A Novel μCT Analysis Reveals Different Responses of Bioerosion and Secondary Accretion to Environmental Variability

Corals build reefs through accretion of calcium carbonate (CaCO₃) skeletons, but net reef growth also depends on bioerosion by grazers and borers and on secondary calcification by crustose coralline algae and other calcifying invertebrates. However, traditional field methods for quantifying secondary accretion and bioerosion confound both processes, do not measure them on the same time-scale, or are restricted to 2D methods. In a prior study, we compared multiple environmental drivers of net erosion using pre- and post-deployment micro-computed tomography scans (μCT; calculated as the % change in volume of experimental CaCO₃ blocks) and found a shift from net accretion to net erosion with increasing ocean acidity. Here, we present a novel μCT method and detail a procedure that aligns and digitally subtracts pre- and post-deployment μCT scans and measures the simultaneous response of secondary accretion and bioerosion on blocks exposed to the same environmental variation over the same time-scale. We tested our method on a dataset from a prior study and show that it can be used to uncover information previously unattainable using traditional methods. We demonstrated that secondary accretion and bioerosion are driven by different environmental parameters, bioerosion is more sensitive to ocean acidity than secondary accretion, and net erosion is driven more by changes in bioerosion than secondary accretion.     

Influence of Diadema antillarum populations (Echinodermata: Diadematidae) on algal community structure in Jardines de la Reina, Cuba

The 1983-1984 mass mortality of Diadema antillarum produced severe damages on Caribbean reefs contributing to substantial changes in community structure that still persist. Despite the importance of Diadema grazing in structuring coral reefs, available information on current abundances and algal-urchin interactions in Cuba is scarce. We analyzed spatial variations in Diadema abundance and its influence on algal community structure in 22 reef sites in Jardines de la Reina, in June/2004 and April/2005. Urchins were counted in five 30 x 2m transects per site, and algal coverage was estimated in randomly located 0.25m side quadrats (15 per site). Abundances of Diadema were higher at reef crests (0.013-1.553 ind/m2), while reef slope populations showed values up to three orders of magnitude lower and were overgrown by macroalgae (up to 87%, local values). Algal community structure at reef slopes were dominated by macroalgae, especially Dictyota, Lobophora and Halimeda while the most abundant macroalgae at reef crests were Halimeda and Amphiroa. Urchin densities were negatively and positively correlated with mean coverage of macroalgae and crustose coralline algae, respectively, when analyzing data pooled across all sites, but not with data from separate habitats (specially reef crest), suggesting, along with historical fish biomass, that shallow reef community structure is being shaped by the synergistic action of other factors (e.g. fish grazing) rather than the influence of Diadema alone. However, we observed clear signs of Diadema grazing at reef crests and decreased macroalgal cover according to 2001 data, what suggest that grazing intensity at this habitat increased at the same time that Diadema recruitment began to be noticeable. Furthermore, the excessive abundance of macroalgae at reef slopes and the scarcity of crustose coralline algae seems to be due by the almost complete absence of D. antillarum at mid depth reefs, where local densities of this urchin were predominantly low.     

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.     

Holocene initiation and development of New Caledonian fringing reefs,SW Pacific

The analysis of 8 selected cores through fringing reefs in New Caledonia reveals that accretion in the Holocene has been less than 6 m. The cores exhibit three main facies: branching coral (Acropora, dominantly), massive coral heads (Porites, mainly) and coral sand/ rubble, principally made up of acroporid fragments. Subordinate facies are composed of coralline algae and alcyonarian spiculite. The initiation of growth varies according to location. The southern reefs (i.e. early settled reefs) generally began to grow first, prior to 5000 y BP. The northern structures (i.e. more recently settled reefs) are younger, occurring after 4200 y BP. This retardation could be ascribed to differences in local physical conditions (nature of substrate, wave energy). Vertical accretion rates were generally higher in areas of lower energy (3.25–6.4 mm·y^-1) versus those exposed to higher energy conditions (1.4–3.1 mm·y^-1). Vertical development through time was accompanied by changes in composition of biological assemblages which reflect changes in hydrodynamics. The basal Acropora-dominated facies was replaced upwards by a Porites-dominated framework. The New Caledonian fringing reefs reached the sea surface generally between 5000 and 2500 y BP after the stabilization of sea level. Hence all of these reefs can be classified as catch-up reefs.     

Coralline algal maerl frameworks-Islands within the phaeophytic kelp belt

Summary In the subtropical belt highly productive ecosystems are formed by coral reefs in oligotrophic seas. Towards more eutrophic conditions, coral reefs diminish and are subsequently replaced by highly productive kelp forests. In high latitudes framework constructing carbonate production is enhanced by the growth of branching coralline algae which predominantly generate maerl-type deposits. On a global view, these coralline algal ecosystems show an island-like distribution pattern within the phaeophytic kelp belt. Compared to kelp ecosystems, coralline-algaldominated ecosystems have low rates of productivity. Therefore, it is reasonable to seek the pronounced competitive value of the extremely slow-growing corallines. Due to their low annual growth increment, the coralline algae studied are very endangered by abiotic physical disturbances and by overgrowth of rapidly growing filamentous algae or sessile invertebrates. To overcome fouling pressure and storm-triggered physical disturbances, coralline algae thrive well in wave-sheltered headlands or skerry areas and generate characteristic ‘denuded areas’ by intense herbivory. This general distributional pattern is also true for high-boreal to subarctic coralline algal bioherms in northern Norway. Such a complex biological feedback maintains a high potential of self-regulation or self-organization in the algal reef bioherms. The different proponents involved in feedback processes include bacterial colonization, diatom microfouling and selective induction of larval metamorphosis. The negative impact of diatom microfouling and the important role of herbivores are relevant activities in the feedback system on a microscopic scale. Macroscopically, intense herbivory on coralline algae create denuded conditions, which are a widespread phenomenon in coralline algal ecosystems.     

Holocene reef evolution in a macrotidal setting: Buccaneer Archipelago,Kimberley Bioregion,Northwest Australia

This study uses information derived from cores to describe the Holocene accretion history of coral reefs in the macrotidal (up to 11 m tidal range) Buccaneer Archipelago of the southern Kimberley coast, Western Australia. The internal architecture of all cored reefs is broadly similar, constituting well-preserved detrital coral fragments, predominantly branching Acropora, in a poorly sorted sandy mud matrix. However, once the reefs reach sea level, they diverge into two types: low intertidal reefs that maintain their detrital character and develop relatively narrow, horizontal or gently sloping reef flats at approximately mean low water spring, and high intertidal reefs that develop broad coralline algal-dominated reef flats at elevations between mean low water neap and mean high water neap. The high intertidal reefs develop where strong, ebb-dominated, tidal asymmetry retains seawater over the low tide and allows continued accretion. Both reef types are ultimately constrained by sea level but differ in elevation by 3–4 m.     

The relationship between habitat structure and fish faunas on New Zealand reefs

Reef fish abundances were sampled at 11 shallow reef localities extending over 1000 km of coastline in northern New Zealand. Sampling was restricted to the 4–10-m depth stratum and included six coastal and five island localities. These were either coralline reef flats dominated by echinoids, or algal reefs with high densities of laminarian and fucoid algae. Reefs dominated by macroscopic algae supported large numbers of small fishes, mainly labrids, and few large benthic-feeding fishes. Echinoid-dominated reefs supported a different fish fauna with more large benthic-feeding species. Additional sampling of echinoiddominated reefs and algal stands in deeper water provided confirmation of these findings. A second sampling programme was carried out at a series of eight sites within a single locality covering 5 km of coastline. These spanned a moderate exposure gradient and ranged from algal dominated reefs to typical coralline reef flats with high densities of grazing invertebrates. The relationship between habitat structure and reef fish species composition and size frequency was similar to that of the large-scale sampling programme. Thirdly, observations on reef fish foraging and feeding patterns within a single reef site suggested that larger benthic-feeding reef fishes were less likely to feed within macroscopic algal stands. Experimental reductions of grazing invertebrates designed to produce brown algal stands on echinoid-dominated reef flats supported these observations. Larger individuals capable of removing echinoids and grazing gastropods did not frequent or feed in laminarian and fucoid algal stands. This pattern is discernible at several spatial scales. Our conclusion is that the type of shallow reef habitat, echinoid- as opposed to algal-dominated, will have an important rôle in determining the associated reef fish fauna.     

Two decades of carbonate budget change on shifted coral reef assemblages: are these reefs being locked into low net budget states?

The ecology of coral reefs is rapidly shifting from historical baselines. One key-question is whether under these new, less favourable ecological conditions, coral reefs will be able to sustain key geo-ecological processes such as the capacity to accumulate carbonate structure. Here, we use data from 34 Caribbean reef sites to examine how the carbonate production, net erosion and net carbonate budgets, as well as the organisms underlying these processes, have changed over the past 15 years in the absence of further severe acute disturbances. We find that despite fundamental benthic ecological changes, these ecologically shifted coral assemblages have exhibited a modest but significant increase in their net carbonate budgets over the past 15 years. However, contrary to expectations this trend was driven by a decrease in erosion pressure, largely resulting from changes in the abundance and size-frequency distribution of parrotfishes, and not by an increase in rates of coral carbonate production. Although in the short term, the carbonate budgets seem to have benefitted marginally from reduced parrotfish erosion, the absence of these key substrate grazers, particularly of larger individuals, is unlikely to be conducive to reef recovery and will thus probably lock these reefs into low budget states.     

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.     

Calcification rates of the massive coral Siderastrea siderea and crustose coralline algae along the Florida Keys (USA) outer-reef tract

Coral reefs are degrading on a global scale, and rates of reef-organism calcification are predicted to decline due to ocean warming and acidification. Systematic measurements of calcification over space and time are necessary to detect change resulting from environmental stressors. We established a network of calcification monitoring stations at four managed reefs along the outer Florida Keys Reef Tract (FKRT) from Miami to the Dry Tortugas. Eighty colonies (in two sequential sets of 40) of the reef-building coral, Siderastrea siderea, were transplanted to fixed apparatus that allowed repetitive detachment for buoyant weighing every 6 months. Algal-recruitment tiles were also deployed during each weighing interval to measure net calcification of the crustose coralline algal (CCA) community. Coral-calcification rates were an order of magnitude greater than those of CCA. Rates of coral calcification were seasonal (summer calcification was 53 % greater than winter), and corals in the Dry Tortugas calcified 48 % faster than those at the other three sites. Linear extension rates were also highest in the Dry Tortugas, whereas percent area of the coral skeletons excavated by bioeroding fauna was lowest. The spatial patterns in net coral calcification revealed here correlate well with Holocene reef thickness along the FKRT and, in part, support the “inimical waters hypothesis” proposed by Ginsburg, Hudson, and Shinn almost 50 yrs ago to explain reef development in this region. Due to the homogeneity in coral-calcification rates among the three main Keys sites, we recommend refinement of this hypothesis and suggest that water-quality variables (e.g., carbonate mineral saturation state, dissolved and particulate organic matter, light attenuation) be monitored alongside calcification in future studies. Our results demonstrate that our calcification monitoring network presents a feasible and worthwhile approach to quantifying potential impacts of ocean acidification, warming, and/or deteriorating water quality on the process of calcification.     

Extreme spatial heterogeneity in carbonate accretion potential on a Caribbean fringing reef linked to local human disturbance gradients

The capacity of coral reefs to maintain their structurally complex frameworks and to retain the potential for vertical accretion is vitally important to the persistence of their ecological functioning and the ecosystem services they sustain. However, datasets to support detailed along‐coast assessments of framework production rates and accretion potential do not presently exist. Here, we estimate, based on gross bioaccretion and bioerosion measures, the carbonate budgets and resultant estimated accretion rates (EAR) of the shallow reef zone of leeward Bonaire – between 5 and 12 m depth – at unique fine spatial resolution along this coast (115 sites). Whilst the fringing reef of Bonaire is often reported to be in a better ecological condition than most sites throughout the wider Caribbean region, our data show that the carbonate budgets of the reefs and derived EAR varied considerably across this ~58 km long fringing reef complex. Some areas, in particular the marine reserves, were indeed still dominated by structurally complex coral communities with high net carbonate production (>10 kg CaCO₃ m^?2 year^?1), high live coral cover and complex structural topography. The majority of the studied sites, however, were defined by relatively low budget states (<2 kg CaCO₃ m^?2 year^?1) or were in a state of net erosion. These data highlight the marked spatial heterogeneity that can occur in budget states, and thus in reef accretion potential, even between quite closely spaced areas of individual reef complexes. This heterogeneity is linked strongly to the degree of localized land‐based impacts along the coast, and resultant differences in the abundance of reef framework building coral species. The major impact of this variability is that those sections of reef defined by low‐accretion rates will have limited capacity to maintain their structural integrity and to keep pace with current projections of climate change induced sea‐level rise (SLR), thus posing a threat to reef functioning and biodiversity, potentially leading to trophic cascades. Since many Caribbean reefs are more severely degraded than those found around Bonaire, it is to be expected that the findings presented here are rather the rule than the exception, but the study also highlights the need for similar high spatial resolution (along‐coast) assessments of budget states and accretion rates to meaningfully explore increasing coastal risk at the country level. The findings also more generally underline the significance of reducing local anthropogenic disturbance and restoring framework building coral assemblages. Appropriately focussed local preservation efforts may aid in averting future large‐scale above reef water depth increases on Caribbean coral reefs and will limit the social and economic implications associated with the loss of reef goods and services.     

Biodeposition in a juvenile mussel bed of the east Frisian Wadden Sea (Southern North Sea)

Seasonal trends of biodeposition and animal coverage in backbarrier beds of the musselMytilus edulis in the East Frisian Wadden Sea were documented and quantified to assess the effects of physical disturbance by storms and to develop a conceptual model for the interpretation of biodeposits in the stratigraphic record. Accretion and erosion of biogenic muds (faeces and pseudofaeces) were quantified by repeated precision levelling of a 12 m² test area. Animal coverage was documented by vertical photography prior to each levelling survey. Levelling and photography were conducted from a bridge and rail system to avoid physical disturbance by trampling. Four surveys, covering a period of different weather conditions, showed variable mean accretion/erosion rates of biodeposits ranging from +0.56 mm d⁻¹ (accretion) to −1.34 mm d⁻¹ (erosion) and areal coverage of mussels ranging from 39.1% to 0%. A pronounced seasonal trend in the biodepositional pattern was observed. Mean accretion rates were high (>0.5 mm d⁻¹) during the summer season, whereas in the winter season net deposition decreased to zero or showed erosional trends. Longterm sediment budgets are thus composed of highly variable mean annual rates which, in turn, comprise highly variable seasonal and even monthly rates, depending on the local weather pattern. This forms the basis for a conceptual model, in terms of which the stratigraphic record of fossil biodeposits can be meaningfully interpreted.     

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.     

Ingestion and transformation of algal turf by Echinometra mathaei on Tiahura fringing reef (French Polynesia)

The sea urchin Echinometra mathaei is the most abundant herbivore on many tropical reefs. We studied the ingestion and digestion diel rhythms, transformation of algal turf and bioerosion attributable to this species on the Tiahura fringing reef in French Polynesia. Ingestion rates showed a circadian rhythm with most feeding taking place during the night. Absorption of food occurred throughout the day with urchins digesting food outside of the feeding period. A total of 73% of the faecal pellets consisted of CaCO(3) eroded from the reef, 20% consisted of organic matter and 7% the refractory organic matter. Of the organic matter, lipids, carbohydrates and chlorophyll were digested and absorbed and proteins were expelled in the faecal pellets. An average individual bioerosion of 0.32 g day(-1) was estimated for E. mathaei from approximately a 35-mm test diameter on the Tiahura fringing reef. We further estimated that E. mathaei release 70.5 g m(2) y(-1) of carbohydrates, 43.8 g m(2) y(-1) of lipid, 23.3 g m(2) y(-1) of protein and 2.0 g m(2) y(-1) of total chlorophyll pigments.     

Tracking paleoenvironmental changes in coralline algal-dominated carbonates of the Lower Oligocene Calcareniti di Castelgomberto formation (Monti Berici, Italy)

Lower Oligocene, shallow-water carbonates of the Calcareniti di Castelgomberto formation (Monti Berici, Italy, Southern Alps) are studied in detail with respect to fabric and component distributions in order to trace paleoecological changes along a monotonous sedimentary stacking pattern. The carbonates are dominated by coralline algal rudstones with a packstone to wackestone matrix. Non-geniculate coralline algae include six genera: Lithoporella melobesioides, Mesophyllum, Neogoniolithon, Spongites, Sporolithon, and Subterraniphyllum. The algae are found in the form of encrusting thalli, rhodoliths, and coralline debris. Non-algal components include larger, small benthic, and planktonic foraminifera associated with bryozoans, zooxanthellate corals, and echinoderms. Four carbonate facies are distinguished: (1) coralline algal facies, (2) coralline algal-coral facies, (3) coralline algal-larger foraminiferal facies, and (4) coralline algal debris facies. Marly horizons also occur in the section. The facies and coralline algal content are interpreted with respect to light intensity, hydrodynamic energy, biotic interactions, and substrate stability. Facies development along the studied section shows systematic variations, suggesting asymmetric sea-level changes with rapid regressions and gradual transgressions.     

Succession and growth rates of encrusting crustose coralline algae (Rhodophyta,Cryptonemiales) in the upper fore-reef environment off Ishigaki Island,Ryukyu Islands

Observations were made on the succession and growth rates of crustose coralline algae growing in situ on artificial substrata in a shallow fore-reef environment on Ishigaki Island, Ryukyu Islands. Succession in well-illuminated environments manifests itself as a gradual replacement of species having very thin thalli by those having larger and thicker thalli. The species Porolithon onkodes, Paragoniolithon conicum and Lithophyllum insipidum achieved dominance by competitive interactions of overgrowing margins. The thicker species recruit quickly (within the first few months), but because of their slow growth rate do not displace the pioneer species that have very thin thalli until after the latter begin to die. Regardless of seasonal temperature fluctuations, which exceed 10 °C, the coralline algal succession is the same for each season. The maximum lateral growth rates of the major species range between 2.9 and 3.9 mm/month. Vertical growth rates of Porolithon onkodes, the thickest species, are the most rapid (more than 2 mm/year at maximum) relative to those of other species. Accretion rates of entire coralline algal cover on ungrazed substrata range from 1.0 to 1.2 mm/year (not allowing any lag time for recruitment), whereas those of grazed substrata are lower. These results are consistent with species which are ecological equivalents and live in similar environments on Caribbean reefs.     

Plaeocene reef sediments from the maiella carbonate platform,Italy

Summary Upper Cretaceous and Paleocene reef limestones from the Maiella carbonate platform show how reefs evolved during a time of faunal turn-over. Biostratigraphy and facies analysis of the reef limestones reveal the details of reef growth, composition, and age. Rudists disappeared as reef builders from the Maiella platform shortly before the Cretaceous/Tertiary boundary. Small coral-algal reefs became established in the Danian to Late Thanetian. These scleractinian-red algal dominated boundstones and framestones represent two periods of reef sedimentation and the subsequent interruption of reef growth by emersion and erosion, controlled primarily by fluctuations of relative sea-level. The coral-algal reefs evolved as the taxonomic composition of reef organisms changed. The Paleocene reef sediments are preserved as large slide blocks and as boulders redeposited from the shallow-water platform onto the slope during the course of the Paleocene.     

‘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.     

Patterns and rates of erosion in dead Porites across the Great Barrier Reef (Australia) after 2 years and 4 years of exposure

Rates and agents of erosion were investigated experimentally at six sites located along a cross shelf transect from the northern Queensland coast out into the Coral Sea. Rates of internal and external erosion of coral blocks, and accretion by coralline algae were measured after 2 years and 4 years of exposure. Blocks were cut from live colonies of Porites sp., which were collected from the outer barrier reef in north Queensland. They were then washed, dried, measured, weighed and attached to grids that were firmly attached to dead coral substrate at depths of 7–10 m. Significant differences in all three parameters were found within and among sites, and rates increased with increasing duration of exposure. Inshore sites were characterized by low rates of external erosion compared to offshore sites. Agents responsible for internal erosion differed among sites, with boring sponges being most abundant on the two inshore reefs, and molluscs most abundant at the offshore sites. Deposit-feeding polychaetes were more abundant at the two inshore sites, while filter and surface deposit feeders were more common at the offshore sites. Net erosion rates varied among sites (1.090±0.499 to 7.846±3.218 kg m²), and the relative importance of the components of erosion changed markedly along the cross-shelf transect.     

Black reefs: iron-induced phase shifts on coral reefs

The Line Islands are calcium carbonate coral reef platforms located in iron-poor regions of the central Pacific. Natural terrestrial run-off of iron is non-existent and aerial deposition is extremely low. However, a number of ship groundings have occurred on these atolls. The reefs surrounding the shipwreck debris are characterized by high benthic cover of turf algae, macroalgae, cyanobacterial mats and corallimorphs, as well as particulate-laden, cloudy water. These sites also have very low coral and crustose coralline algal cover and are call black reefs because of the dark-colored benthic community and reduced clarity of the overlying water column. Here we use a combination of benthic surveys, chemistry, metagenomics and microcosms to investigate if and how shipwrecks initiate and maintain black reefs. Comparative surveys show that the live coral cover was reduced from 40 to 60% to <10% on black reefs on Millennium, Tabuaeran and Kingman. These three sites are relatively large (>0.75 km²). The phase shift occurs rapidly; the Kingman black reef formed within 3 years of the ship grounding. Iron concentrations in algae tissue from the Millennium black reef site were six times higher than in algae collected from reference sites. Metagenomic sequencing of the Millennium Atoll black reef-associated microbial community was enriched in iron-associated virulence genes and known pathogens. Microcosm experiments showed that corals were killed by black reef rubble through microbial activity. Together these results demonstrate that shipwrecks and their associated iron pose significant threats to coral reefs in iron-limited regions.