Metabolic plasticity of the corals Porites lutea and Diploastrea heliopora exposed to large amplitude internal waves

The metabolic plasticity of the two mounding coral species Porites lutea (Milne-Edwards and Haime, 1860) and Diploastrea heliopora (Lamarck, 1816) was investigated in the Similan Islands (Thailand), an offshore Andaman Sea island group subjected to large amplitude internal waves (LAIW). Nutrient concentrations were highly correlated with LAIW intensity and contributed to 3- and 10-fold higher symbiont densities in P. lutea and D. heliopora, respectively, along with elevated pigment concentrations, protein content, host tissue, and symbiont biomass. The comparison of LAIW-exposed and LAIW-sheltered island faces, and LAIW-intense and LAIW-weak years suggests a species-specific metabolic plasticity to LAIW, where D. heliopora benefits more from increased nutrient and organic matter availability than P. lutea. The ubiquitous LAIW in Southeast Asia and beyond may provide so far unexplored clues to coral acclimatization to disturbances on various scales, and hence, a potential key to coral resilience to climate change.     

Differential Impact of Monsoon and Large Amplitude Internal Waves on Coral Reef Development in the Andaman Sea

The Andaman Sea and other macrotidal semi-enclosed tropical seas feature large amplitude internal waves (LAIW). Although LAIW induce strong fluctuations i.e. of temperature, pH, and nutrients, their influence on reef development is so far unknown. A better-known source of disturbance is the monsoon affecting corals due to turbulent mixing and sedimentation. Because in the Andaman Sea both, LAIW and monsoon, act from the same westerly direction their relative contribution to reef development is difficult to discern. Here, we explore the framework development in a number of offshore island locations subjected to differential LAIW- and SW-monsoon impact to address this open question. Cumulative negative temperature anomalies – a proxy for LAIW impact – explained a higher percentage of the variability in coral reef framework height, than sedimentation rates which resulted mainly from the monsoon. Temperature anomalies and sediment grain size provided the best correlation with framework height suggesting that so far neglected subsurface processes (LAIW) play a significant role in shaping coral reefs.     

Benthic Reef Primary Production in Response to Large Amplitude Internal Waves at the Similan Islands (Andaman Sea,Thailand)

Coral reefs are facing rapidly changing environments, but implications for reef ecosystem functioning and important services, such as productivity, are difficult to predict. Comparative investigations on coral reefs that are naturally exposed to differing environmental settings can provide essential information in this context. One prevalent phenomenon regularly introducing alterations in water chemistry into coral reefs are internal waves. This study therefore investigates the effect of large amplitude internal waves (LAIW) on primary productivity in coral reefs at the Similan Islands (Andaman Sea, Thailand). The LAIW-exposed west sides of the islands are subjected to sudden drops in water temperature accompanied by enhanced inorganic nutrient concentrations compared to the sheltered east. At the central island, Ko Miang, east and west reefs are only few hundred meters apart, but feature pronounced differences. On the west lower live coral cover (-38 %) coincides with higher turf algae cover (+64 %) and growth (+54 %) compared to the east side. Turf algae and the reef sand-associated microphytobenthos displayed similar chlorophyll a contents on both island sides, but under LAIW exposure, turf algae exhibited higher net photosynthesis (+23 %), whereas the microphytobenthos displayed reduced net and gross photosynthesis (-19 % and -26 %, respectively) accompanied by lower respiration (-42 %). In contrast, the predominant coral Porites lutea showed higher chlorophyll a tissues contents (+42 %) on the LAIW-exposed west in response to lower light availability and higher inorganic nutrient concentrations, but net photosynthesis was comparable for both sides. Turf algae were the major primary producers on the west side, whereas microphytobenthos dominated on the east. The overall primary production rate (comprising all main benthic primary producers) was similar on both island sides, which indicates high primary production variability under different environmental conditions.     

Large-amplitude internal waves benefit corals during thermal stress

Tropical scleractinian corals are particularly vulnerable to global warming as elevated sea surface temperatures (SSTs) disrupt the delicate balance between the coral host and their algal endosymbionts, leading to symbiont expulsion, mass bleaching and mortality. While satellite sensing of SST has proved a reliable predictor of coral bleaching at the regional scale, there are large deviations in bleaching severity and mortality on the local scale that are poorly understood. Here, we show that internal waves play a major role in explaining local coral bleaching and mortality patterns in the Andaman Sea. Despite a severe region-wide SST anomaly in May 2010, frequent upslope intrusions of cold sub-pycnocline waters due to breaking large-amplitude internal waves (LAIW) mitigated coral bleaching and mortality in shallow waters. In LAIW-sheltered waters, by contrast, bleaching-susceptible species suffered severe bleaching and total mortality. These findings suggest that LAIW benefit coral reefs during thermal stress and provide local refugia for bleaching-susceptible corals. LAIW are ubiquitous in tropical stratified waters and their swash zones may thus be important conservation areas for the maintenance of coral diversity in a warming climate. Taking LAIW into account can significantly improve coral bleaching predictions and provide a valuable tool for coral reef conservation and management.     

Sponge bioerosion accelerated by ocean acidification across species and latitudes?

In many marine biogeographic realms, bioeroding sponges dominate the internal bioerosion of calcareous substrates such as mollusc beds and coral reef framework. They biochemically dissolve part of the carbonate and liberate so-called sponge chips, a process that is expected to be facilitated and accelerated in a more acidic environment inherent to the present global change. The bioerosion capacity of the demosponge Cliona celata Grant, 1826 in subfossil oyster shells was assessed via alkalinity anomaly technique based on 4 days of experimental exposure to three different levels of carbon dioxide partial pressure (pCO₂) at ambient temperature in the cold-temperate waters of Helgoland Island, North Sea. The rate of chemical bioerosion at present-day pCO₂ was quantified with 0.08–0.1 kg m^?2 year^?1. Chemical bioerosion was positively correlated with increasing pCO₂, with rates more than doubling at carbon dioxide levels predicted for the end of the twenty-first century, clearly confirming that C. celata bioerosion can be expected to be enhanced with progressing ocean acidification (OA). Together with previously published experimental evidence, the present results suggest that OA accelerates sponge bioerosion (1) across latitudes and biogeographic areas, (2) independent of sponge growth form, and (3) for species with or without photosymbionts alike. A general increase in sponge bioerosion with advancing OA can be expected to have a significant impact on global carbonate (re)cycling and may result in widespread negative effects, e.g. on the stability of wild and farmed shellfish populations, as well as calcareous framework builders in tropical and cold-water coral reef ecosystems.     

Biological degradation of coral framework in a turbid lagoon environment,Discovery Bay,north Jamaica

The potentially negative effects of increased sedimentation on corals are well documented, whereas, the impacts upon early diagenetic processes, such as bioerosion, remain poorly understood. This study examined macroboring through image analysis of coral slabs from two high sedimentation and turbid reefs, Columbus Park and Red Buoy, within Discovery Bay, north Jamaica. Infestation of coral framework by macroborers was significant at both Columbus Park and Red Buoy for all depth zones sampled: 0–8 m (6.5 versus 8.3%), 8–16 m (11.4 versus 10.7%), and 16–25 m (6.2 versus 18.5%), with only the deepest zone significantly different (P<0.001). Bioeroding communities exhibit a shift from mainly sponge-dominated (>90%) assemblages in clear-water settings towards a greater relative importance of worms (up to 17.2%) and bivalves (up to 40.5%) with increasing sedimentation. The high infestation levels of the bivalve Lithophaga spp. offset the reduced sponge bioerosion. As a result, macroboring infestation levels are comparable to those reported from adjacent clear-water reef sites. This study indicates that macroboring of coral framework continues under environmental conditions previously inferred to be detrimental to coral growth and survival.     

Large-amplitude internal waves sustain coral health during thermal stress

Ocean warming is a major threat for coral reefs causing widespread coral bleaching and mortality. Potential refugia are thus crucial for coral survival. Exposure to large-amplitude internal waves (LAIW) mitigated heat stress and ensured coral survival and recovery during and after an extreme heat anomaly. The physiological status of two common corals, Porites lutea and Pocillopora meandrina, was monitored in host and symbiont traits, in response to LAIW-exposure throughout the unprecedented 2010 heat anomaly in the Andaman Sea. LAIW-exposed corals of both species survived and recovered, while LAIW-sheltered corals suffered partial and total mortality in P. lutea and P. meandrina, respectively. LAIW are ubiquitous in the tropics and potentially generate coral refuge areas. As thermal stress to corals is expected to increase in a warming ocean, the mechanisms linking coral bleaching to ocean dynamics will be crucial to predict coral survival on a warming planet.     

Macrobioerosion in Porites corals in subtropical northern South China Sea: a limiting factor for high-latitude reef framework development

Bioerosion is an important limiting factor in carbonate accretion and reef framework development; however, few studies have quantified the direct impact of macroborers on high-latitude coral communities, which are viewed as potential refuge during a period of global warming. In this study, internal macrobioerosion of Porites corals was examined at Daya Bay, subtropical northern South China Sea. The principal borers were the bivalve Lithophaga spp. and the sponges Cliona spp. and Cliothosa spp. (≥80 %), while sipunculid and polychaete worms and barnacles accounted for small amounts of bioerosion (≤20 %). Porites corals were heavily bioeroded in areas impacted by aquacultural and urban activities (10.34–27.55 %) compared with corals in relatively unpolluted areas (2.18–6.76 %). High levels of bioerosion, especially boring bivalve infestation, significantly weaken the corals and increase their susceptibility to dislodgement and fragmentation in typhoons, limiting accumulation of limestone framework. This study implies that carbonate accretion and reef development for high-latitude coral communities may be limited in future high-CO₂ and eutrophication-stressed environments.     

Decadal‐scale rates of reef erosion following El Niño‐related mass coral mortality

As the frequency and intensity of coral mortality events increase under climate change, understanding how declines in coral cover may affect the bioerosion of reef frameworks is of increasing importance. Here, we explore decadal‐scale rates of bioerosion of the framework building coral Orbicella annularis by grazing parrotfish following the 1997/1998 El Niño‐related mass mortality event at Long Cay, Belize. Using high‐precision U‐Th dating and CT scan analysis, we quantified in situ rates of external bioerosion over a 13‐year period (1998–2011). Based upon the error‐weighted average U‐Th age of dead O. annularis skeletons, we estimate the average external bioerosion between 1998 and 2011 as 0.92 ± 0.55 cm depth. Empirical observations of herbivore foraging, and a nonlinear numerical response of parrotfish to an increase in food availability, were used to create a model of external bioerosion at Long Cay. Model estimates of external bioerosion were in close agreement with U‐Th estimates (0.85 ± 0.09 cm). The model was then used to quantify how rates of external bioerosion changed across a gradient of coral mortality (i.e., from few corals experiencing mortality following coral bleaching to complete mortality). Our results indicate that external bioerosion is remarkably robust to declines in coral cover, with no significant relationship predicted between the rate of external bioerosion and the proportion of O. annularis that died in the 1998 bleaching event. The outcome was robust because the reduction in grazing intensity that follows coral mortality was compensated for by a positive numerical response of parrotfish to an increase in food availability. Our model estimates further indicate that for an O. annularis‐dominated reef to maintain a positive state of reef accretion, a necessity for sustained ecosystem function, live cover of O. annularis must not drop below a ~5–10% threshold of cover.     

Comments on coral reef regeneration, bioerosion, biogeography, and chemical ecology: future directions

This paper represents a brief review of three processes operating on coral reefs and the results of studies of a fourth process, and how those results may be applied to the benefit of humankind. The areas are reef regeneration processes; bioerosion; dispersal, recruitment and biogeography of corals; and chemical ecology and natural products chemistry of reef organisms. Possible future directions for research will also be considered in each area. Regarding reef regeneration processes, coral reefs are degrading rapidly on a global scale due to over-fishing, fishing techniques causing habitat destruction, deforestation, mass mortalities of key reef species, nutrient enrichment and sedimentation. Seeding of reefs with the larvae of corals and other key reef organisms, such as echinoids, may help to promote and enhance reef regeneration in the future. Such techniques will be made possible by studies of the embryology, larval ecology, dispersal and recruitment processes, and related local physical oceanographic processes. Regarding bioerosion, both internal and external bioerosion are affected by grazers and predators. Bioerosion is also affected by nutrient enrichment, as shown through correlative studies (Great Barrier Reef) and studies of opportunity (Kaneohe Bay). Ongoing experiments such as ENCORE will help to answer questions about the role of dissolved nutrients in enhancing internal bioerosion. Questions still remain, however, regarding the role of particulates in promoting internal bioerosion and the resultant weakening of and negative growth in the reef framework. Regarding dispersal, recruitment and the biogeography of corals, it is now known that most species of coral reproduce via broadcasting, although there appear to be proportionally more brooders in the Caribbean than in the western Pacific. Differential extinctions in the western Pacific vs. the western Atlantic have contributed to the biogeographic distribution of corals we observe today and the concentric isoclines of species diversity in numerous reef organisms in the western Pacific. The role of reproductive mode in contributing to these patterns is, however, still not understood. Investigations into the roles of different larval longevities and reproductive modes may help us answer questions regarding their differential distribution and the potential effects of major perturbations such as global warming on future distributions. With respect to the chemical ecology of alcyonacean octocorals (soft corals), four functions have been determined thus far for secondary metabolites in this group, anti-predation, anti-competition (allelopathy), anti-fouling, and enhancement of reproductive success. Investigations of alcyonacean reproduction has revealed that it may be necessary for several secondary metabolites to be present simultaneously before a function may be realized or fully manifested. This raises questions regarding the manner in which novel compounds are tested by medical laboratories for bioactivity using a single compound. Simultaneously testing of multiple compounds derived from a single organism may be necessary in the future to reveal potential valuable synergistic bioactivity. Also, some novel secondary metabolites may have other valuable commercial applications, as is the case with the UV-absorbing compounds of corals and other reef organisms found on the Great Barrier Reef. In order to avoid overlooking medically or commercially valuable functions of these compounds, broader testing may be necessary.     

Ocean Acidification Accelerates Reef Bioerosion

In the recent discussion how biotic systems may react to ocean acidification caused by the rapid rise in carbon dioxide partial pressure (pCO₂) in the marine realm, substantial research is devoted to calcifiers such as stony corals. The antagonistic process – biologically induced carbonate dissolution via bioerosion – has largely been neglected. Unlike skeletal growth, we expect bioerosion by chemical means to be facilitated in a high-CO₂ world. This study focuses on one of the most detrimental bioeroders, the sponge Cliona orientalis, which attacks and kills live corals on Australia’s Great Barrier Reef. Experimental exposure to lowered and elevated levels of pCO₂ confirms a significant enforcement of the sponges’ bioerosion capacity with increasing pCO₂ under more acidic conditions. Considering the substantial contribution of sponges to carbonate bioerosion, this finding implies that tropical reef ecosystems are facing the combined effects of weakened coral calcification and accelerated bioerosion, resulting in critical pressure on the dynamic balance between biogenic carbonate build-up and degradation.     

Effects of grazing and damselfish territoriality on internal bioerosion of dead corals : indirect effects

An experiment was performed on Britomart Reef, Great Barrier Reef (central region), to determine the relationship between fish grazing, damselfish territoriality, and internal bioerosion of dead coral substratum. The damselfish Hemiglyphidodon plagiometopon Bleeker (Pomacentridae) was used for the study because it actively excludes herbivorous fish, particularly scarids and acanthurids, from its territories, creating undergrazed patches in the environment. Its territories simulated conditions of naturally reduced grazing. Freshly killed pieces of the plating coral Pachyseris speciosa Dana were placed under four experimental conditions: (1) within cages, excluding grazing fish; (2) within damselfish territories; (3) beneath shade tops to control for light; and (4) outside damselfish territories, fully exposed to grazers. Internal bioeroders were identified by pattern of substratum excavation and characterization of borings, and were quantified by digitizing x-ray radiographs of the substratum. Three major categories of borers were identified: Cliothosa hancocki Topsent, “other sponges” (of the Cliona viridis Schmidt species complex), and “worms” (including polychaetes and sipunculids). Variations in grazing pressure were found to significantly alter the taxonomic composition of the bioeroder community. Bioerosion by C. hancocki, a boring sponge with large exposed papillae, was found to increase significantly when grazing was reduced within damselfish territories. By contrast, other boring sponges of the C. viridis complex decreased in abundance; they were not affected by higher sedimentation in cages. The response of bioerosion by “worms” was less clear but increased slightly within damselfish territories. This was due primarily to a shift in taxonomic composition and dominance from polychaetes to sipunculids (particularly Cleosiphon aspergillum Quatrefages). The effects of grazing on the internal bioeroder community were often altered or obscured in the caged treatments; this was most likely due to caging artifacts such as increased sedimentation and decreased light. In general, bioerosion rates of the substratum P. speciosa were low in comparison to rates established or estimated for corals with less dense skeletons. Total internal bioerosion rates did not vary significantly with changes in grazing pressure. This study implies that, overall, reduced grazing pressure will lead to production of fine sediments derived from internal bioeroders. Under high grazing pressures, the addition of external bioerosion effected directly by grazers would also produce coarse sediment, resulting in an increase in total bioerosion rates (internal and external) and an increased contribution of both coarse and fine sediments to the reef environment.     

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.     

Bitten down to size: Fish predation determines growth form of the Caribbean coral reef sponge Mycale laevis

Interactions between organisms add complexity to ecosystem function, particularly on coral reefs. The Caribbean orange icing sponge Mycale laevis is semi-cryptic, often growing under coral colonies or between coral branches. This association is reportedly a mutualism, with the sponge deterring boring sponges from invading the coral skeleton and the coral providing an expanding surface for sponge growth. But is there an alternative explanation for the proximity of sponge and coral? We examined the importance of fish predation on the growth of the sponge. While the semi-cryptic growth form of M. laevis predominates on reefs off the Florida Keys and the Bahamas Islands, M. laevis grows with a non-cryptic, erect morphology off Bocas del Toro, Panama. Surveys revealed that sponge-eating fishes were rare or absent at Bocas del Toro compared to sites in the Florida Keys. Because past studies were inconsistent about the palatability of M. laevis to fish predators, we conducted feeding experiments with sponges from all three sites. Crude organic extracts of M. laevis from all three sites were palatable to generalist fish predators in aquarium assays, and field feeding assays and caging experiments conducted in the Florida Keys confirmed that spongivorous fishes readily ate exposed fragments of M. laevis. Our results suggest that M. laevis is restricted to its semi-cryptic growth form by spongivorous predators, with corals providing a physical refuge from predation. This alternative explanation supports the broader hypothesis that Caribbean reef sponges can be categorized on the basis of chemical defense into defended, palatable, and preferred species, the last of which are restricted to refugia.     

Sponge borehole size as a relative measure of bioerosion and paleoproductivity

Bioerosion intensity has been proposed as a measure of paleoproductivity in fossil reefs, but it is difficult to measure directly because fossil corals are often incomplete and because it is difticult to infer the length of time a given coral was exposed to bioeroding organisms. Both nutrient availability and taphonomic factors can affect bioerosion intensity as measured in dead corals. Here, we examine these two effects separately using data from previous studies on bioerosion in modern and fossil corals. Size of individual sponge borings accurately reflects total bioerosion in modern massive and branching corals on the Great Barrier Reef. Total bioerosion in both massive and branching corals decreases outward across the continental shelf, paralleling trends in nutrient availability. Size of individual Cliothosa hancocki borings decreases across the shelf in branching Acropora but not in massive Porites. Fossil sponge borings Entobia convoluta and Uniglobites glomerata in massive corals from Oligocene and Miocene reefs in Puerto Rico are smallest in Oligocene shelf-edge reefs, intermediate in Oligocene patch reefs, and largest in Miocene patch reefs. Both facies-related influence, represented by Oligocene shelf-edge reefs vs. Oligocene patch reefs, and nutrient-related influence, represented by Oligocene vs. Miocene patch reefs, were reflected in the size of sponge boreholes. Size of sponge borings also varies among species of host corals, apparently in relation to skeletal architecture. Borehole size is inversely correlated with skeletal density as measured by the relative proportion of skeleton and pore space in transverse thin section. There is a weak positive correlation between borehole size and corallite diameter. These findings contradict reported positive correlations between total bioerosion and bulk density in modern corals. Borehole size appears accurately to reflect intensity of total internal bioerosion in fossil corals. Facies-controlled taphonomic overprints and influence of skeletal differences between coral species limit the use of sponge borehole size to a rough indicator of paleoproductivity in fossil coral reef environments.     

Boring sponges,an increasing threat for coral reefs affected by bleaching events

Coral bleaching is a stress response of corals induced by a variety of factors, but these events have become more frequent and intense in response to recent climate‐change‐related temperature anomalies. We tested the hypothesis that coral reefs affected by bleaching events are currently heavily infested by boring sponges, which are playing a significant role in the destruction of their physical structure. Seventeen reefs that cover the entire distributional range of corals along the Mexican Pacific coast were studied between 2005/2006, and later between 2009/2010. Most of these coral reefs were previously impacted by bleaching events, which resulted in coral mortalities. Sponge abundance and species richness was used as an indicator of bioerosion, and coral cover was used to describe the present condition of coral reefs. Coral reefs are currently highly invaded (46% of the samples examined) by a very high diversity of boring sponges (20 species); being the coral reef framework the substrate most invaded (56%) followed by the rubbles (45%), and the living colonies (36%). The results also indicated that boring sponges are promoting the dislodgment of live colonies and large fragments from the framework. In summary, the eastern coral reefs affected by bleaching phenomena, mainly provoked by El Niño, present a high diversity and abundance of boring sponges, which are weakening the union of the colony with the reef framework and promoting their dislodgment. These phenomena will probably become even more intense and severe, as temperatures are projected to continue to rise under the scenarios for future climate change, which could place many eastern coral reefs beyond their survival threshold.     

‘Ten Years After’—a long-term settlement and bioerosion experiment in an Arctic rhodolith bed (Mosselbukta,Svalbard)

Rhodolith beds and bioherms formed by ecosystem engineering crustose coralline algae support the northernmost centres of carbonate production, referred to as polar cold-water carbonate factories. Yet, little is known about biodiversity and recruitment of these hard-bottom communities or the bioeroders degrading them, and there is a demand for carbonate budgets to include respective rates of polar carbonate build-up and bioerosion. To address these issues, a 10-year settlement and bioerosion experiment was carried out at the Arctic Svalbard archipelago in and downslope of a rhodolith bed. The calcifiers recorded on experimental settlement tiles (56 taxa) were dominated by bryozoans, serpulids and foraminiferans. The majority of the bioerosion traces (30 ichnotaxa) were microborings, followed by attachment etchings and grazing traces. Biodiversity metrics show that calcifier diversity and bioerosion ichnodiversity are both elevated in the rhodolith bed, if compared to adjacent aphotic waters, but these differences are statistically insignificant. Accordingly, there were only low to moderate dissimilarities in the calcifier community structure and bioerosion trace assemblages between the two depth stations (46 and 127 m), substrate orientations (up- and down-facing) and substrate types (PVC and limestone), in that order of relevance. In contrast, surface coverage as well as the carbonate accretion and bioerosion rates were all significantly elevated in the rhodolith bed, reflecting higher abundance or size of calcifiers and bioerosion traces. All three measures were highest for up-facing substrates at 46 m, with a mean coverage of 78.2% (on PVC substrates), a mean accretion rate of 24.6 g m^?2  year^?1 (PVC), and a mean bioerosion rate of ?35.1 g m^?2 year^?1 (limestone). Differences in these metrics depend on the same order of factors than the community structure. Considering all limestone substrates of the two platforms, carbonate accretion and bioerosion were nearly in balance at a net rate of ?2.5 g m^?2 year^?1. A latitudinal comparison with previous settlement studies in the North Atlantic suggests that despite the harsh polar environment there is neither a depletion in the diversity of hard-bottom calcifier communities nor in the ichnodiversity of grazing traces, attachment etchings and microborings formed by organotrophs. In contrast, microborings produced by phototrophs are strongly depleted because of limitations in the availability of light (condensed photic zonation, polar night, shading by sea ice). Also, macroborings were almost absent, surprisingly. With respect to carbonate production, the Svalbard carbonate factory marks the low end of a latitudinal gradient while bioerosion rates are similar or even higher than at comparable depth or photic regime at lower latitudes, although this might not apply to shallow euphotic waters (not covered in our experiment), given the observed depletion in bioeroding microphytes and macroborers. While echinoid grazing is particularly relevant for the bioerosion in the rhodolith bed, respective rates are far lower than those reported from tropical shallow-water coral reefs. The slow pace of carbonate production but relatively high rates of bioerosion (both promoted by low carbonate supersaturation states in Arctic waters), in concert with high retention of skeletal carbonates on the seafloor and no calcite cements forming in open pore space created by microborers, suggest a low fossilisation potential for polar carbonates, such as those formed in the Mosselbukta rhodolith beds.     

Bioerosion caused by the sea urchin Diadema Mexicanum (Echinodermata: Echinoidea) at Bahías de Huatulco, Western Mexico

Mexican Pacific sea urchin studies have been focused mainly on species distribution, ecology and fisheries. Reef degradation by sea urchin bioerosion has not been studied previously en these reefs. We investigate the importance of Diadema mexicanum as a bioerosive agent of coral carbonate at Bahias de Huatulco, and the relative magnitude of coral accretion and bioerosion. At each of five localities in Bahias de Huatulco, sea urchin density, feeding and mechanical (spine) erosion was determined for three size class intervals. In general, D. mexicanum do not exert any significant role on coral reef community structure (live coral, dead coral or algal coverage) at the Huatulco area, probably because they are generally small (2.9-4 cm test size) and few in number (1.0-6.8 ind.m-2). Mean bioerosion rates are consistent with those measured for other diadematoids, as well as other urchin species in various eastern Pacific localities. However, the degree of bioerosive impact depends on species, test size, and population density of urchins. Coral carbonate removal by D. mexicanum erosion varies from 0.17 to 3.28 kgCaCO3m(-2)yr(-1). This represents a carbonate loss of < 5% of the annual coral carbonate production at Jicaral Chachacual, San Agustín and Isla Cacaluta, but 16 and 27% at Isla Montosa and La Entrega. On balance, coral accretion exceeds sea urchin erosion at all sites examined at Huatulco. At Bahias de Huatulco coral reef communities are actively growing, though in the coming years, it might be necessary to investigate the local effects of the interaction among erosion, and environmental and human induced perturbations.     

Growth models of Bajocian coral-microbialite reefs of Chargey-lès-Port (eastern France): palaeoenvironmental interpretations

Very large amount of microbialites, up to 70% of the reef volume takes part in the edification of Lower Bajocian coral reefs in the Chargey-lès-Port quarry (Haute-Saône, France). Such high amounts of microbialites were unknown within bioconstructions of Middle Jurassic age. Along the 16 m-thick section, seven successive biohermal or biostromal units developed on a shallow platform. Bioconstructions display a first coral growth phase with either constratal or superstratal growth fabrics. Coral fauna is relatively poorly diversified and is dominated by massive forms (Isastrea, Thamnasteria, and Periseris) or branched phaceloid (Cladophyllia) and ramose (Dendraraea) colonies. Corals can be heavily encrusted by microbialites of diverse forms and fabrics (leiolitic, thrombolitic, and stromatolitic). According to the coral growth fabrics, microbialite crusts developed on top of or at the underside of coral colonies, forming a coral-microbialite elementary unit. Microbialites show a multiphase development: (i) directly at the coral surface, a first and mm-scale microbialite layer locally developed; (ii) a second, cm-scale microbialite layer (up to 8 cm thick) covered the entire coral reef framework and assumed the main building role; and (iii) a third, mm- to cm-scale, laminated microbialite layer may also be observed onlapping previous reef structures, before having been progressively buried under sediments. Contemporaneously to the coral growth phase, the first microbialite layer developed on dead portions of coral colonies. The transition between coral growth and microbialite development (i.e., second layer of microbialites) is interpreted as a result of a coral reef crisis, probably reflecting more nutrient-rich conditions. The passage to a stromatolitic (third) layer suggests a control of the accumulation rate. Composition and architecture of coral-microbialite reef units of Chargey-lès-Port highlight the relations between high-frequency fluctuating environmental factors (mainly accumulation rate and trophic conditions) and reef development.