a set of samples taken in 1992 to provide a foraminiferal baseline for future studies of environmental change. Our study provides the first island-wide analysis of shallow benthic foraminifera from around Moorea (Society Archipelago). We analyzed the composition, species richness, patterns of distribution and abundance of unstained foraminiferal assemblages from bays, fringing reefs, nearshore and back- and fore-reef environments. A total of 380 taxa of foraminifera were recorded, a number that almost doubles previous species counts. Spatial patterns of foraminiferal assemblages are characterized by numerical abundances of individual taxa, cluster groups and gradients of species richness, as documented by cluster, Fisher α, ternary plot and Principal Component Analyses (PCA). The inner bay inlets are dominated by stress-tolerant, mostly thin-shelled taxa of Bolivina, Bolivinella, Nonionoides, Elongobula, and Ammonia preferring low-oxygen and/or nutrient-rich habitats influenced by coastal factors such as fresh-water runoff and overhanging mangroves. The larger symbiont-bearing foraminifera (Borelis, Amphistegina, Heterostegina, Peneroplis) generally live in the oligotrophic, well-lit back- and fore-reef environments. Amphisteginids and peneroplids were among the few taxa found in the bay environments, probably due to their preferences for phytal substrates and tolerance to moderate levels of eutrophication. The fringing reef environments along the outer bay are characterized by Borelis schlumbergeri, Heterostegina depressa, Textularia spp. and various miliolids which represent a hotspot of diversity within the complex reef-lagoon system of Moorea. The high foraminiferal Fisher α and species richness diversity in outer bay fringing reefs is consistent with the disturbance-mosaic (microhabitat heterogeneity) hypothesis.Calculations of the FORAM Index (FI), a single metric index to assess reef vitality, indicate that all fore- and most back-reef environments support active carbonate accretion and provide habitat suitability for carbonate producers dependent on algal symbiosis. Lowest suitability values were recorded within the innermost bays, an area where natural and increasing anthropogenic influences continue to impact the reefs. The presence of habitat specific assemblages and numerical abundance values of individual taxa show that benthic foraminifera are excellent recorders of environmental perturbations and good indicators useful in modern and ancient ecological and environmental studies.