A geospatial assessment of the relationship between reef flat community calcium carbonate production and wave energy

The ability of benthic communities inhabiting coral reefs to produce calcium carbonate underpins the development of reef platforms and associated sedimentary landforms, as well as the fixation of inorganic carbon and buffering of diurnal pH fluctuations in ocean surface waters. Quantification of the relationship between reef flat community calcium carbonate production and wave energy provides an empirical basis for understanding and managing this functionally important process. This study employs geospatial techniques across the reef platform at Lizard Island, Great Barrier Reef, to (1) map the distribution and estimate the total magnitude of reef community carbonate production and (2) empirically ascertain the influence of wave energy on community carbonate production. A World-View-2 satellite image and a field data set of 364 ground referencing points are employed, along with data on physical reef characteristics (e.g. bathymetry, rugosity) to map and validate the spatial distribution of the four major community carbonate producers (live coral, carbonate sand, green calcareous macroalgae and encrusting calcified algae) across the reef platform. Carbonate production is estimated for the complete reef platform from the composition of these community components. A synoptic model of wave energy is developed using the Simulating WAves Nearshore (SWAN) two-dimensional model for the entire reef platform. The relationship between locally derived measures of carbonate production and wave energy is evaluated at both the global scale and local scale along spatial gradients of wave energy traversing the reef platform. A wave energy threshold is identified, below which carbonate production levels appear to increase with wave energy and above which mechanical forcing reduces community production. This implies an optimal set of hydrodynamic conditions characterized by wave energy levels of approximately 300 J m^?2, providing an empirical basis for management of potential changes in community carbonate production associated with climate change-driven increases in wave energy.     

Carbonate production of an emergent reef platform, Warraber Island, Torres Strait, Australia

Complex relationships exist between tropical reef ecology, carbonate (CaCO₃) production and carbonate sinks. This paper investigated census-based techniques for determining the distribution and carbonate production of reef organisms on an emergent platform in central Torres Strait, Australia, and compared the contemporary budget with geological findings to infer shifts in reef productivity over the late Holocene. Results indicate that contemporary carbonate production varies by several orders of magnitude between and within the different reef-flat sub-environments depending on cover type and extent. Average estimated reef-flat production was 1.66 ± 1.78 kg m⁻² year⁻¹ (mean ± SD) although only 23% of the area was covered by carbonate producers. Collectively, these organisms produce 17,399 ± 18,618 t CaCO₃ year⁻¹, with production dominated by coral (73%) and subordinate contributions by encrusting coralline algae (18%) articulated coralline algae, molluscs, foraminifera and Halimeda (<4%). Comparisons between the production of these organisms across the different reef-flat zones, surface sediment composition and accumulation rates calculated from cores indicate that it is necessary to understand the spatial distribution, density and production of each major organism when considering the types and amounts of carbonate available for storage in the various reef carbonate sinks. These findings raise questions as to the reliability of using modal production rates in global models independent of ecosystem investigation, in particular, indicating that current models may overestimate reef productivity in emergent settings. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Silurian carbonate platforms and extinction events—ecosystem changes exemplified from Gotland, Sweden

Recent and ancient carbonate platforms are major marine ecosystems, built by various carbonate-secreting organisms with different sensitivity for environmental change. For this reason, carbonate platforms are excellent sensors for changes in contemporaneous marine environments. A variety of ecosystem changes in carbonate platforms have previously been recognised in the aftermath of mass extinction events. This paper addresses how two Silurian extinction events among graptolites, conodonts, and pentamerid brachiopods can be related to changes in the style of carbonate production and general evolution of low latitude carbonate platforms in a similar way as previously reported from the major five mass extinctions of the Phanerozoic. Strata formed on Gotland during the Mulde and Lau events share remarkably many similarities but are strikingly different in composition compared to other strata on the island. The event-related strata is characterised by the sudden appearance of widespread oolites, deviating reef composition, flat-pebble conglomerates, abundant micro- and macro-oncoids, stromatolites, and other microbial facies suggesting decreased bioturbation levels in contemporaneous shelf seas. Importantly, these changes can be tied to high-resolution biostratigraphic frameworks and global stable isotope excursions. The anomalous intervals may therefore be searched for elsewhere in order to test their regional or global significance.     

Distribution of gravel-sized empty tests of large benthic foraminifers as practical depositional indicators in tropical reef and shelf carbonate environments

To explore the utility of gravel-sized tests of large benthic foraminifers (LBFs) as practical paleoenvironmental indicators of tropical reef and shelf carbonate environments, depth and spatial distributions of gravel-sized empty tests of LBFs were examined using 39 surface sediment samples collected from depths shallower than 200 m off the west coast of Miyako Island (Ryukyu Islands, northwest Pacific). Distributions of the LBF tests were mainly related to water depth, topography, and substrate type. Q-mode cluster analysis based on the binary (presence/absence) data of LBF associations (4–2-mm size fraction) clearly delineates four depositional environments: bay, back reef to fore reef, flat shelf, and shelf slope. Application of this modern dataset to fossil LBF data from larger foraminiferal limestones of the Pleistocene Ryukyu Group indicate that a test section was deposited in an outer flat shelf at depths between 54 and 99 m. Comparisons of these results with previous reports suggest that our foraminiferal analysis using gravel-sized tests is methodologically easier than conventional analyses including smaller sized tests to distinguish similar levels of depositional environments. However, taxonomic and environmental similarities make the applicability of this dataset to fossil LBF data from Quaternary tropical carbonate environments in the northwest Pacific.     

Controls on reef development and the terrigenous-carbonate interface on a shallow shelf,Nicaragua (Central America)

Marine geology and physical oceanographic data collected during two field projects (∼4 months) on the Caribbean shelf of Nicaragua indicate a surprising dominance of carbonate deposition and reef growth on a shelf that is receiving an abnormally large volume of terrigenous sediments. High rainfall rates (∼400–500 cm/year), coupled with a warm tropical climate, encourage rapid denudation of the country’s central volcanic highland and transport of large volumes of terrigenous sediment and fresh water to the coast. Estimates suggest that three times more fresh water and fifteen times more sediment are introduced per unit length of coastline than on the east coast of the United States. Distribution of the terrigenous facies, development of carbonate sediment suites, and the location and quality of viable reefs are strongly controlled by the dynamic interaction near the coasts of highly turbid fresh to brackish water effluents from thirteen rivers with clear marine waters of the shelf. Oceanic water from the central Caribbean drift current intersects the shelf and moves slowely in a dominant northwest direction toward the Yucatan Channel. A sluggish secondary gyre moves to the south toward Costa Rica. In contrast, the turbid coastal water is deflected to the south in response to density gradients, surface water slopes, and momentum supplied by the steady northeast trade winds. A distinct two-layered flow is commonly present in the sediment-rich coastal boundary zone, which is typically 10–20 km wide. The low-salinity upper layer is frictionally uncoupled from the ambient shelf water and therefore can expand out of the normally coherent coastal boundary zone during periods of abnormal flooding or times when instability is introduced into the northeast trades. Reef distribution, abruptness of the terrigenous-carbonate interface, and general shelf morphology reflect the long-term dynamic structure of the shelf waters. A smooth-bottomed ramp of siliciclastic sands to silts and clays mantles the inner shelf floor in a linear belt paralleling the coast. This belt generally corresponds to the western flank of the coastal boundary zone. Occurrence of reefs is generally confined to areas outside this zone. Terrigenous clays and silts of the inner shelf are abruptly (<20 km from the coast) replaced byHalimeda-rich sediment of the middle and outer shelf. Within the carbonate facies belt, reef complexes thrive as small, isolated masses; large, reef-capped platforms; reef fringes around islands; and shelfedge structures with vertical relief that can exceed 25 m. In general, the frequency and proliferation of reefs increase away from the turbid coastal boundary layer and toward the cooler and saltier water that upwells at the shelf margin.     

Observations of the thermal environment on Red Sea platform reefs: a heat budget analysis

Hydrographic measurements were collected on nine offshore reef platforms in the eastern Red Sea shelf region, north of Jeddah, Saudi Arabia. The data were analyzed for spatial and temporal patterns of temperature variation, and a simple heat budget analysis was performed with the goal of advancing our understanding of the physical processes that control temperature variability on the reef. In 2009 and 2010, temperature variability on Red Sea reef platforms was dominated by diurnal variability. The daily temperature range on the reefs, at times, exceeded 5°C—as large as the annual range of water temperature on the shelf. Additionally, our observations reveal the proximity of distinct thermal microclimates within the bounds of one reef platform. Circulation on the reef flat is largely wave driven. The greatest diurnal variation in water temperature occurs in the center of larger reef flats and on reefs protected from direct wave forcing, while smaller knolls or sites on the edges of the reef flat tend to experience less diurnal temperature variability. We found that both the temporal and spatial variability in water temperature on the reef platforms is well predicted by a heat budget model that includes the transfer of heat at the air–water interface and the advection of heat by currents flowing over the reef. Using this simple model, we predicted the temperature across three different reefs to within 0.4°C on the outer shelf using only information about bathymetry, surface heat flux, and offshore wave conditions.     

Depositional environment and constraining factors on the facies architecture of the Qom Formation,Central Basin,Iran

In the Central Iran Basin, the mixed carbonate–siliciclastic deposits of the C member of the Qom Formation were deposited on a carbonate platform which is dominated by rhodalgal associations occurring in tropical–subtropical environment. The biogenic rhodalgal association is dominated by bryozoa, coralline red algae, bivalves and echinoids together with smaller amounts of photo-dependent biota including large benthic foraminifera and corals. The abundance of heterozoan association and the bloom of suspension-feeding organisms are the result of an increase in nutrient availability which has profound controlling effect on the biotic system. The low occurrence of symbiont-bearing benthic foraminifera and coral, typical of stable, oligotrophic condition, represents their low tolerance to unstable, nutrient-rich environment. In the investigated Oligocene–Miocene shallow marine carbonate succession, 10 different microfacies were distinguished through depositional texture and biotic components. The rock sequences investigated are referred to an open shelf carbonate platform in which the depositional environments range from outer shelf to inner shelf conditions.     

Latitudinal gradients in Atlantic reef fish communities: trophic structure and spatial use patterns

Trophic strategies and spatial use habits were investigated in reef fish communities. The results supported the hypothesis of differential use of food resources among tropical and higher latitude reef fishes, i.e . the number of species and relative abundance of fishes relying on relatively low‐quality food significantly decreased from tropical to temperate latitudes. The species : genus ratio of low‐quality food consumers increased toward the tropics, and was higher than the overall ratio considering all fishes in the assemblages. This supports the view that higher speciation rates occurred among this guild of fishes in warm waters. It was also demonstrated that density of herbivorous fishes (the dominant group relying on low‐quality food resources) in the western Atlantic decreased from tropical to temperate latitudes. Spatial use and mobility varied with latitude and consequently reef type and complexity. Fishes with small‐size home ranges predominated on tropical coral reefs.     

Siliciclastic‐carbonate ramp,a hostile world for carbonate shelf fauna? The example of the Givetian Ardenne Platform

The Givetian platform of the Ardennes Massif records several alternations between a siliciclastic‐carbonate ramp and a carbonate shelf. Usually these depositional contexts are considered as a major disruption implying a perturbation of many ecological parameters. We established the impact of these variations on the biodiversity structure through the study of the trophic organisation. Thanks to a previous microfacies analysis, 550 levels of the Mont d'Haurs section in Givet and ten associated environments were precisely defined. Seven palaeotrophic levels are recognized from micropalaeontological data, including benthic, planktonic, heterotroph and autotroph organisms. The spatial and temporal distributions of these levels have been analysed through means of multivariate analyses. The statistic results show that the distribution of the palaeotrophic levels during periods characterized by a mixed ramp is not significantly different than during carbonate shelf influences. These environmental modifications do not affect the community‐type. These results support recent studies performed on different benthic communities occurring in the Givetian of the Ardennes Massif. Indeed, trilobite and ostracod faunas of this period appear more affected by global environmental changes as the Ka?ák (uppermost Eifelian) and Taghanic (late middle Givetian) events. Moreover, along the proximal‐distal transect on the platform, the reef constitutes the most singular environment. However, contrary with the Frasnian reefs of the Ardennes Massif, which consist of carbonate mud mounds laying on a deep mixed ramp, there is no trilobite community that appears restricted to Givetian reefs. Thus global bioevents during the Middle Devonian have a more important control on the biodiversity than the platform morphology. These results support the view that at wide scales (spatial and temporal), the biodiversity responds more positively to ecological disturbances.     

Taphonomy of Early Permian benthic assemblages (Carnic Alps, Austria): carbonate dissolution versus biogenic carbonate precipitation

During the Early Permian, in the area of the Carnic Alps, a quartz-gravelly beach fringed a mixed siliciclastic-carbonate lagoon with fleshy algal meadows and oncoids; seaward, an ooid shoal belt graded down dip to a low-energy carbonate inner shelf with phylloid algal meadows. In limestones, foraminiferal biomurae and bioclast preservation record tapholoss by rotting of non-calcified organisms (interpreted as fleshy algae) and by dissolution of aragonitic fossils. Carbonate loss by dissolution was counteracted and, locally, perhaps exceeded by carbonate precipitation of encrusting foraminifera and as oncoids. Sites of abrasion and carbonate dissolution (beach), sites with tapholoss by rotting and dissolution, but with microbialite/foraminiferal carbonate precipitation (lagoon, inner shelf), and sites only of carbonate precipitation (ooid shoals) co-existed on discrete shelf compartments. Compartmentalized, contemporaneous carbonate dissolution and precipitation, to total amounts yet difficult to quantify, impede straightforward estimates of ancient carbonate sediment budget.     

Habitat effects and sampling bias on Phanerozoic reef distribution

Incomplete preservation, heterogeneous geographic sampling, uncertainties in palaeogeographic reconstructions and inconsistencies of reef definitions bias global reef patterns observed in the geological record. This sampling bias is added to a biological habitat area effect, which is thought to be of paramount importance for modern reefs. To evaluate the importance of sampling bias of ancient reefs, I first tested the habitat area effect and sampling bias for modern tropical reefs and then evaluated these factors for pre-Pleistocene Phanerozoic reefs. Results suggest that habitat area, although significantly affecting Phanerozoic reef patterns, is considerably less important than sampling bias. Sampling bias is more controlled by socioeconomic factors than by geological processes such as subduction of oceanic crust or sea-level fluctuations. Reefs are more likely to be sampled in rich countries, irrespective of geological and ecological controls. The numeric and geographic distribution of reefs as currently recorded in the published literature is thus probably largely artifactual, which may explain the scarcity of significant correlations between reef distribution patterns and inferred physico-chemical controls.     

Biostratigraphy and paleoecology of the Oligo-Miocene succession in Fars and Khuzestan areas (Zagros Basin,SW Iran)

Paleontological and biostratigraphical studies on carbonate platform succession from southwest Iran documented a great diversity of shallow-water benthic foraminifera during the Oligocene–Miocene. Larger foraminifera are the main means for the stratigraphic zonation of carbonate sediments. The distributions of larger benthic foraminifera in two outcrop sections (Abolhayat and Lali) in the Zagros Basin, Iran, are used to determine the age of the Asmari Formation. Four assemblage zones have been recognized by distribution of the larger benthic foraminifera in the study areas. Assemblage 3 (Aquitanian age) and 4 (Burdigalian age) have not been recognized in the Abolhayat section (Fars area), due to sea-level fall. The end Chattian sea-level fall restricted marine deposition in the Abolhayat section and Asmari Formation replaced laterally by the Gachsaran Formation. This suggests that the Miocene part of the formation as recognized in the Lali section (Khuzestan area) of the Zagros foreland basin is not present in the Abolhayat outcrop. The distribution of the Oligocene larger benthic foraminifera indicates that shallow marine carbonate sediments of the Asmari Formation at the study areas have been deposited in the photic zone of tropical to subtropical oceans. Based on analysis of larger benthic foraminiferal assemblages and microfacies features, three major depositional environments are identified. These include inner shelf, middle shelf and outer shelf. The inner shelf facies is characterized by wackestone–packstone, dominated by various taxa of imperforate foraminifera. The middle shelf is represented by packstone–grainstone to floatstone with a diverse assemblage of larger foraminifera with perforate wall. Basinwards is dominated by argillaceous wackestone characterized by planktonic foraminifera and large and flat nummulitidae and lepidocyclinidae. Planktonic foraminifera wackestone is the dominant facies in the outer shelf.     

Assessing the contribution of foraminiferan protists to global ocean carbonate production

Larger symbiont-bearing foraminifera are prominent and important producers of calcium carbonate in modern tropical environments. With an estimated production of at least 130 million tons of CaCO(3) per year, they contribute almost 5% of the annual present-day carbonate production in the world's reef and shelf areas (0-200 m) and approximately 2.5% of the CaCO(3) of all oceans. Together with non-symbiont-bearing smaller foraminifera, all benthic foraminifera are estimated to annually produce 200 million tons of calcium carbonate worldwide. The majority of foraminiferal calcite in modern oceans is produced by planktic foraminifera. With an estimated annual production of at least 1.2 billion tons, planktic foraminifera contribute more than 21% of the annual global ocean carbonate production. Total CaCO(3) of benthic and planktic foraminifera together amounts to 1.4 billion tons of calcium carbonate per year. This accounts to almost 25% of the present-day carbonate production of the oceans, and highlights the importance of foraminifera within the CaCO(3) budget of the world's oceans.     

Carbonate production by scleractinian corals at Aqaba,Gulf of Aqaba,Red Sea

Summary In a fringing reef at Aqaba at the northern end of the Gulf of Aqaba (29°26′N) growth rates, density, and the calcification rate ofPorites were investigated in order to establish calculations of gross carbonate production for the reefs in this area. Colony accretion ofPorites decreases with depth as a function of decreasing growth rates. The calcification rate ofPorites is highest in shallow water (0–5 m depth) with 0.9 g·cm⁻²·yr⁻¹ and falls down to 0.5 g·cm⁻²·yr⁻¹ below 30 m. Scleractinian coral gross production is calculated from potential productivity and coral coverage. It is mainly dependent on living coral cover and to a lesser extent on potential productivity. Total carbonate production on the reef ranged from 0 to 2.7 kg/m² per year, with a reef-wide average of 1.6 kg/m² perycar. Maximum gross carbonate production by corals at Aqaba occurs at the reef crest and in the middle fore-reef from 10 to 15 m water depth. Production is low in sandy reef parts. Below 30 m depth values still reach ca. 50% of shallow water values. Mean potential production of colonies and gross carbonate production of the whole reef community at Aqaba is lower than in tropical reefs. However, carbonate production is higher than in reef areas at the same latitude in the Pacific, indicating a northward shift of reef production in the Red Sea.     

Quantifying production rates and size fractions of parrotfish‐derived sediment: A key functional role on Maldivian coral reefs

Coral reef fish perform numerous important functional roles on coral reefs. Of these, carbonate sediment production, as a by‐product of parrotfish feeding, is especially important for contributing to reef framework construction and reef‐associated landform development. However, only limited data exist on: (i) how production rates vary among reef habitats as a function of parrotfish assemblages, (ii) the relative importance of sediment produced from eroded, reworked, and endogenous sources, or (iii) the size fractions of sediment generated by different parrotfish species and size classes. These parameters influence not only overall reef‐derived sediment supply, but also influence the transport potential and depositional fate of this sedimentary material. Here, we show that parrotfish sediment production varies significantly between reef‐platform habitats on an atoll‐margin Maldivian reef. Highest rates of production (over 0.8 kg m⁻² year⁻¹) were calculated in three of the eight platform habitats; a rubble‐dominated zone, an Acropora spp. dominated zone, and a patch reef zone. Habitat spatial extent and differences in associated parrotfish assemblages strongly influenced the total quantities of sediment generated within each habitat. Nearly half of total parrotfish sediment production occurred in the rubble habitat, which comprised only 8% of the total platform area. Over 90% of this sedimentary material originated from eroded reef framework as opposed to being reworked existing or endogenously produced sediment, and comprised predominantly coral sands (predominantly 125–1000 µm in diameter). This is comparable to the dominant sand types and size fractions found on Maldivian reef islands. By contrast, nearly half of the sediment egested by parrotfish in the Acropora spp. dominated and patch reef habitats resulted from reworked existing sediments. These differences between habitats are a result of the different parrotfish assemblages supported. Endogenous carbonate production was found to be insignificant compared to the quantity of eroded and reworked material. Our findings have important implications for identifying key habitats and species which act as major sources of sediment for reef‐island systems.     

Spatial resilience of the Great Barrier Reef under cumulative disturbance impacts

In the face of increasing cumulative effects from human and natural disturbances, sustaining coral reefs will require a deeper understanding of the drivers of coral resilience in space and time. Here we develop a high‐resolution, spatially explicit model of coral dynamics on Australia's Great Barrier Reef (GBR). Our model accounts for biological, ecological and environmental processes, as well as spatial variation in water quality and the cumulative effects of coral diseases, bleaching, outbreaks of crown‐of‐thorns starfish (Acanthaster cf. solaris), and tropical cyclones. Our projections reconstruct coral cover trajectories between 1996 and 2017 over a total reef area of 14,780 km², predicting a mean annual coral loss of ?0.67%/year mostly due to the impact of cyclones, followed by starfish outbreaks and coral bleaching. Coral growth rate was the highest for outer shelf coral communities characterized by digitate and tabulate Acropora spp. and exposed to low seasonal variations in salinity and sea surface temperature, and the lowest for inner‐shelf communities exposed to reduced water quality. We show that coral resilience (defined as the net effect of resistance and recovery following disturbance) was negatively related to the frequency of river plume conditions, and to reef accessibility to a lesser extent. Surprisingly, reef resilience was substantially lower within no‐take marine protected areas, however this difference was mostly driven by the effect of water quality. Our model provides a new validated, spatially explicit platform for identifying the reefs that face the greatest risk of biodiversity loss, and those that have the highest chances to persist under increasing disturbance regimes.     

Early Permian facies and paleogeography of the Southeastern Russian craton

Summary During the Early Permian deep-water basins existed in the southeastern part of the Russian craton. North and west of the Cis-Ural foredeep and the Precaspian depression (micro-ocean) carbonate platforms were formed on a shallow-marine shelf during the Asselian, Sakmarian and Early Artinskian. Reefs developed on the margin of these platforms along the slopes of the Cis-Ural foredeep and the Precaspian depression. The reefs shifted platform ward in the eastern areas, due to the tectonic subsidence of the platform margin and at the same time, prograded basinward in the south. Movements of continental blocks from the south during the Late Artinskian and Kungurian caused the separation of the Early Permian basin of the Russian craton from the Palaeo-Tethys, followed by evaporite sedimentation in the restricted basins. The existence of source rocks (bituminous deep-water sediments), thick reservoir rocks (limestones and dolostones), evaporitic seals and structural as well as stratigraphic traps are responsible for large productive gas and oil fields (e.g., Orenburg field), some of which are distinctly associated with reef carbonates.     

Late Triassic reefs from the Northwest and South Tethys: distribution, setting, and biotic composition

The paleolatitudinal distribution patterns during Ladinian and Carnian time are characterized by an increasing expansion of reefs from the northern to the southern hemisphere. The optimum of reef diversity and frequency in the Norian is associated with the development of extended attached or isolated carbonate platforms. Norian-Rhaetian sponge and coral reefs of the Northern Calcareous Alps developed (1) as reef belt composed of patch reefs in platform-edge positions facing the open-marine northwestern Tethys basins and (2) as patch reefs in intraplatform basins as well as in ramp positions.Carnian and Norian-Rhaetian sponge and coral reefs of the Arabian Peninsula are formed (1) as reef complexes at the margins of carbonate platforms on the tops of volcanic seamounts in the southern Tethyan ocean, as small biostromes on these isolated platforms, and (2) as transgressive reef complexes on the attached platform of the Gondwana margin. The Norian Gosaukamm reefal breccia of the NW Tethys is a counterpart of Jabal Wasa reefal limestone of the Gondwana margin with similarities in geological setting and biotic composition. Rhaetian coral biostromes of low diversity known from the Austrian Koessen basin resemble to the time equivalent Ala biostromes of the isolated Kawr platform in the southern Neo-Tethys by forming a discontinuous layer in shallow intraplatform basin setting.     

广西南丹拉要泥盆纪礁组合剖面的古生态特征

广西南丹拉要泥盆纪生物礁发育在台地的边缘,是一个礁、滩、开阔台地沉积互相交替的叠置礁。礁组合中出现5个主要的生物群落。其中以层孔虫、床板珊瑚为主的 Stromatopora-Alveolites群落是最重要的造礁群落。这些群落有规律地分布在礁组合的不同部位。它们在剖面上的垂直分布显示礁组合的发育过程分成3个大的旋回,其中包括9个次一级的成礁旋回。旋回反映了海水深浅的变化。