Linkages between coral assemblages and coral proxies of terrestrial exposure along a cross-shelf gradient on the southern Great Barrier Reef

Coral core records, combined with measurements of coral community structure, were used to assess the long-term impact of multiple environmental stressors on reef assemblages along an environmental gradient. Multiple proxies (luminescent lines, Ba/Ca, δ¹⁵N) that reflect different environmental conditions (freshwater discharge, sediment delivery to the nearshore, nutrient availability and transformations) were measured in Porites coral cores collected from nearshore reefs at increasing distance from the intensively agricultural region of Mackay (Queensland, Australia). The corals provide a record (1968–2002) of the frequency and intensity of exposure to terrestrial runoff and fertilizer-derived nitrogen and were used to assess how the present-day coral community composition may have been influenced by flood-related disturbance. Reefs closest to the mainland (5–32 km offshore) were characterized by low hard coral cover (≤10%), with no significant differences among locations. Distinct annual luminescent lines and elevated Ba/Ca values (4.98 ± 0.63 μmol mol⁻¹; mean ± SD) in the most inshore corals (Round Top Island; 5 km offshore) indicated chronic, sub-annual exposure to freshwater and resuspended terrestrial sediment that may have historically prevented reef formation. By contrast, corals from Keswick Island (32 km offshore) indicated episodic, high-magnitude exposure to Pioneer River discharge during extreme flood events (e.g., 1974, 1991), with strongly luminescent lines and substantially enriched coral skeletal δ¹⁵N (12–14‰). The reef assemblages at Keswick and St. Bees islands were categorically different from all other locations, with high fleshy macroalgal cover (80.1 ± 7.2% and 62.7 ± 7.1%, respective mean ± SE) overgrowing dead reef matrix. Coral records from Scawfell Island (51 km offshore) indicated little exposure to Pioneer catchment influence: all locations from Scawfell and further offshore had total hard and soft coral cover comparable to largely undisturbed nearshore to middle shelf reefs of the southern Great Barrier Reef.     

Spatial and seasonal reef calcification in corals and calcareous crusts in the central Red Sea

The existence of coral reef ecosystems critically relies on the reef carbonate framework produced by scleractinian corals and calcareous crusts (i.e., crustose coralline algae). While the Red Sea harbors one of the longest connected reef systems in the world, detailed calcification data are only available from the northernmost part. To fill this knowledge gap, we measured in situ calcification rates of primary and secondary reef builders in the central Red Sea. We collected data on the major habitat-forming coral genera Porites, Acropora, and Pocillopora and also on calcareous crusts (CC) in a spatio-seasonal framework. The scope of the study comprised sheltered and exposed sites of three reefs along a cross-shelf gradient and over four seasons of the year. Calcification of all coral genera was consistent across the shelf and highest in spring. In addition, Pocillopora showed increased calcification at exposed reef sites. In contrast, CC calcification increased from nearshore, sheltered to offshore, exposed reef sites, but also varied over seasons. Comparing our data to other reef locations, calcification in the Red Sea was in the range of data collected from reefs in the Caribbean and Indo-Pacific; however, Acropora calcification estimates were at the lower end of worldwide rates. Our study shows that the increasing coral cover from nearshore to offshore environments aligned with CC calcification but not coral calcification, highlighting the potentially important role of CC in structuring reef cover and habitats. While coral calcification maxima have been typically observed during summer in many reef locations worldwide, calcification maxima during spring in the central Red Sea indicate that summer temperatures exceed the optima of reef calcifiers in this region. This study provides a foundation for comparative efforts and sets a baseline to quantify impact of future environmental change in the central Red Sea.

     

Environmental Records from Great Barrier Reef Corals: Inshore versus Offshore Drivers

The biogenic structures of stationary organisms can be effective recorders of environmental fluctuations. These proxy records of environmental change are preserved as geochemical signals in the carbonate skeletons of scleractinian corals and are useful for reconstructions of temporal and spatial fluctuations in the physical and chemical environments of coral reef ecosystems, including The Great Barrier Reef (GBR). We compared multi-year monitoring of water temperature and dissolved elements with analyses of chemical proxies recorded in Porites coral skeletons to identify the divergent mechanisms driving environmental variation at inshore versus offshore reefs. At inshore reefs, water Ba/Ca increased with the onset of monsoonal rains each year, indicating a dominant control of flooding on inshore ambient chemistry. Inshore multi-decadal records of coral Ba/Ca were also highly periodic in response to flood-driven pulses of terrigenous material. In contrast, an offshore reef at the edge of the continental shelf was subject to annual upwelling of waters that were presumed to be richer in Ba during summer months. Regular pulses of deep cold water were delivered to the reef as indicated by in situ temperature loggers and coral Ba/Ca. Our results indicate that although much of the GBR is subject to periodic environmental fluctuations, the mechanisms driving variation depend on proximity to the coast. Inshore reefs are primarily influenced by variable freshwater delivery and terrigenous erosion of catchments, while offshore reefs are dominated by seasonal and inter-annual variations in oceanographic conditions that influence the propensity for upwelling. The careful choice of sites can help distinguish between the various factors that promote Ba uptake in corals and therefore increase the utility of corals as monitors of spatial and temporal variation in environmental conditions.     

珊瑚礁区碳循环研究进展

珊瑚礁是海洋中生产力水平最高的生态系统之一,其碳循环受到有机碳代谢(光合作用/呼吸作用)和无机碳代谢(钙化/溶解)两大代谢过程的共同作用,过程十分复杂.珊瑚礁植物的光合作用保证了有机碳的有效补充,动物摄食及微生物降解等生物过程驱动了珊瑚礁区有机碳高效循环,只有不超过7％的有机碳进入沉积物,而向大洋区水平输出的有机碳通量变化幅度较大,主要受到水动力条件的影响.珊瑚礁区碳酸盐沉积(无机碳代谢)是全球碳酸盐库的重要组成部分,年累积量达到全球CaCO3年累积量的23%～26%,是影响大气CO2浓度的重要组成;珊瑚礁是大气CO2源或汇则取决于净有机生产力与净无机生产力的比值(ROI),当ROI <0.6时,珊瑚礁区是大气CO2的源,反之,则是大气CO2的汇.     

Environmental factors associated with the spatial distribution of crustose coralline algae on the Great Barrier Reef

Crustose coralline algae (CCA) fulfill two key functional roles in coral reef ecosystems: they contribute significantly to reef calcification, and they induce larval settlement of many benthic organisms. Percentage cover of CCA, and environmental conditions, were visually estimated on 144 reefs of the Great Barrier Reef between 10 and 24° latitude S. Reefs were located across the shelf and ranged from turbid near-shore reefs close to rivers to clean-water reefs hundreds of kilometers from coastal influences. On each reef, two sites were surveyed between 0.5 and 18 m depth. Strong cross-shelf trends occurred in cover of CCA, amount of sediment deposited, water clarity, and slope angle. Relative distance across the shelf and sedimentation jointly explained 84% of variation in CCA cover. Three regions running parallel to the shore were identified, with a mean CCA cover of <1% on the inner third of the shelf, and >20% cover on the outer half of the shelf, with a narrow transition region between the two. Within each region, the cover of CCA was unrelated to distance across the shelf, but was related to the sedimentary environment, being relatively higher on reefs with low sediment deposits. On the inner third of the shelf, the most sediment-exposed reefs were unsuitable habitats for CCA. The inverse relationship between CCA and sediment has implications for the recruitment of CCA-specialised organisms, and for rates of reef calcification.     

Dissolved inorganic carbon and total alkalinity of a Hawaiian fringing reef: chemical techniques for monitoring the effects of ocean acidification on coral reefs

There is an interest in developing approaches to “ecosystem-based” management for coral reefs. One aspect of ecosystem performance is to monitor carbon metabolism of whole communities. In an effort to explore robust techniques to monitor the metabolism of fringing reefs, especially considering the possible effects of ocean acidification, a yearlong study of the carbonate chemistry of a nearshore fringing reef in Hawaii was conducted. Diurnal changes in seawater carbonate chemistry were measured once a week in an algal-dominated and a coral-dominated reef flat on the Waimanalo fringing reef, Hawaii, from April of 2010 until May of 2011. Calculated rates of gross primary production (GPP) and net community calcification (G) were similar to previous estimates of community metabolism for other coral reefs (GPP 971 mmol C m^?2 d^?1; G 186 mmol CaCO₃ m^?2 d^?1) and indicated that this reef was balanced in terms of organic metabolism, exhibited net calcification, and was a net source of CO₂ to the atmosphere. Average slopes of total alkalinity versus dissolved inorganic carbon (TA–DIC slope) for the coral-dominated reef flat exhibited a greater calcification-to-net photosynthesis ratio than for the algal-dominated reef flat (coral slope vs. algal slope). Over the course of the time series, TA–DIC slopes remained significantly different between sites and were not correlated with diurnal averages in reef-water residence time or solar irradiance. These characteristic slopes for each reef flat reflect the relationship between carbon and carbonate community metabolism and can be used as a tool to monitor ecosystem function in response to ocean acidification.     

The 1991–1992 rapid ecological assessment of Palau’s coral reefs

At the request of the Palau and US governments, a team of 30 scientists under the leadership of the Nature Conservancy completed a rapid ecological assessment (REA) of nearshore marine resources in Palau in 1992. The REA provided ecological input to Palau's ongoing master plan for economic development and identified 45 marine sites worthy of special protection. The REA relied on previous literature, 1992 aerial photography, interviews, and field observations. A combination of qualitative and quantitative techniques were used to assess stony corals, other reef invertebrates, reef and shore fishes, macroscopic algae, seagrasses, sea turtles and other marine organisms. The REA covered a variety of coral reef habitats including beaches, seagrass beds, fringing reefs, lagoons, passes, channels, reef holes, patch and pinnacle reefs, barrier reefs, atolls, submerged reefs, mangroves, and rock islands. Major stresses to Palau's coral reefs include sedimentation from soil erosion, overfishing, and damage from periodic storms and waves. Minor stresses include dredge-and fill activities, sewage pollution, anchor damage, tourism use, ship groundings, aquarium fish collecting, and minor crown-of-thorns (Acanthaster) infestations.     

珊瑚共生体碳代谢特征研究进展

珊瑚礁作为一种典型的海洋生态系统,具有巨大的固碳和储碳潜力。然而,目前对于珊瑚礁的净碳能力(碳释放与碳吸收)仍存在争议,主要归因于珊瑚共生体碳代谢的多样性和复杂性。珊瑚礁在生物钙化、呼吸过程中向大气释放二氧化碳(CO₂);但在生物合成和沉积过程中却可以将碳进行固定与埋藏;为此,珊瑚礁的碳源碳汇身份还有待明确。现有部分研究表明,共生体通过碳代谢可以促进珊瑚礁吸收大气中的CO₂。此外,珊瑚礁和海岸带蓝碳生态系统通常表现出很强的连通性,珊瑚共生体碳代谢能有效提高海岸带盐沼植被、海草床、海洋浮游植物等生物的碳汇功能。为了加深对珊瑚礁碳源-碳汇功能的理解,综述了珊瑚共生体的碳代谢特征,梳理了共生体中碳的关键生态过程(有机碳的迁移、无机碳的转化、两者的赋存状态),总结了细菌-虫黄藻-病毒在共生体碳代谢中的作用,评述了珊瑚礁碳源-碳汇特征及影响因子。旨在阐明珊瑚共生体碳代谢的关键过程,并基于此寻求有效的珊瑚礁碳增汇技术,形成以碳增量为主的珊瑚保护与修复技术,提升珊瑚礁在蓝碳生态系统中的贡献。     

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

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

Survival of hurricane-generated coral fragments and a disturbance model of reef calcification/growth rates

Summary Hurricane Gerta, with winds reaching 150 km/h, crossed the Belize barrier reef on September 18, 1978. Breakage and scouring of corals occurred in all zones of the reef to a depth of approximately 25 m. Survivorship of storm-generated coral fragments and detached colonies is strongly size dependent, conforming to the power function Y=4.44X^0.66 where Y is the percent of fragments and X is the fragment size. Forty-six percent of detached Acropora palmata branches, which are larger ( =37.6 cm long) than fragments of other species ( =16.7 cm long), survived. Overall, 39% of fragments and detached colonies survived. This high survivorship, which probably increased the total number of colonies present, and redistribution of corals may explain the rapid recovery of reefs from all but the severest hurricanes. Storms appear to prevent coral reefs from reaching a mature state characterized by low calcification and growth rates. Therefore, we suggest that long-term reef calcification and growth rates are highest on reefs periodically distrubed by storms of intermediate intensity.Smithsonian Tropical Research Institute, Box 2072, Balboa, Republic of Panama     

How will coral reef fish communities respond to climate-driven disturbances? Insight from landscape-scale perturbations

Global climate change is rapidly altering disturbance regimes in many ecosystems including coral reefs, yet the long-term impacts of these changes on ecosystem structure and function are difficult to predict. A major ecosystem service provided by coral reefs is the provisioning of physical habitat for other organisms, and consequently, many of the effects of climate change on coral reefs will be mediated by their impacts on habitat structure. Therefore, there is an urgent need to understand the independent and combined effects of coral mortality and loss of physical habitat on reef-associated biota. Here, we use a unique series of events affecting the coral reefs around the Pacific island of Moorea, French Polynesia to differentiate between the impacts of coral mortality and the degradation of physical habitat on the structure of reef fish communities. We found that, by removing large amounts of physical habitat, a tropical cyclone had larger impacts on reef fish communities than an outbreak of coral-eating sea stars that caused widespread coral mortality but left the physical structure intact. In addition, the impacts of declining structural complexity on reef fish assemblages accelerated as structure became increasingly rare. Structure provided by dead coral colonies can take up to decades to erode following coral mortality, and, consequently, our results suggest that predictions based on short-term studies are likely to grossly underestimate the long-term impacts of coral decline on reef fish communities.     

River runoff reconstructions from novel spectral luminescence scanning of massive coral skeletons

Inshore massive corals often display bright luminescent lines that have been linked to river flood plumes into coastal catchments and hence have the potential to provide a long-term record of hinterland precipitation. Coral luminescence is thought to result from the incorporation of soil-derived humic acids transported to the reef during major flood events. Corals far from terrestrial sources generally only exhibit dull relatively broad luminescence bands, which are attributed to seasonal changes in coral density. We therefore tested the hypothesis that spectral ratios rather than conventional luminescence intensity provide a quantitative proxy record of river runoff without the confounding effects of seasonal density changes. For this purpose, we have developed a new, rapid spectral luminescence scanning (SLS) technique that splits emission intensities into red, green and blue domains (RGB) for entire cores with an unprecedented linear resolution of 71.4 μm. Since humic acids have longer emission wavelength than the coral aragonite, normalisation of spectral emissions should yield a sensitive optical humic acid/aragonite ratio for humic acid runoff, i.e., G/B ratio. Indeed, G/B ratios rather than intensities are well correlated with Ba/Ca, a geochemical coral proxy for sediment runoff, and with rainfall data, as exemplified for coral records from Madagascar. Coral cores also display recent declining trends in luminescence intensity, which are also reported in corals elsewhere. Such trends appear to be associated with a modern decline in skeletal densities. By contrast, G/B spectral ratios not only mark the impact of individual cyclones but also imply that humic acid runoff increased in Madagascar over the past few decades while coral skeletal densities decreased. Consequently, the SLS technique deconvolves the long-term interplay between humic acid incorporation and coral density that have confounded earlier attempts to use luminescence intensities as a proxy for river runoff.     

Sediment suppresses herbivory across a coral reef depth gradient

Sediments are a ubiquitous feature of all coral reefs, yet our understanding of how they affect complex ecological processes on coral reefs is limited. Sediment in algal turfs has been shown to suppress herbivory by coral reef fishes on high-sediment, low-herbivory reef flats. Here, we investigate the role of sediment in suppressing herbivory across a depth gradient (reef base, crest and flat) by observing fish feeding following benthic sediment reductions. We found that sediment suppresses herbivory across all reef zones. Even slight reductions on the reef crest, which has 35 times less sediment than the reef flat, resulted in over 1800 more herbivore bites (h⁻¹ m⁻²). The Acanthuridae (surgeonfishes) were responsible for over 80 per cent of all bites observed, and on the reef crest and flat took over 1500 more bites (h⁻¹ m⁻²) when sediment load was reduced. These findings highlight the role of natural sediment loads in shaping coral reef herbivory and suggest that changes in benthic sediment loads could directly impair reef resilience.     

Synergistic impacts of global warming on the resilience of coral reefs

Recent epizootics have removed important functional species from Caribbean coral reefs and left communities vulnerable to alternative attractors. Global warming will impact reefs further through two mechanisms. A chronic mechanism reduces coral calcification, which can result in depressed somatic growth. An acute mechanism, coral bleaching, causes extreme mortality when sea temperatures become anomalously high. We ask how these two mechanisms interact in driving future reef state (coral cover) and resilience (the probability of a reef remaining within a coral attractor). We find that acute mechanisms have the greatest impact overall, but the nature of the interaction with chronic stress depends on the metric considered. Chronic and acute stress act additively on reef state but form a strong synergy when influencing resilience by intensifying a regime shift. Chronic stress increases the size of the algal basin of attraction (at the expense of the coral basin), whereas coral bleaching pushes the system closer to the algal attractor. Resilience can change faster—and earlier—than a change in reef state. Therefore, we caution against basing management solely on measures of reef state because a loss of resilience can go unnoticed for many years and then become disproportionately more difficult to restore.     

The role of turtles as coral reef macroherbivores

Herbivory is widely accepted as a vital function on coral reefs. To date, the majority of studies examining herbivory in coral reef environments have focused on the roles of fishes and/or urchins, with relatively few studies considering the potential role of macroherbivores in reef processes. Here, we introduce evidence that highlights the potential role of marine turtles as herbivores on coral reefs. While conducting experimental habitat manipulations to assess the roles of herbivorous reef fishes we observed green turtles (Chelonia mydas) and hawksbill turtles (Eretmochelys imbricata) showing responses that were remarkably similar to those of herbivorous fishes. Reducing the sediment load of the epilithic algal matrix on a coral reef resulted in a forty-fold increase in grazing by green turtles. Hawksbill turtles were also observed to browse transplanted thalli of the macroalga Sargassum swartzii in a coral reef environment. These responses not only show strong parallels to herbivorous reef fishes, but also highlight that marine turtles actively, and intentionally, remove algae from coral reefs. When considering the size and potential historical abundance of marine turtles we suggest that these potentially valuable herbivores may have been lost from many coral reefs before their true importance was understood.     

The Changing Health of Coral Reefs

Throughout their entire global range coral reefs are in decline. Coral bleaching, macroalgal overgrowth and coral diseases — responses signaling the declining health of coral reefs — have occurred with increasing frequency and intensity in recent decades. Decreased calcification may also be affecting coral reefs over longer time scales. Declines in coral reef health have been attributed to various natural and anthropogenic processes, but assignment of causality has proved problematic. Coral bleaching has been observed during extreme climate events such as El Niño; furthermore, there are indications that exposure to UV radiation, air, infectious microbes, and elevated temperature plays a role in the dramatic increase of coral bleaching since the mid-1970s. Macroalgal overgrowth is usually ascribed to eutrophi-cation and coral diseases to weakening of the coral host resistance by anthropogenic pollution. An issue precluding a strict anthropogenic cause of coral reef decline is that both overgrowth and coral diseases are known to occur, although less frequently, on reefs remote from human development. While its causes are still being unraveled, the overall decline in coral reef health sends an unambiguous signal that the coral reef system is losing its ability to withstand sudden or persistent environmental changes.     

Framework of barrier reefs threatened by ocean acidification

To date, studies of ocean acidification (OA) on coral reefs have focused on organisms rather than communities, and the few community effects that have been addressed have focused on shallow back reef habitats. The effects of OA on outer barrier reefs, which are the most striking of coral reef habitats and are functionally and physically different from back reefs, are unknown. Using 5‐m long outdoor flumes to create treatment conditions, we constructed coral reef communities comprised of calcified algae, corals, and reef pavement that were assembled to match the community structure at 17 m depth on the outer barrier reef of Moorea, French Polynesia. Communities were maintained under ambient and 1200 μatm pCO₂ for 7 weeks, and net calcification rates were measured at different flow speeds. Community net calcification was significantly affected by OA, especially at night when net calcification was depressed ~78% compared to ambient pCO₂. Flow speed (2–14 cm s^?1) enhanced net calcification only at night under elevated pCO₂. Reef pavement also was affected by OA, with dissolution ~86% higher under elevated pCO₂ compared to ambient pCO₂. These results suggest that net accretion of outer barrier reef communities will decline under OA conditions predicted within the next 100 years, largely because of increased dissolution of reef pavement. Such extensive dissolution poses a threat to the carbonate foundation of barrier reef communities.     

Coral reefs in an urban embayment in Hawaii: a complex case history controlled by natural and anthropogenic stress

The effects of natural and anthropogenic stress need to be separated before coral reef ecosystems can be effectively managed. In this paper, a 25 year case history of coral reefs in an urban embayment (Mamala Bay) off Honolulu, Hawaii is described and differences between natural and man-induced stress are distinguished. Mamala Bay is a 30 km long shallow coastal bay bordering the southern (leeward) shore of Oahu and the city of Honolulu in the Hawaiian Islands. During the last 25 years, this area has been hit by two magnitude 5 hurricane events (winds >240 km/h) generating waves in excess of 7.5 m. Also during this period, two large sewer outfalls have discharged up to 90 million gallons per day (mgd) or (360x10⁶ L/day) of point source pollution into the bay. Initially the discharge was raw sewage, but since 1977 it has received advanced brimary treatment. Non-point source run-off from the Honolulu watershed also enters the bay on a daily basis. The results of the study show that discharge of raw sewage had a serious but highly localized impact on shallow (10m) reef corals in the bay prior to 1977. After 1977, when treatment was upgraded to the advanced primary level and outfalls were extended to deep water (>65 m), impacts to reef corals were no longer significant. No measurable effects of either point or non-point source pollution on coral calcification, growth, species composition, diversity or community structure related to pollution can now be detected. Conversely the effects of hurricane waves in 1982 and 1992 together caused major physical destruction to the reefs. In 1982, average coral cover of well-developed offshore reefs dropped from 60–75% to 5–15%. Only massive species in high relief areas survived. Today, recovery is occurring, and notwithstanding major future disturbance events, long-term biological processes should eventually return the coral ecosystems to a more mature successional stage. This case history illustrates the complex nature of the cumulative effects of natural and anthropogenic stress on coral reefs and the need for a long-term data base before the status of a coral reef can be properly interpreted.     

Short-Term Coral Bleaching Is Not Recorded by Skeletal Boron Isotopes

Coral skeletal boron isotopes have been established as a proxy for seawater pH, yet it remains unclear if and how this proxy is affected by seawater temperature. Specifically, it has never been directly tested whether coral bleaching caused by high water temperatures influences coral boron isotopes. Here we report the results from a controlled bleaching experiment conducted on the Caribbean corals Porites divaricata, Porites astreoides, and Orbicella faveolata. Stable boron (δ¹¹B), carbon (δ¹³C), oxygen (δ¹⁸O) isotopes, Sr/Ca, Mg/Ca, U/Ca, and Ba/Ca ratios, as well as chlorophyll a concentrations and calcification rates were measured on coral skeletal material corresponding to the period during and immediately after the elevated temperature treatment and again after 6 weeks of recovery on the reef. We show that under these conditions, coral bleaching did not affect the boron isotopic signature in any coral species tested, despite significant changes in coral physiology. This contradicts published findings from coral cores, where significant decreases in boron isotopes were interpreted as corresponding to times of known mass bleaching events. In contrast, δ¹³C and δ¹⁸O exhibited major enrichment corresponding to decreases in calcification rates associated with bleaching. Sr/Ca of bleached corals did not consistently record the 1.2°C difference in seawater temperature during the bleaching treatment, or alternatively show a consistent increase due to impaired photosynthesis and calcification. Mg/Ca, U/Ca, and Ba/Ca were affected by coral bleaching in some of the coral species, but the observed patterns could not be satisfactorily explained by temperature dependence or changes in coral physiology. This demonstrates that coral boron isotopes do not record short-term bleaching events, and therefore cannot be used as a proxy for past bleaching events. The robustness of coral boron isotopes to changes in coral physiology, however, suggests that reconstruction of seawater pH using boron isotopes should be uncompromised by short-term bleaching events.     

Response of triassic reef coral communities to sea-level fluctuations, storms and sedimentation: Evidence from a spectacular outcrop (Adnet, Austria)

Summary The Upper Rhaetian coral limestone of Adnet, southeast of Salzburg Austria has been repeatedly referred to as one of the most spectacular examples of an ancient ‘autochthonous’ coral reef structure. The ‘Tropfbruch’ quarry is probably the best outcrop for interpreting the distributional patterns of biotic successions and communities of a late Triassic patch reef. Our study is based on the interpretation of a) outcrop photographs, b) reef maps resulting from quadrat transects, and c) the analysis of quantitative data describing the distribution and frequency of reef organisms and sediment. A new methodological approach (combination of reef mapping and photo-transects) is used to obtain quantitative field data which can be compared in greater detail with data from modern coral reefs investigated by corresponding quantitative surveys. Three unconformities and three well-defined ‘reef growth stages’ reflecting the vertical and lateral development of the reef structure were differrentiated using transects: Stage 1, representing the reef growth optimum, is characterized by laterally differentiated coral reef knobs with corals in growth position. Criteria supporting this interpretation are the extraordinary size of the corals, their preservation in situ and the great thickness of this interval. The massive coralPamiroseris grew under higher energy conditions at the rim of the reef knob, whereas branchingRetiophyllia colonies preferred less agitated water in the center. Vertical changes are reflected by an increase in frequency of the dasycladacean algaDiplopora adnetensis and by the decreasing size ofRetiophyllia. These sedimentological and biological criteria together with the unconformity above indicate a fall in the sea level as a major control mechanism. Stage 2, separated from stage 1 by an unconformity caused by partial subaerial exposure and karstification, is characterized by vertically stacked coral successions with diverse reef debris. Facies heterogeneity is reflected by differences in the diversity, taphonomy and packing density of reef-building organisms as well as by differences in sediment input from the platform. Water depths and accommodation space were lower, therefore minor sea level fluctuations had a stronger effect on the biotic composition. The high percentage of coral debris and corals reworked by storms and the increase in the input of platform sediment led to a reduction of reef growth. Stage 3, again separated at the base by an unconformity associated with karstification, is characterized by bioclastic sediments with isolated reefbuilders forming a level-bottom community. The distribution of different coral morphotypes suggests that sea level fluctuations were not the only controlling factor. Variations in the substrate were caused by differences in the input of platform sediment. The three-step development seen in Adnet documents the response of low-diverse coral associations to variations caused by small-scale sea level changes, storm activity and sedimentation. The vertical changes in reef community structures correspond to a sequence of ‘allogenic replacements’. The Adnet reef structure should not be regarded as a general model of Alpine Upper Rhaetian reefs, because of the particular setting of the patch reef. Only the ‘capping beds’ of the Upper Rhaetian Reef Limestone of the Steinplatte exhibit criteria similar to Adnet. Potential modern analogues of features seen in the coral communities of Adnet are the internal structure of theRetiophyllia thickets, the key role of branching corals within the communities, the scattered distribution and low and even diversity of corals subsequent to breaks in settlement, segration patterns of corals indicating ‘contact avoidance’, toppling of large coral colonies by intensive boring, and decreasing coral coverage from deeper and sheltered settings to more shallower water depths.