ECOLOGY AND MORPHOLOGY OF RECENT CORAL REEFS

1. The classical ‘coral reef problem’ concerned the geological relationships of reefs as major topographical features; modern coral studies consider reefs both as complex biological systems of high productivity and as geological structures forming a framework for and being modified by coral growth. 2. Deep borings in reefs have conclusively confirmed the general arguments of Darwin, that oceanic reefs developed by progressive subsidence of their foundations. Darwin failed to take account of Pleistocene changes in sea level and their effect on the present surface features of reefs. Daly's alternative ‘glacial control theory’ was based on false assumptions concerning marine erosion rates during glacial periods, but if sea level during the Holocene was higher than at present, as Daly also supposed, the effects on reef features would be profound. 3. Reefs are complex biological systems in tropical seas, dominated by scleractinian corals. Coral faunas are larger and more diverse in the Indo-Pacific than in the Atlantic. Hermatypic corals are restricted to shallow water by the light requirements of their symbiotic algae, but temperature is a major control of worldwide distributions. Temperature, salinity and sediment tolerances of corals are wider than formerly supposed, and corals can survive brief emersion except when it coincides with heavy rainfall. Water turbulence is an important ecological control, but difficult to measure. 4. The trophic status of corals is still unclear, but in spite of their anatomical and physiological specialization as carnivores it is likely that they derive some nutrient substances from zooxanthellae. Suggestions that filamentous algae in coral heads play a major part in the economy of the corals have not been supported by later work, but biomass pyramids constructed on the basis by Odum and Odum remain the only ones available. Most reefs are apparently autotrophic, with 1500–3500 g. Carbon being fixed per m.² per year. 5. Few animals eat corals, which may account for their success. Important predators are fish and the echinoderm Acanthaster. Quantitative estimates of biogenic erosion of organic skeletons on reefs are high. Fish affect not only corals but other invertebrates, algae and marine phanerogams. 6. Corals may be killed by ‘dark water’, intense rain or river floodwaters, earth movements, human interference and especially hurricanes. Reef recovery after hurricanes may take 10–20 years. 7. In addition to fringing, barrier and atoll reefs, intermediate types are recognised. The main types may consist of linear reefs or faros. Smaller lagoon reefs include pinnacles, patches and platforms, and submerged knolls. Complex cellular or mesh reef patterns are also found. 8. Reefs are conspicuously zoned, both laterally in response to changing exposure to waves to form windward and leeward reefs, and transversely, as a result of steep environmental gradients across reef flats from sea to lagoon. Topographic and ecological zones may be characterized by particular coral species, but these vary widely from reef to reef. A major distinction can be made between reefs with and without algal ridges, which are common on open-ocean trade-wind reefs, in the Indo-Pacific, but are absent on Caribbean reefs and on Indo-Pacific reefs in more sheltered waters. gorgonians are common on Caribbean reefs, alcyonaceans in the Indo-Pacific. 9. Much of the difficulty in comparing reefs stems from the lack of uniformity in surveying methods. Problems of describing the complex three-dimensional patterns of organisms on reefs have yet to be solved, and hence little progress has been made in explanation of these patterns. Explanation in terms of simple environmental controls is inadequate. 10. Understanding the distribution of corals is made more difficult both by taxo-nomic problems and by the plasticity of growth form in different situations. 11. Growth of corals and reefs may be estimated by measuring the growth of individual colonies, measuring rates of calcium carbonate deposition in the skeleton, measuring topographic change on the reef and deducing net rates of reef growth from geological evidence. Massive corals may increase in diameter by 1 cm./year, branches of branching corals may increase in length by 10 cm./year. Study of deposition rates shows variation within colonies, between species, in light and dark, and seasonally. Rates of reef growth extrapolated from colony measurements reach 2–5 cm./year, and contrast with figures as low as 0–02 cm/year averaged over 70 million years from borehole data. Both colony growth rates and geological data suggest worldwide variations in rates of reef growth. 12. In spite of clear evidence of long-continued subsidence, present surface features of reefs, often only thinly veneered by modern corals, have been much affected by recent sea level fluctuations. Many slightly raised reefs at 2–10 m. above sea level date at 90–160 thousand years B.P.; there is evidence for a sea level at about the present level at 30–35 thousand years B.P.; and controversy continues over whether sea level has stood higher than the present at any time since the last sea level rise began about 20,000 years ago. Evidence from many reefs suggests a slightly higher sea level in the last 4000 years, but on other reefs such evidence is lacking. 13. Several reef features (submerged terraces, groove-spur systems, algal ridge, reef flat, reef blocks and reef islands) have been interpreted either as relict features dating from a higher sea level in the last 5000 years, or contemporary features developed in response to present processes. In some cases the evidence is equivocal; in others it is clear that diverse features are being grouped together under the same name. If such features are referable to a higher sea level, this may have been of last Interglacial or even Interstadial age rather than Holocene. 14. A reef consists of a rigid framework defining several major depositional environments within and around it. Sediments are of biological, mainly skeletal origin, except in unusual environments such as the Bahama Banks. The characteristics of sediments derived from organisms depend partly on the breakdown patterns of particular skeletons, partly on transportation and sorting processes. Fine sediments may be either detrital, or physicochemical precipitates. 15. Organisms affect sediments after deposition, by disturbance, transportation and probably comminution. Fish and holothurians have been studied in detail. 16. While new theories of coral reefs are proposed from time to time, the need is less for new theories than for standardised procedures to ensure comparability of reef studies and the identification of variations in reefs both on local and regional scales. While reefs as biological systems adjust relatively rapidly to changes, reefs as geological systems adjust much more slowly. Because of the magnitude and recency of Pleistocene fluctuations in sea level, many biological features of reefs are out of phase with inherited geological features, and this had led to much controversy.     

Sponge-mediated coral reef growth and rejuvenation

Sponges mediate consolidation of Porites furcata rubble on shallow Caribbean reefs by quickly adhering to rubble and stabilizing it until carbonate secreting organisms can grow and consolidate it to the reef. Experimental investigations demonstrate that the entire cycle from (1) temporary binding of rubble by sponges, through (2) rubble consolidation by encrusting coralline algae, to (3) colonization of consolidated rubble by corals, can be completed within 10 months. Bound rubble both adds to vertical reef growth and also provides stable substrata for colonization by corals. Corals that colonize stabilized rubble are damaged less and survive better than on unstable rubble. Rubble that is not temporarily stabilized by sponges does not become bound to the reef, because continuous movement disturbs the consolidation process, and does not provide suitable substrata for settlement and growth of corals. Sponge-mediated consolidation of rubble may increase rates of reef growth and enhance reef recovery after damage. This new role for sponges in reef growth is not obvious from examination of the internal fabric of a reef frame. Spongemediated consolidation may help to explain geographic and temporal differences in growth and morphology among shallow reefs of ramose corals.     

Indo-Pacific and Atlantic spurs and grooves revisited: the possible effects of different Holocene sea-level history,exposure, and reef accretion rate in the shallow fore reef

Shallow fore-reef areas worldwide are usually characterized by spurs and grooves. A comparison of examples from the three world oceans suggests that Indo-Pacific spurs and grooves are shaped predominantly by erosion, whereas western Atlantic spur and groove systems are largely a product of constructive processes. I propose that this difference is caused by regional differences in Holocene sea-level change, which controlled exposure to waves and currents, and reef-accretion rates. The transgressive–regressive sea-level curve in the Indo-Pacific realm, i.e., the Mid-to-Late Holocene sea-level fall in these areas has maintained high-energy conditions in the shallow fore reef. Higher exposure to waves and currents favors erosion and leads to a dominance of crustose coralline algae that have relatively slow growth rates. In the western Atlantic, the transgressive Holocene sea level has caused Mid-to-Late Holocene deepening and has maintained accommodation space for reef accretion. Fast-growing acroporid corals thrive under lower exposure and are more common than coralline algae. The fossil record of the spur and groove system is rather poor, which is probably a consequence of the need of excellent, three-dimensional outcrops for identification.     

Coral growth with thermal stress and ocean acidification: lessons from the eastern tropical Pacific

The rapid growth of scleractinian corals is responsible for the persistence of coral reefs through time. Coral growth rates have declined over the past 30 years in the western Pacific, Indian, and North Atlantic Oceans. The spatial scale of this decline has led researchers to suggest that a global phenomenon like ocean acidification may be responsible. A multi-species inventory of coral growth from Pacific Panamá confirms that declines have occurred in some, but not all species. Linear extension declined significantly in the most important reef builder of the eastern tropical Pacific, Pocillopora damicornis, by nearly one-third from 1974 to 2006. The rate of decline in skeletal extension for P. damicornis from Pacific Panamá (0.9% year⁻¹) was nearly identical to massive Porites in the Indo-Pacific over the past 20–30 years (0.89–1.23% year⁻¹). The branching pocilloporid corals have shown an increased tolerance to recurrent thermal stress events in Panamá, but appear to be susceptible to acidification. In contrast, the massive pavonid corals have shown less tolerance to thermal stress, but may be less sensitive to acidification. These differing sensitivities will be a fundamental determinant of eastern tropical Pacific coral reef community structure with accelerating climate change that has implications for the future of reef communities worldwide.     

Late Holocene reef growth and relative sea-level changes in Atol das Rocas, equatorial South Atlantic

Shallow drilling provided the first detailed record of vertical reef accretion rates for the last 4,000 years from the oceanic atoll Atol das Rocas. Six cores up to1-m long from windward, leeward, and intertidal hardground environments were radiocarbon dated. Frameworks are dominated by the coralline alga Porolithon cf. pachydermum with minor contributions of Lithophyllum sp. Coralline bindstone and framestone facies were identified. Vertical accretion rates (VAR) form three groups: group A frameworks were formed between 3,490±45 years BP and 2,770±45 years BP, and VAR are 0.85, 1.4, and 1.6 mm/year; group B frameworks were formed between 2,510±45 year BP and 490±45 year BP, and VAR are 0.25, 0.46, and 0.42 mm/year; group C frameworks were formed between 900±50 year BP and 655±45 year BP, and VAR are 3.2, 9.75, and 18.4 mm/year. Results indicate that coralline-algal reefs may display a catch-down response to a falling sea level similar to the way corals respond to a rising sea level. In this case, present day reef topography may be the result of late Holocene SW Atlantic sea-level changes. The calculated VAR of 18.4 mm/year is the highest rate known to date for a coralline-algal reef and close to the maximum rates recorded for corals.     

Estimation of photosynthesis and calcification rates at a fringing reef by accounting for diurnal variations and the zonation of coral reef communities on reef flat and slope: a case study for the Shiraho reef, Ishigaki Island, southwest Japan

Seven coral reef communities were defined on Shiraho fringing reef, Ishigaki Island, Japan. Net photosynthesis and calcification rates were measured by in situ incubations at 10 sites that included six of the defined communities, and which occupied most of the area on the reef flat and slope. Net photosynthesis on the reef flat was positive overall, but the reef flat acts as a source for atmospheric CO₂, because the measured calcification/photosynthesis ratio of 2.5 is greater than the critical ratio of 1.67. Net photosynthesis on the reef slope was negative. Almost all excess organic production from the reef flat is expected to be effused to the outer reef and consumed by the communities there. Therefore, the total net organic production of the whole reef system is probably almost zero and the whole reef system also acts as a source for atmospheric CO₂. Net calcification rates of the reef slope corals were much lower than those of the branching corals. The accumulation rate of the former was approximately 0.5 m kyr⁻¹ and of the latter was ~0.7–5 m kyr⁻¹. Consequently, reef slope corals could not grow fast enough to keep up with or catch up to rising sea levels during the Holocene. On the other hand, the branching corals grow fast enough to keep up with this rising sea level. Therefore, a transition between early Holocene and present-day reef communities is expected. Branching coral communities would have dominated while reef growth kept pace with sea level rise, and the reef was constructed with a branching coral framework. Then, the outside of this framework was covered and built up by reef slope corals and present-day reefs were constructed.     

Cycles of coral reef ‘turn‐on’, rapid growth and ‘turn‐off’ over the past 8500 years: a context for understanding modern ecological states and trajectories

Human activities threaten reef ecosystems globally, forcing ecological change at rates and scales regarded as unprecedented in the Holocene. These changes are so profound that a cessation of reef accretion (reef ‘turn‐off’) and net erosion of reef structures is argued by many as the ultimate and imminent trajectory. Here, we use a regional scale reef growth dataset, based on 76 core records (constrained by 211 radiometric dates) from 22 reefs along and across the inner‐shelf of the Great Barrier Reef, Australia, to examine the timing of different phases of reef initiation (‘turn‐on’), growth and ‘turn‐off’ during the Holocene. This dataset delineates two temporally discrete episodes of reef‐building over the last 8500 years: the first associated with the Holocene transgression‐early highstand period [～8.5–5.5 k calibrated years bp (cal ybp )]; the second since ～2.3 k cal ybp . During both periods, reefs accreted rapidly to sea level before entering late evolutionary states – states naturally characterized by reduced coral cover and low accretion potential – and a clear hiatus occurs between these reef‐building episodes for which no records of reef initiation exist. These transitions mimic those projected under current environmental disturbance regimes, but have been driven entirely by natural forcing factors. Our results demonstrate that, even through the late Holocene, reef health and growth has fluctuated through cycles independent of anthropogenic forcing. Consequently, degraded reef states cannot de facto be considered to automatically reflect increased anthropogenic stress. Indeed, in many cases degraded or nonaccreting reef communities may reflect past reef growth histories (as dictated by reef growth–sea level interactions) as much as contemporary environmental change. Recognizing when changes in reef condition reflect these natural ‘turn‐on’– growth –‘turn‐off’ cycles and how they interact with on‐going human disturbance is critical for effective coral reef management and for understanding future reef ecological trajectories.     

Competition between corals and algal turfs along a gradient of terrestrial influence in the nearshore central Great Barrier Reef

Competition between benthic algae and corals is a key process in the community ecology of reefs, especially during reef degradation. However, there have been very few experimental tests for competition between corals and benthic algae, despite widespread assumptions that algae are generally superior competitors, especially in eutrophic conditions. This study tested for competition for space between the massive coral Porites lobata and algal filamentous turfs on three reefs along a cross-shelf gradient of terrestrial influence, by experimentally removing or damaging either corals or algae. The corals and algae were competing for space, but, significantly, the algae appeared to have little effect on coral growth. In contrast, corals significantly inhibited algal growth, suggesting Porites was the competitive superior. Importantly, coral growth was generally positive, even on the reef with the greatest terrestrial influence. Competitive outcomes did not support the argument that algae are more successful competitors in more eutrophic conditions.     

Holocene coral reef accretion in Hawaii: a function of wave exposure and sea level history

 In the high Hawaiian Islands, significant accretion due to coral reef growth is limited by wave exposure and sea level. Holocene coral growth and reef accretion was measured at four stations off Oahu, Hawaii, chosen along a gradient in wave energy from minimum to maximum exposures. The results show that coral growth of living colonies (linear extension) at optimal depths is comparable at all stations (7.7–10.1 mm/y), but significant reef accretion occurs only at wave sheltered stations. At wave sheltered stations in Hanauma Bay and Kaneohe Bay, rates of long term reef accretion are about 2.0 mm/y. At wave exposed stations, off Mamala Bay and Sunset Beach, reef accretion rates are virtually zero in both shallow (1 m) and deeper (optimal) depths (12 m). At wave sheltered stations, such as Kaneohe Bay and Hanauma Bay, Holocene reef accretion is on the order of 10–15 m thick. At wave exposed stations, Holocene accretion is represented by only a thin veneer of living corals resting on antecedent Pleistocene limestone foundations. Modern coral communities in wave exposed environments undergo constant turnover associated with mortality and recruitment or re-growth of fragmented colonies and are rarely thicker than a single living colony. Breakage, scour, and abrasion of living corals during high wave events appears to be the major source of mortality and ultimately limits accretion to wave sheltered environments. Depth is particularly important as a modulator of wave energy. The lack of coral reef accretion along shallow open ocean coastlines may explain the absence of mature barrier reefs in the high Hawaiian Islands. Accepted: 14 May 1998     

Holocene growth of a mid-Pacific atoll: Tarawa,Kiribati

Cores from ten holes, drilled to a maximum depth of 30 m, on Tarawa atoll in the central Pacific have been utilised in a study of the Holocene development of the atoll. Four dominant lithologies, in descending order, are cay rock, unconsolidated sediment, corals and leached limestone. Petrographic and radiometric age analyses indicate that the Holocene reef has developed on a previous (last interglacial) reef; the latter shows the effects of both vadose and phreatic freshwater diagenesis. Hydrological investigations beneath the present islands indicate the presence of freshwater lenses up to 29 m thick; the modern lenses are unrelated to freshwater diagenetic imprints preserved within the limestones. Vertical accretion rates of 5–8 m/1000 years for the Holocene reef section on Tarawa are significantly higher than rates measured for other Pacific atolls. The dated coral sequences suggest a more rapid rate of sea level rise during the early Holocene, and a relatively earlier stabilisation of sea level than has been suggested previously.     

Sporadic Disturbances in Fluctuating Coral Reef Environments: El Nino and Coral Reef Development in the Eastern Pacific

The disastrous effects of the intense 1982–83 El Niño-SouthernOscillation (ENSO) bring new insight into the long-term developmentof eastern Pacific coral reefs. The 1988–83 ENSO sea surfacewarming event caused extensive reef coral bleaching (loss ofsymbiotic zooxanthellae), resulting in up to 70–95% coralmortality on reefs in Costa Rica, Panama, Colombia and Ecuador.In the Galapagos Islands (Ecuador), most coral reefs experienced>95% coral mortality. Also, several coral species experiencedextreme reductions in population size, and local and regionalextinctions. The El Niño event spawned secondary disturbances,such as increased predation and bioerosion, that continue toimpact reef-building corals. The death of Pocillopora colonieswith their crustacean guards eliminated coral barriers now allowingthe corallivore Acanthaster planci access to formerly protectedcoral prey. Sea urchins and other organisms eroded disturbedcorals at rates that exceed carbonate production, potentiallyresulting in the elimination of existing reef buildups. In otherreefbuilding regions following extensive, catastrophic coralmortality, rapid recovery often occurs through the growth ofsurviving corals, recruitment of new corals from nearby sourcepopulations, and survival of consolidated reef surfaces. Inthe eastern Pacific, however, the return of upwelling conditionsand the survival of coral predators and bioeroders hamper coralreef recovery by reducing recruitment success and eroding coralreef substrates. Thus, coral reef growth that occurs betweendisturbance events is not conserved. Repeated El Niñodisturbances, which have occurred throughout the recent geologichistory of the eastern Pacific, prevent coral communities fromincreasing in diversity and limit the development and persistenceof significant reef features. The poor development of easternPacific coral reefs throughout Holocene and perhaps much ofPleistocene time may result from recurrent thermal disturbancesof the intensity of the 1982–83 El Niño event.     

Bathymetric zonation of modern microborers in dead coral substrates from New Caledonia—Implications for paleodepth reconstructions in Holocene corals

Microboring endoliths were studied in dead modern corals collected at various depths and in growth position on La Surprise atoll in New Caledonia. Eight taxa of microborers were regularly observed: three cyanobacteria (Hyella caespitosa, Mastigocoleus testarum, and Plectonema terebrans), two chlorophytes (Ostreobium quekettii and Phaeophila dendroides) and three fungal forms (Dodgella priscus, and two unidentified fungi). The relative abundance of each taxon was determined in the modern corals throughout the studied sampling interval (from 0 down to 40 m water depth). This allowed us to propose for the first time a bathymetric model based on microendolith assemblages colonizing in situ hard-substrates of corals. In this model, the occurrence of the cyanobacteria Hyella and Mastigocoleus indicates a bathymetric range from 0 down to a water depth of 3 and 6 m respectively. The chlorophyta O. quekettii, associated with the cyanobacterium P. terebrans and fungi, is dominant from 10 to 40 m depth, which is the lower limit of our sampling. The distribution ranges of individual taxa observed in the dead coral substrates from New Caledonia are much narrower than those reported in the literature for other substrates.The bathymetric model based on distribution of microborers in modern corals was then applied to interpret the trace assemblages of microborers observed in Holocene corals collected by drilling on two reef sites of New Caledonia : La Surprise islet, in the vicinity of the sampling sites of the modern corals, and the barrier reef of Bayes, on the NE coast of Grande Terre. The paleodepth interpretations performed at various levels along the cores allow identification of several shallowing-upward sequences during the last 7000 years of reef growth in New Caledonia. Differences in the thickness of these sequences between La Surprise and Bayes sites were interpreted as differences in the space available for accommodation between the two sites, in turn related to the differential subsidence that affected New Caledonia during Holocene times. The use of the microendolith trace assemblages in paleodepth reconstructions of Holocene reefs from New Caledonia allowed detection of relative sea level variations finer than those deduced from changes in coralgal assemblages.     

Otolith growth of Springer's demoiselle, Chrysiptera springeri (Pomacentridae, Allen & Lubbock), on a protected and non-protected coral reef

The structural complexity of coral reefs is important for their function as shelter and feeding habitats for coral reef fishes, but physical disturbance by human activities often reduce complexity of the reefs by selectively destroying fragile and more complex coral species. The damselfish Springer's demoiselle Chrysiptera springeri primarily utilize complex coral heads for shelter and are hence vulnerable to human disturbance. In order to evaluate the potential effect of habitat degradation on juvenile fish growth, coral reef cover, fish age at settling and otolith growth, juvenile Springer's demoiselle was investigated on a protected and non‐protected coral reef in Darvel Bay, Borneo. The protected reef had higher coverage of complex branching corals and exhibited a more complex 3‐dimensional structure than the non‐protected reef. Springer's demoiselle settled at the same age on non‐protected and protected reefs. The growth rates of the otoliths from Springer's demoiselle were similar during the pre‐settlement period on the two reefs (manova , P > 0.05), but from age 20 to 48 days (post‐settlement period) the otolith growth rate of juveniles on the non‐protected reef was reduced compared to those from the protected reef (manova , P = 0.017). However, the differences in the otolith size, and by inference, fish size, after 48 days were small. The small effect of habitat degradation on growth is likely related to the fact that the Springer's demoiselles collected on the non‐protected reef were associated with the few remaining complex coral heads. Increased foraging‐predation tradeoffs on the non‐protected reef may decrease food intake and growth of juvenile Springer's demoiselle, but the main effect of habitat degradation on their abundance is likely to be related to lack of suitable shelter, and consequently reduced carrying capacity, on disturbed reefs.     

Coral communities and reef growth in the southern Great Barrier Reef

 Fringing reef development is limited around 22° S along the inner Great Barrier Reef, although there is substantial development north and south of this latitude. This study examined the relationships among coral communities and the extent of reef development. Reefs were examined to determine coral composition, colony abundance, colony size and growth form between the latitudes 20°S and 23°S. Major reef framework builders (scler- actinian genus Acropora and families Faviidae and Poritidae) dominated reefs north and south of 22°S, but declined significantly at 22°S where foliose and encrusting corals (Turbinaria and Montipora spp.) were most common. Porites spp. were present at 22° S but had encrusting morphologies. Consistently high turbidity at this latitude, caused by a 10 m tidal range and strong tidal flows, resuspends silts from the shallow shelf, and appears to have precluded reef development throughout the Holocene, by limiting the abundance, stunting the growth, and shortening the life expectancies of reef framework corals. The distinctions between ‘natural’ and ‘human-induced’ degradation may be interpreted on the basis of the relationship between Holocene development and current benthic community longevity. A mismatch between substantial past reef building capacity (a broad and/or thick reef) and non-existent or limited present reef-building capacity could signify anything from a long-period, natural cycle to an unprecedented deterioration in ecosystem function caused by human influence. Accepted: 29 July 1996     

Symbiotic crabs maintain coral health by clearing sediments

Stony corals are the foundation of coral reef ecosystems and form associations with other reef species. Many of these associations may be ecologically important and play a role in maintaining the health and diversity of reef systems, rendering it critical to understand the influence of symbiotic organisms in mediating responses to perturbation. This study demonstrates the importance of an association with trapeziid crabs in reducing adverse effects of sediments deposited on corals. In a field experiment, mortality rates of two species of branching corals were significantly lowered by the presence of crabs. All outplanted corals with crabs survived whereas 45–80% of corals without crabs died within a month. For surviving corals that lacked crabs, growth was slower and tissue bleaching and sediment load were higher. Laboratory experiments revealed that corals with crabs shed substantially more of the sediments deposited on coral surfaces, but also that crabs were most effective at removing grain sizes that were most damaging to coral tissues. The mechanism underlying this symbiotic relationship has not been recognized previously, and its role in maintaining coral health is likely to become even more critical as reefs worldwide experience increasing sedimentation.     

The composition and structure of a protected coral reef in Cam Ranh Bay in the South China Sea

A nonstructural reef at Hon Nai Island in Cam Ranh Bay (southern Vietnam) was investigated. In comparison with most of the coastal continental and island coral reefs of this region, it is characterized by high species richness of reef-building corals, among them scleractinians. A total of 34 species of Acropora were found, which represent 80% of the total species composition of this scleractinian genus on the reefs of Vietnam and 25% on the reefs of the Indo-Pacific. Among 169 species of scleractinians found on the reef of Hon Nai, Favia sp. nov. was previously unknown to science. The vertical bionomic zonality of the reef corresponds to a zonal distribution of environmental factors and is similar to that on reefs of the Gulf of Siam and various areas of the Pacific and the Caribbean Basin. The thriving of the Hon Nai island reef may be connected with protective measures undertaken by the Government of Vietnam and the minimization of anthropogenic impacts due to the activities of the Sanest Co.     

Predator Crown-of-Thorns Starfish (Acanthaster planci) Outbreak,Mass Mortality of Corals,and Cascading Effects on Reef Fish and Benthic Communities

Outbreaks of the coral-killing seastar Acanthaster planci are intense disturbances that can decimate coral reefs. These events consist of the emergence of large swarms of the predatory seastar that feed on reef-building corals, often leading to widespread devastation of coral populations. While cyclic occurrences of such outbreaks are reported from many tropical reefs throughout the Indo-Pacific, their causes are hotly debated, and the spatio-temporal dynamics of the outbreaks and impacts to reef communities remain unclear. Based on observations of a recent event around the island of Moorea, French Polynesia, we show that Acanthaster outbreaks are methodic, slow-paced, and diffusive biological disturbances. Acanthaster outbreaks on insular reef systems like Moorea''s appear to originate from restricted areas confined to the ocean-exposed base of reefs. Elevated Acanthaster densities then progressively spread to adjacent and shallower locations by migrations of seastars in aggregative waves that eventually affect the entire reef system. The directional migration across reefs appears to be a search for prey as reef portions affected by dense seastar aggregations are rapidly depleted of living corals and subsequently left behind. Coral decline on impacted reefs occurs by the sequential consumption of species in the order of Acanthaster feeding preferences. Acanthaster outbreaks thus result in predictable alteration of the coral community structure. The outbreak we report here is among the most intense and devastating ever reported. Using a hierarchical, multi-scale approach, we also show how sessile benthic communities and resident coral-feeding fish assemblages were subsequently affected by the decline of corals. By elucidating the processes involved in an Acanthaster outbreak, our study contributes to comprehending this widespread disturbance and should thus benefit targeted management actions for coral reef ecosystems.     

Establishment and development of patch reefs in the intracratonic Ordovician sequence near Chicoutimi, Quebec

Patch reefs occur near the top of the transgressive sequence of Ordovician Trenton Group limestones in the Chicoutimi area of Quebec, eastern Canada. Despite their small sue, these reefs comprise diverse assemblages dominated by bryozoans, corals, stromatoporoids and receptaculitid algae. Pelmatozoans and gastropods are also conspicuous. The reefs were initiated and grew in a fully marine, open shelf setting. Available substrates varied from loose skeletal lenses to soft, firm or hardened bioturbated wackestones, and the earliest stages of reef growth reflect this heterogeneity. Loose or less firm substrates were colonised by bryozoans and pelmatozoans and/or by receptaculitids, which, together with accessory organisms, stabilised the sediments and provided the basis for further reef development. The resultant firmer, slightly elevated substrates provided sites for attachment of stromatoporoids and colonial corals which spread over earlier reef organisms and sediments and dominated the later stages of reef growth. On hardened areas of sediment, stromatoporoids and corals colonised the surface directly and the early stabilising stage of reef growth is absent. The compositions and developmental stages of these Trenton Group reefs are comparable with those seen in broadly contemporaneous and often larger reefs elsewhere, and are among the earliest in which corals played an important role.     

Geomorphology and Late Quaternary development of Middleton and Elizabeth Reefs

Middleton and Elizabeth Reefs are two mid-latitude, annular reefs within the Lord Howe linear chain of volcanic islands and seamounts in the southwestern Pacific Ocean. Drilling, vibrocoring, seismic profiling, and dating indicate that each has a rim of Holocene reef framework, enclosing a lagoon partly filled by prograding sand sheets composed of fragments of coral, coralline algae, foraminifers, and other skeletal debris. The reefs lie close to the latitudinal limits for coral growth and the reef framework is very porous, dominated by branching rather than massive corals. Coralline algae are the principal binding agent in the upper reef framework. Holocene reef growth began on a foundation of Pleistocene reefal limestone encountered at a depth of 8 m in cores on the windward side of Middleton Reef. Holocene corals became established on this foundation around 6,700 radiocarbon yr B.P., implying little if any lag after inundation of the platform by the post-glacial sea-level rise. Windward reef growth tracked sea-level rise (keep-up mode), and a prominent reef crest was established on both reefs by 5,000 yr B.P. Leeward margins appear to have been characterized by catch-up growth. Development of cays is limited, and has been restricted by the paucity of coarse coralline debris or cemented conglomerate on which islands could become established. The morphology and development of Middleton and Elizabeth Reefs has been similar to that of tropical atolls, although the rate of subsidence appears to have been relatively slow reflecting their position on the margin of the foundered continental crust of the Lord Howe Rise.