Presence of competitors influences photosynthesis, but not growth, of the hard coral Porites cylindrica at elevated seawater CO2

Changes in environmental conditions, such as those caused by elevated carbon dioxide (CO2), potentially alter the outcome of competitive interactions between species. This study aimed to understand how elevated CO2 could influence competitive interactions between hard and soft corals, by investigating growth and photosynthetic activity of Porites cylindrica (a hard coral) under elevated CO2 and in the presence of another hard coral and two soft coral competitors. Corals were collected from reefs around Orpheus and Pelorus Islands on the Great Barrier Reef, Australia. They were then exposed to elevated pCO2 for 4 weeks with two CO2 treatments: intermediate (pCO2 648) and high (pCO2 1003) compared with a control (unmanipulated seawater) treatment (pCO2 358). Porites cylindrica growth did not vary among pCO2 treatments, regardless of the presence and type of competitors, nor was the growth of another hard coral species, Acropora cerealis, affected by pCO2 treatment. Photosynthetic rates of P. cylindrica were sensitive to variations in pCO2, and varied between the side of the fragment facing the competitors vs. the side facing away from the competitor. However, variation in photosynthetic rates depended on pCO2 treatment, competitor identity, and whether the photosynthetic yields were measured as maximum or effective photosynthetic yield. This study suggests that elevated CO2 may impair photosynthetic activity, but not growth, of a hard coral under competition and confirms the hypothesis that soft corals are generally resistant to elevated CO2. Overall, our results indicate that shifts in the species composition in coral communities as a result of elevated CO2 could be more strongly related to the individual tolerance of different species rather than a result of competitive interactions between species.

Seawater carbonate chemistry and calcification during tropical reef studies at Moorea (French Polynesia), 1997

Community metabolism and air-sea carbon dioxide (CO2) fluxes were investigated in July 1992 on a fringing reef at Moorea (French Polynesia). The benthic community was dominated by macroalgae (85% substratum cover) and comprised of Phaeophyceae Padina tenuis (Bory), Turbinaria ornata (Turner) J. Agardh, and Hydroclathrus clathratus Bory (Howe); Chlorophyta Halimeda incrassata f. ovata J. Agardh (Howe); and Ventricaria ventricosa J. Agardh (Olsen et West), as well as several Rhodophyta (Actinotrichia fragilis Forskál (Børgesen) and several species of encrusting coralline algae). Algal biomass was 171 g dry weight/m**2. Community gross production (Pg), respiration (R), and net calcification (G) were measured in an open-top enclosure. Pg and R were respectively 248 and 240 mmol Co2/m**2/d, and there was a slight net dissolution of CaCO3 (0.8 mmol/m**2/d). This site was a sink for atmospheric CO2 (10 ± 4 mmol CO2/m**2/d), and the analysis of data from the literature suggests that this is a general feature of algal-dominated reefs. Measurement of air-sea CO2 fluxes in open water close to the enclosure demonstrated that changes in small-scale hydrodynamics can lead to misleading conclusions. Net CO2 evasion to the atmosphere was measured on the fringing reef due to changes in the current pattern that drove water from the barrier reef (a C02 source) to the study site.

Seawater carbonate chemistry during a Ishigaki Island (Japan) coral reef seasonal observations, 2005

Monitoring seawater CO2 for a full year with seasonal observations of community metabolism in Ishigaki Island, Japan, revealed seasonal variation and anomalous values owing to the bleaching event in 1998. The daily average pCO2 showed a seasonal pattern on an annual scale, 280 to 320 ?atm in winter and 360 to 400 ?atm in summer, which was determined primarily by the seasonal change in seawater temperature. By contrast, the range in the diel variation in pCO2, 400 to 500 ?atm in summer 200 to 300 ?atm in winter, was attributed to the seasonal variation in community metabolism: Gross primary production (P g ) and respiration (R) were high in summer and low in winter. During the 1998 bleaching event, although P g and R increased, community excess organic production (E) decreased by three quarters compared with the same month in 1999, when the coral community showed high recovery. This change in metabolism led to large diel range and increased average value of pCO2 levels in the seawater on the reef flat. The decrease in the range and increase in the average value of pCO2 were observed by monitoring the Palau barrier reef flat, where overall mortality of corals occurred after the bleaching. All the metabolic parameters, P g , R, E and calcification (G) were reduced by half after the bleaching, which increased the average pCO2 value by 10 ?atm and decreased its diel range from 200-400 ?atm to 100-200 ?atm. Bleaching and resultant mortality of coral reefs led to degradation of their metabolic performance, and thus resulted in the loss of their active interaction with the carbon cycle.

Elemental contents, non-carbonate, fatty acids and alcohols analysis from different Holes of IODP Expedition 310

During IODP Expedition 310 (Tahiti Sea Level), drowned Pleistocene-Holocene barrier-reef terraces were drilled on the slope of the volcanic island. The deglacial reef succession typically consists of a coral framework encrusted by coralline algae and later by microbialites; the latter make up < 80% of the rock volume. Lipid biomarkers were analyzed in order to identify organisms involved in reef-microbialite formation at Tahiti, as the genesis of deglacial microbialites and the conditions favoring their formation are not fully understood. Sterols plus saturated and monounsaturated short-chain fatty acids predominantly derived from both marine primary producers (algae) and bacteria comprise 44 wt% of all lipids on average, whereas long-chain fatty acids and long-chain alcohols derived from higher land plants represent an average of only 24 wt%. Bacterially derived mono-O-alkyl glycerol ethers (MAGEs) and branched fatty acids (10-Me-C16:0; iso- and anteiso-C15:0 and -C17:0) are exceptionally abundant in the microbial carbonates (average, 19 wt%) and represent biomarkers of intermediate-to-high specificity for sulfate-reducing bacteria. Both are relatively enriched in 13C compared to eukaryotic lipids. No lipid biomarkers indicative of cyanobacteria were preserved in the microbialites. The abundances of Al, Si, Fe, Mn, Ba, pyroxene, plagioclase, and magnetite reflect strong terrigenous influx with Tahitian basalt as the major source. Chemical weathering of the basalt most likely elevated nutrient levels in the reefs and this fertilization led to an increase in primary production and organic matter formation, boosting heterotrophic sulfate reduction. Based on the observed biomarker patterns, sulfate-reducing bacteria were apparently involved in the formation of microbialites in the coral reefs off Tahiti during the last deglaciation.

R-codes, and coral abundance, water temperature and transect characteristics of nearshore reefs in Belize

Coral reefs are increasingly threatened by global and local anthropogenic stressors, such as rising seawater temperature and nutrient enrichment. These two stressors vary widely across the reef face and parsing out their influence on coral communities at reef system scales has been particularly challenging. Here, we investigate the influence of temperature and nutrients on coral community traits and life history strategies on lagoonal reefs across the Belize Mesoamerican Barrier Reef System (MBRS). A novel metric was developed using ultra-high-resolution sea surface temperatures (SST) to classify reefs as enduring low (lowTP), moderate (modTP), or extreme (extTP) temperature parameters over 10 years (2003 to 2012). Chlorophyll-a (chl a) records obtained for the same interval were employed as a proxy for bulk nutrients and these records were complemented with in situ measurements to "sea truth" nutrient content across the three reef types. Chl a concentrations were highest at extTP sites, medial at modTP sites and lowest at lowTP sites. Coral species richness, abundance, diversity, density, and percent cover were lower at extTP sites compared to lowTP and modTP sites, but these reef community traits did not differ between lowTP and modTP sites. Coral life history strategy analyses showed that extTP sites were dominated by hardy stress-tolerant and fast-growing weedy coral species, while lowTP and modTP sites consisted of competitive, generalist, weedy, and stress-tolerant coral species. These results suggest that differences in coral community traits and life history strategies between extTP and lowTP/modTP sites were driven primarily by temperature differences with differences in nutrients across site types playing a lesser role. Dominance of weedy and stress-tolerant genera at extTP sites suggests that corals utilizing these two life history strategies may be better suited to cope with warmer oceans and thus may warrant further protective status during this climate change interval. Data associated with this project are archived here, including: -SST data -Satellite Chl a data -Nutrient measurements -Raw coral community survey data For questions contact Justin Baumann (j.baumann3 gmail.com)

Seawater carbonate chemistry and net ecosystem calcification and production in a coral community Kaneohe Bay, Oahu, Hawaii, 2011

Net ecosystem calcification rates (NEC) and net photosynthesis (NP) were determined from CO2 seawater parameters on the barrier coral reef of Kaneohe Bay, Oahu, Hawaii. Autosamplers were deployed to collect samples on the barrier reef every 2 hours for six 48-hour deployments, two each in June 2008, August 2009, and January/February 2010. NEC on the Kaneohe Bay barrier reef increased throughout the day and decreased at night. Net calcification continued at low rates at night except for six time periods when net dissolution was measured. The barrier reef was generally net photosynthetic (positive NP) during the day and net respiring (negative NP) at night. NP controlled the diel cycles of the partial pressure of CO2 (pCO2) and aragonite saturation state resulting in high daytime aragonite saturation state levels when calcification rates were at their peak. However, the NEC and NP diel cycles can become decoupled for short periods of time (several hours) without affecting calcification rates. On a net daily basis, net ecosystem production (NEP) of the barrier reef was found to be sometimes net photosynthetic and sometimes net respiring and ranged from -378 to 80 mmol m-2 d-1 when calculated using simple box models. Daily NEC of the barrier reef was positive (net calcification) for all deployments and ranged from 174 to 331 mmol CaCO3 m-2 d-1. Daily NEC was strongly negatively correlated with average daily pCO2 (R2 = 0.76) which ranged from 431 to 622 µatm. Daily NEC of the Kaneohe Bay barrier reef is similar to or higher than daily NEC measured on other coral reefs even though aragonite saturation state levels (mean aragonite saturation state = 2.85) are some of the lowest measured in coral reef ecosystems. It appears that while calcification rate and ?arag are correlated within a single coral reef ecosystem, this relationship does not necessarily hold between different coral reef systems. It can be expected that ocean acidification will not affect coral reefs uniformly and that some may be more sensitive to increasing pCO2 levels than others.

Composition of bioclastic sands, carbonates and pyroclastic rocks of the Great Meteor and Josephine Seamounts, eastern North Atlantic

1. Great Meteor Seamount (GMS) is a very large (24,000 km**3) guyot with a flat summit plateau at 330-275 m; it has a volcanic core, capped by 150-600 m of post-Middle-Miocene carbonate and pyroclastic rocks, and is covered by bioclastic sands. The much smaller Josephine Seamount (JS, summit 170- 500 m w. d.) consists mainly of basalt which is only locally covered by limestones and bioclastic sands. 2. The bioclastic sands are almost free of terrigenous components, and are well sorted, unimodal medium sands. (1) "Recent pelagic sands" are typical of water depths > 600 m (JS) or > 1000 m (GMS). (2) "Sands of mixed relict-recent origin" (10-40% relict) and (3) "relict sands" (> 40% relict) are highly reworked, coarse lag deposits from the upper flanks and summit tops in which recent constituents are mixed with Pleistocene or older relict material. 3. From the carbonate rocks of both seamounts, 12 "microfacies" (MF-)types were distinguished. The 4 major types are: (1) Bio(pel)sparites (MF 1) occur on the summit plateaus and consist of magnesian calcite cementing small pellets and either redeposited planktonic bioclasts or mixed benthonic-planktonic skeletal debris ; (2) Porous biomicrites (MF 2) are typical of the marginal parts of the summit plateaus and contain mostly planktonic foraminifera (and pteropods), sometimes with redeposited bioclasts and/or coated grains; (3) Dense, ferruginous coralline-algal biomicrudites with Amphistegina sp. (MF 3.1), or with tuffaceous components (MF 3.2); (4) Dense, pelagic foraminiferal nannomicrite (MF 4) with scattered siderite rhombs. Corresponding to the proportion and mineralogical composition of the bioclasts and of the (Mgcalcitic) peloids, micrite, and cement, magnesian calcite (13-17 mol-% MgCO3) is much more abundant than low-Mg calcite and aragonite in rock types (1) and (2). Type (3) contains an "intermediate" Mg-calcite (7-9 mol-X), possibly due to an original Mg deficiency or to partial exsolution of Mg during diagenesis. The nannomicrite (4) consists of low-Mg calcite only. 4. Three textural types of volcanic and associated gyroclastic rocks were distinguished: (1) holohyaline, rapidly chilled and granulated lava flows and tuffs (palagonite tuff breccia and hyaloclastic top breccia); (2) tachylitic basalts (less rapidly chilled; with opaque glass); and (3) "slowly" crystallized, holocrystalline alkali olivine basalts. The carbonate in most mixed pyroclastic-carbonate sediments at the basalt contact is of "post-eruptive" origin (micritic crusts etc.); "pre-eruptive" limestone is recrystallized or altered at the basalt contact. A deuteric (?hydrothermal) "mineralX", filling vesicles in basalt and cementing pyroclastic breccias is described for the first time. 5. Origin and development of GMS andJS: From its origin, some 85 m. y. ago, the volcano of GMS remained active until about 10 m. y. B. P. with an average lava discharge of 320 km**3/m. y. The volcanic origin of JS is much younger (?Middle Tertiary), but the volcanic activity ended also about 9 m. y. ago. During L a t e Miocene to Pliocene times both volcanoes were eroded (wave-rounded cobbles). The oldest pyroclastics and carbonates (MF 3.1, 3.2) were originally deposited in shallow-water (?algal reef hardground). The Plio (-Pleisto) cene foraminiferal nannomicrites (MF 4) suggest a meso- to bathypelagic environment along the flanks of GMS. During the Quaternary (?Pleistocene) bioclastic sands were deposited in water depths beyond wave base on the summit tops, repeatedly reworked, and lithified into loosely consolidated biopelsparites and biomicrites (MF 1 and 2; Fig. 15). Intermediate steps were a first intragranular filling by micrite, reworking, oncoidal coating, weak consolidation with Mg-calcite cemented "peloids" in intergranular voids and local compaction of the peloids into cryptocrystalline micrite with interlocking Mg-calcite crystals up to 4p. The submarine lithification process was frequently interrupted by long intervals of nondeposition, dissolution, boring, and later infilling. The limestones were probably never subaerially exposed. Presently, the carbonate rocks undergo biogenic incrustation and partial dissolution into bioclastic sands. The irregular distribution pattern of the sands reflects (a) the patchy distribution of living benthonic organisms, (b) the steady rain of planktonic organism onto the seamount top, (c) the composition of disintegrating subrecent limestones, and (d) the intensity of winnowing and reworking bottom current

Sediment facies of the Indian-Pakistan continental margin, Indian Ocean Expedition

The studies described here base mainly on sedimentary material collected during the "Indian Ocean Expedition" of the German research vessel "Meteor" in the region of the Indian-Pakistan continental margin in February and March 1965. Moreover,samples from the mouth of the Indus-River were available, which were collected by the Pakistan fishing vessel "Machhera" in March 1965. Altogether, the following quantities of sedimentary material were collected: 59.73 m piston cores. 54.52 m gravity cores. 33 box grab samples. 68 bottom grab samples Component analyses of the coarse fraction were made of these samples and the sedimentary fabric was examined. Moreover, the CaCO3 and Corg contents were discussed. From these investigations the following history of sedimentation can be derived: Recent sedimentation on the shelf is mainly characterized by hydrodynamic processes and terrigenous supply of material. In the shallow water wave action and currents running parallel to the coast, imply a repeated reworking which induces a sorting of the grains and layering of the sediments as well as a lack of bioturbation. The sedimentation rate is very high here. From the coast-line down to appr. 50 m the sediment becomes progressively finer, the conditions of deposition become less turbulent. On the outer shelf the sediment is again considerably coarser. It contains many relicts of planktonic organisms and it shows traces of burrowing. Indications for redeposition are nearly missing, a considerable part of the fine fraction of the sediments is, however, whirled up and carried away. In wide areas of the outer shelf this stirring has gained such a degree that recent deposits are nearly completely missing. Here, coarse relict sands rich in ooids are exposed, which were formed in very shallow stirred water during the time when the sea reached its lowest level, i.e. at the turn of the Pleistocene to the Holocene. Below the relict sand white, very fine-grained aragonite mud was found at one location (core 228). This aragonite mud was obviously deposited in very calm water of some greater depth, possibly behind a reef barrier. Biochemic carbonate precipitation played an important part in the formation of relict sands and aragonite muds. In postglacial times the relict sands were exposed for long periods to violent wave action and to areal erosion. In the present days they are gradually covered by recent sediments proceeding from the sides. On the continental margin beyond the shelf edge the distribution of the sediments is to a considerable extent determined by the morphology of the sea bottom. The material originating from the continent and/or the shelf, is less transported by action of the water than by the force of gravity. Within the range of the uppermost part of the continental slope recent sedimentation reaches its maximum. Here the fine material is deposited which has been whirled up in the zone of the relict sands. A laminated fine-grained sediment is formed here due to the very high sedimentation rate as well as to the extremely low O2-content in the bottom water, which prevents life on the bottom of the sea and impedes thus also bioturbation. The lamination probaly reflects annual variation in deposition and can be attributed to the rhythm of the monsoon with its effects on the water and the weather conditions. In the lower part of the upper continental slope sediments are to be found which show in varying intensity, intercalations of fine material (silt) from the shelf, in large sections of the core. These fine intercalations of allochthonous material are closely related to the autochthonous normal sediment, so that a great number of small individual depositional processes can be inferred. In general the intercalations are missing in the uppermost part of the cores; in the lower part they can be met in different quantities, and they reach their maximum frequency in the upper part of the lower core section. The depositions described here were designated as turbid layer sediments, since they get their material from turbid layers, which transport components to the continental slope which have been whirled up from the shelf. Turbidites are missing in this zone. Since the whole upper continental slope shows a low oxygen-content of the bottom water the structure of the turbid layer sediments is more or less preserved. The lenticular-phacoidal fine structure does, however, not reflect annual rhythms, but sporadic individual events, as e.g. tsunamis. At the lower part of the continental slope and on the continental rise the majority of turbidites was deposited, which, during glacial times and particularly at the beginning of the post-glacial period, transported material from the zone of relict sands. The Laccadive Ridge represented a natural obstacle for the transport of suspended sediments into the deep sea. Core SIC-181 from the Arabian Basin shows some intercalations of turbidites; their material, however, does not originate from the Indian Shelf, but from the Laccadive Ridge. Within the range of the Indus Cone it is surprising that distinct turbidites are nearly completely missing; on the other hand, turbid layer sediments are to be found. The bottom of the sea is showing still a slight slope here, so that the turbidites funneled through the Canyon of the Swatch probably rush down to greater water depths. Due to the particularly large supply of suspended material by theIndus River the turbid layer sediments show farther extension than in other regions. In general the terrigenous components are concentrated on the Indus Cone. It is within the range of the lower continental slope that the only discovery of a sliding mass (core 186) has been located. It can be assumed that this was set in motion during the Holocene. During the period of time discussed here the following development of kind and intensity of the deposition of allochthonous material can be observed on the Indian-Pakistan continental margin: At the time of the lowest sea level the shelf was only very narrow, and the zone in which bottom currents were able to stir up material by oscillating motion, was considerably confined. The rivers flowed into the sea near to the edge of the shelf. For this reason the percentage of terrigenous material, quartz and mica is higher in the lower part of many cores (e.g. cores 210 and 219) than in the upper part. The transition from glacial to postglacial times caused a series of environmental changes. Among them the rise of the sea level (in the area of investigation appr. 150 m) had the most important influence on the sedimentation process. In connection with this event many river valleys became canyons, which sucked sedimentary material away from the shelf and transported it in form of turbidites into the deep sea. During the rise of the sea level a situation can be expected with a maximum area of the comparatively plane shelf being exposed to wave action. During this time the process of stirring up of sediments and formation of turbid layers will reach a maximum. Accordingly, the formation of turbidites and turbid layer sediments are most frequent at the same time. This happened in general in the older polstglacial period. The present day high water level results in a reduced supply of sediments into the canyons. The stirring up of sediments from the shelf by wave action is restricted to the finest material. The missing of shelf material in the uppermost core sections can thus be explained. The laminated muds reflect these calm sedimentation conditions as well. In the southwestern part of the area of investigation fine volcanic glass was blown in during the Pleistocene, probably from the southeast. It has thus become possible to correlate the cores 181, 182, 202. Eolian dust from the Indian subcontinent represents probably an important component of the deep sea sediments. The chemism of the bottom as well as of the pore water has a considerable influence on the development of the sediments. Of particular importance in this connection is a layer with a minimum content of oxygen in the sea water (200-1500 m), which today touches the upper part of the continental slope. Above and beyond this oxygen minimum layer somewhat higher O2-values are to be observed at the sea bottom. During the Pleistocene the oxygen minimum layer has obviously been locatedin greater depth as is indicated by the facies of laminated mud occuring in the lower part of core 219. The type of bioturbation is mainly determined by the chemism. Moreover, the chemism is responsible for a considerable selective dissolution, either complete or partial, of the sedimentary components. Within the range of the oxygen minimum layer an alkaline milieu is developed at the bottom. This causes a complete or partial dissolution of the siliceous organisms. Here, bioturbation is in general completely missing; sometimes small pyrite-filled burrowing racks are found. In the areas rich in O2 high pH-values result in a partial dissolution of the calcareous shells. Large, non-pyritized burrowing tracks characterize the type of bioturbation in this environment. A study of the "lebensspuren" in the cores supports the assumption that, particularly within the region of the Laccadive Basin, the oxygen content in the bottom sediments was lower than during the Holocene. This may be attributed to a high sedimentation rate and to a lower O2-content of the bottom water. The composition of the allochthonous sedimentary components, detritus and/or volcanic glass may locally change the chemism to a considerable extent for a certain time; under such special circumstances the type of bioturbation and the state of preservation of the components may be different from those of the normal sediment.

Changes in coral reef communities across a natural gradient in seawater pH

Ocean acidification threatens the survival of coral reef ecosystems worldwide. The negative effects of ocean acidification observed in many laboratory experiments have been seen in studies of naturally low-pH reefs, with little evidence to date for adaptation. Recently, we reported initial data suggesting that low-pH coral communities of the Palau Rock Islands appear healthy despite the extreme conditions in which they live. Here, we build on that observation with a comprehensive statistical analysis of benthic communities across Palau's natural acidification gradient. Our analysis revealed a shift in coral community composition but no impact of acidification on coral richness, coralline algae abundance, macroalgae cover, coral calcification, or skeletal density. However, coral bioerosion increased 11-fold as pH decreased from the barrier reefs to the Rock Island bays. Indeed, a comparison of the naturally low-pH coral reef systems studied so far revealed increased bioerosion to be the only consistent feature among them, as responses varied across other indices of ecosystem health. Our results imply that whereas community responses may vary, escalation of coral reef bioerosion and acceleration of a shift from net accreting to net eroding reef structures will likely be a global signature of ocean acidification.

Seawater carbonate chemistry and community calcification during Shiraha coral reef (Ishigaki Island, Japan) studies, 1995

Only about half of all the CO_2 that has been produced by the burning of fossil fuels now remains in the atmosphere. The CO_2 "missing" from the atmosphere is the subject of an important debate. It was thought that the great majority of the missing CO_2 has invaded the ocean, for this system naturally acts as a giant chemical regulator of the atmosphere. Although it is clear that ocean processes have a major role in the regulation of the carbon dioxide content of the atmosphere through air-sea exchange processes, recent studies of the oceanic carbon cycle and air-sea interaction indicate that oceanic carbon is in a quasi-steady state via the system of biological and physical processes in the ocean interior. It is difficult to determine whether the ocean has the capacity to take up the increasing air-born CO_2 released by human activities over the past five or six decades. To understand this enigma, we need a better understanding of the natural variability of the oceanic carbon cycle.

Diverse coral communities in naturally acidified waters of a Western Pacific reef

Anthropogenic carbon dioxide emissions are acidifying the oceans, reducing the concentration of carbonate ions ([CO32-) that calcifying organisms need to build and cement coral reefs. To date, studies of a handful of naturally acidified reef systems reveal depauperate communities, sometimes with reduced coral cover and calcification rates, consistent with results of laboratory-based studies. Here we report the existence of highly diverse, coral-dominated reef communities under chronically low pH and aragonite saturation state (Omega ar). Biological and hydrographic processes change the chemistry of the seawater moving across the barrier reefs and into Palau's Rock Island bays, where levels of acidification approach those projected for the western tropical Pacific open ocean by 2100. Nevertheless, coral diversity, cover, and calcification rates are maintained across this natural acidification gradient. Identifying the combination of biological and environmental factors that enable these communities to persist could provide important insights into the future of coral reefs under anthropogenic acidification.