Effects of variations in carbonate chemistry on the calcification rates of Madracis auretenra (= Madracis mirabilis sensu Wells, 1973): bicarbonate concentrations best predict calcification rates

Physiological data and models of coral calcification indicate that corals utilize a combination of seawater bicarbonate and (mainly) respiratory CO₂ for calcification, not seawater carbonate. However, a number of investigators are attributing observed negative effects of experimental seawater acidification by CO₂ or hydrochloric acid additions to a reduction in seawater carbonate ion concentration and thus aragonite saturation state. Thus, there is a discrepancy between the physiological and geochemical views of coral biomineralization. Furthermore, not all calcifying organisms respond negatively to decreased pH or saturation state. Together, these discrepancies suggest that other physiological mechanisms, such as a direct effect of reduced pH on calcium or bicarbonate ion transport and/or variable ability to regulate internal pH, are responsible for the variability in reported experimental effects of acidification on calcification. To distinguish the effects of pH, carbonate concentration and bicarbonate concentration on coral calcification, incubations were performed with the coral Madracis auretenra (= Madracis mirabilis sensu Wells, 1973) in modified seawater chemistries. Carbonate parameters were manipulated to isolate the effects of each parameter more effectively than in previous studies, with a total of six different chemistries. Among treatment differences were highly significant. The corals responded strongly to variation in bicarbonate concentration, but not consistently to carbonate concentration, aragonite saturation state or pH. Corals calcified at normal or elevated rates under low pH (7.6–7.8) when the seawater bicarbonate concentrations were above 1800 μm . Conversely, corals incubated at normal pH had low calcification rates if the bicarbonate concentration was lowered. These results demonstrate that coral responses to ocean acidification are more diverse than currently thought, and question the reliability of using carbonate concentration or aragonite saturation state as the sole predictor of the effects of ocean acidification on coral calcification.     

Regional‐scale dominance of non‐framework building corals on Caribbean reefs affects carbonate production and future reef growth

Coral cover on Caribbean reefs has declined rapidly since the early 1980's. Diseases have been a major driver, decimating communities of framework building Acropora and Orbicella coral species, and reportedly leading to the emergence of novel coral assemblages often dominated by domed and plating species of the genera Agaricia, Porites and Siderastrea. These corals were not historically important Caribbean framework builders, and typically have much smaller stature and lower calcification rates, fuelling concerns over reef carbonate production and growth potential. Using data from 75 reefs from across the Caribbean we quantify: (i) the magnitude of non‐framework building coral dominance throughout the region and (ii) the contribution of these corals to contemporary carbonate production. Our data show that live coral cover averages 18.2% across our sites and coral carbonate production 4.1 kg CaCO₃ m^?2 yr^?1. However, non‐framework building coral species dominate and are major carbonate producers at a high proportion of sites; they are more abundant than Acropora and Orbicella at 73% of sites; contribute an average 68% of the carbonate produced; and produce more than half the carbonate at 79% of sites. Coral cover and carbonate production rate are strongly correlated but, as relative abundance of non‐framework building corals increases, average carbonate production rates decline. Consequently, the use of coral cover as a predictor of carbonate budget status, without species level production rate data, needs to be treated with caution. Our findings provide compelling evidence for the Caribbean‐wide dominance of non‐framework building coral taxa, and that these species are now major regional carbonate producers. However, because these species typically have lower calcification rates, continued transitions to states dominated by non‐framework building coral species will further reduce carbonate production rates below ‘predecline’ levels, resulting in shifts towards negative carbonate budget states and reducing reef growth potential.     

Calcium carbonate budgets for two coral reefs affected by different terrestrial runoff regimes, Rio Bueno, Jamaica

A process-based carbonate budget was used to compare carbonate framework production at two reef sites subject to varying degrees of fluvial influence in Rio Bueno, Jamaica. The turbid, central embayment was subjected to high rates of fluvial sediment input, framework accretion was restricted to ≤30 m, and net carbonate production was 1,887 g CaCO₃ m⁻² year⁻¹. Gross carbonate production (GCP) was dominated by scleractinians (97%), particularly by sediment-resistant species, e.g. Diploria strigosa on the reef flat (<2 m). Calcareous encrusters contributed very little carbonate. Total bioerosion removed 265 g CaCO₃ m⁻² year⁻¹ and was dominated by microborers. At the clear-water site, net carbonate production was 1,236 g CaCO₃ m⁻² year⁻¹; the most productive zone was on the fore-reef (10 m). Corals accounted for 82% of GCP, and encrusting organisms 16%. Bioerosion removed 126 g CaCO₃ m⁻² year⁻¹ and was dominated by macroborers. Total fish and urchin grazing was limited throughout (≤20 g CaCO₃ m⁻² year⁻¹). The study demonstrates that: (1) carbonate production and net reef accretion can occur where environmental conditions approach or exceed perceived threshold levels for coral survival; and (2) although live coral cover (and carbonate production rates) were reduced on reef-front sites along the North Jamaican coast, low population densities of grazing fish and echinoids to some extent offset this, thus maintaining positive carbonate budgets.     

Coral reef calcifiers buffer their response to ocean acidification using both bicarbonate and carbonate

Central to evaluating the effects of ocean acidification (OA) on coral reefs is understanding how calcification is affected by the dissolution of CO₂ in sea water, which causes declines in carbonate ion concentration [CO₃²⁻] and increases in bicarbonate ion concentration [HCO₃⁻]. To address this topic, we manipulated [CO₃²⁻] and [HCO₃⁻] to test the effects on calcification of the coral Porites rus and the alga Hydrolithon onkodes, measured from the start to the end of a 15-day incubation, as well as in the day and night. [CO₃²⁻] played a significant role in light and dark calcification of P. rus, whereas [HCO₃⁻] mainly affected calcification in the light. Both [CO₃²⁻] and [HCO₃⁻] had a significant effect on the calcification of H. onkodes, but the strongest relationship was found with [CO₃²⁻]. Our results show that the negative effect of declining [CO₃²⁻] on the calcification of corals and algae can be partly mitigated by the use of HCO₃⁻ for calcification and perhaps photosynthesis. These results add empirical support to two conceptual models that can form a template for further research to account for the calcification response of corals and crustose coralline algae to OA.     

Horizontal transmission of Symbiodinium cells between adult and juvenile corals is aided by benthic sediment

Of all reef-building coral species, 80–85 % initially draw their intracellular symbionts (dinoflagellates of the genus Symbiodinium) from the environment. Although Symbiodinium cells are crucial for the growth of corals and the formation of coral reefs, little is known about how corals first encounter free-living Symbiodinium cells. We report how the supply of free-living Symbiodinium cells to the benthos by adult corals can increase the rate of horizontal symbiont acquisition for conspecific recruits. Three species of newly settled aposymbiotic (i.e., symbiont-free) corals were maintained in an open aquarium system containing: sterilized sediment and adult coral fragments combined; adult coral fragments alone; sterilized sediment alone; or seawater at Heron Island, Great Barrier Reef, Australia. In all instances, the combination of an adult coral and sediment resulted in the highest symbiont acquisition rates by juvenile corals (up to five-fold greater than seawater alone). Juvenile corals exposed to individual treatments of adult coral or sediment produced an intermediate acquisition response (<52 % of recruits), and symbiont acquisition from unfiltered seawater was comparatively low (<20 % of recruits). Additionally, benthic free-living Symbiodinium cells reached their highest densities in the adult coral + sediment treatment (up to 1.2 × 10⁴ cells mL⁻¹). Our results suggest that corals seed microhabitats with free-living Symbiodinium cells suitable for many coral species during the process of coral recruitment.

     

Calcification by juvenile corals under heterotrophy and elevated CO2

Ocean acidification (OA) threatens the existence of coral reefs by slowing the rate of calcium carbonate (CaCO₃) production of framework-building corals thus reducing the amount of CaCO₃ the reef can produce to counteract natural dissolution. Some evidence exists to suggest that elevated levels of dissolved inorganic nutrients can reduce the impact of OA on coral calcification. Here, we investigated the potential for enhanced energetic status of juvenile corals, achieved via heterotrophic feeding, to modulate the negative impact of OA on calcification. Larvae of the common Atlantic golf ball coral, Favia fragum, were collected and reared for 3 weeks under ambient (421 μatm) or significantly elevated (1,311 μatm) CO₂ conditions. The metamorphosed, zooxanthellate spat were either fed brine shrimp (i.e., received nutrition from photosynthesis plus heterotrophy) or not fed (i.e., primarily autotrophic). Regardless of CO₂ condition, the skeletons of fed corals exhibited accelerated development of septal cycles and were larger than those of unfed corals. At each CO₂ level, fed corals accreted more CaCO₃ than unfed corals, and fed corals reared under 1,311 μatm CO₂ accreted as much CaCO₃ as unfed corals reared under ambient CO₂. However, feeding did not alter the sensitivity of calcification to increased CO₂; ? calcification/?Ω was comparable for fed and unfed corals. Our results suggest that calcification rates of nutritionally replete juvenile corals will decline as OA intensifies over the course of this century. Critically, however, such corals could maintain higher rates of skeletal growth and CaCO₃ production under OA than those in nutritionally limited environments.     

Sponge erosion under acidification and warming scenarios: differential impacts on living and dead coral

Ocean acidification will disproportionately impact the growth of calcifying organisms in coral reef ecosystems. Simultaneously, sponge bioerosion rates have been shown to increase as seawater pH decreases. We conducted a 20‐week experiment that included a 4‐week acclimation period with a high number of replicate tanks and a fully orthogonal design with two levels of temperature (ambient and +1 °C), three levels of pH (8.1, 7.8, and 7.6), and two levels of boring sponge (Cliona varians, present and absent) to account for differences in sponge attachment and carbonate change for both living and dead coral substrate (Porites furcata). Net coral calcification, net dissolution/bioerosion, coral and sponge survival, sponge attachment, and sponge symbiont health were evaluated. Additionally, we used the empirical data from the experiment to develop a stochastic simulation of carbonate change for small coral clusters (i.e., simulated reefs). Our findings suggest differential impacts of temperature, pH and sponge presence for living and dead corals. Net coral calcification (mg CaCO₃ cm^?2 day^?1) was significantly reduced in treatments with increased temperature (+1 °C) and when sponges were present; acidification had no significant effect on coral calcification. Net dissolution of dead coral was primarily driven by pH, regardless of sponge presence or seawater temperature. A reevaluation of the current paradigm of coral carbonate change under future acidification and warming scenarios should include ecologically relevant timescales, species interactions, and community organization to more accurately predict ecosystem‐level response to future conditions.     

Coral calcification responds to seawater acidification: a working hypothesis towards a physiological mechanism

The decrease in the saturation state of seawater, Ω, following seawater acidification, is believed to be the main factor leading to a decrease in the calcification of marine organisms. To provide a physiological explanation for this phenomenon, the effect of seawater acidification was studied on the calcification and photosynthesis of the scleractinian tropical coral Stylophora pistillata. Coral nubbins were incubated for 8 days at three different pH (7.6, 8.0, and 8.2). To differentiate between the effects of the various components of the carbonate chemistry (pH, CO₃²⁻, HCO₃⁻, CO₂, Ω), tanks were also maintained under similar pH, but with 2-mM HCO₃⁻ added to the seawater. The addition of 2-mM bicarbonate significantly increased the photosynthesis in S. pistillata, suggesting carbon-limited conditions. Conversely, photosynthesis was insensitive to changes in pH and pCO₂. Seawater acidification decreased coral calcification by ca. 0.1-mg CaCO₃ g⁻¹ d⁻¹ for a decrease of 0.1 pH units. This correlation suggested that seawater acidification affected coral calcification by decreasing the availability of the CO₃²⁻ substrate for calcification. However, the decrease in coral calcification could also be attributed either to a decrease in extra- or intracellular pH or to a change in the buffering capacity of the medium, impairing supply of CO₃²⁻ from HCO₃⁻.     

Neritic carbonate for six submerged coral reefs from northern Australia: Implications for Holocene global carbon dioxide

Total contribution of six recently discovered submerged coral reefs in northern Australia to Holocene neritic CaCO₃ and CO₂ is assessed to address a gap in global budgets. CaCO₃ production for the reef framework and inter-reefal deposits is 0.26–0.28 Mt. Holocene CO₂ production is 0.14–0.16 Mt. Coral and coralline algae are the dominant sources of Holocene CaCO₃ although foraminifers and molluscs are the dominant constituents of inter-reefal deposits. The total amount of Holocene neritic CaCO₃ produced by the six submerged coral reefs is several orders of magnitude smaller than that calculated using accepted CaCO₃ production values because of very low production, a ‘give-up’ growth history, and presumed significant dissolution and exports. The contribution of submerged coral reefs to global Holocene neritic CaCO₃ is estimated to be 0.26–0.62 Gt, which yields 0.15–0.37 Gt of CO₂. This amount of CO₂ is 0.02–0.05% of the 780 Gt added to the atmosphere since 18 kyr BP. Contributions from Australian submerged coral reefs are estimated to be 0.05 Gt CaCO₃ and 0.03 Gt CO₂ for an emergent reef area of 47.9 × 10³ km². Based on the growth history of the submerged coral reefs in the Gulf of Carpentaria, maximum global Holocene CaCO₃ fluxes could have attained 0.3 Gt yr^− 1 between 11 and 7 ka BP. This additional CaCO₃ would have culminated in a maximum CaCO₃ production from all (emergent and submerged) coral reefs of 1.2 Gt yr^− 1 and neritic CaCO₃ production of 2.75 Gt yr^− 1. The dilemma remains that the global area and CaCO₃ mass of submerged coral reefs are currently unknown. It is inevitable that many more submerged coral reefs will be found. Our findings imply that submerged coral reefs are a small but fundamental source of Holocene neritic CaCO₃ and CO₂.     

Effect of zooplankton availability on the rates of photosynthesis, and tissue and skeletal growth in the scleractinian coral Stylophora pistillata

This work investigated the effect of light and feeding on tissue composition as well as on rates of photosynthesis and calcification in the zooxanthellae (zoox) scleractinian coral, Stylophora pistillata. Microcolonies were maintained at three different light levels (80, 200, 300 μmol m⁻² s⁻¹) and subjected to two feeding regimes (starved and fed) over 9 weeks. Corals were fed both natural plankton and Artemia salina nauplii four times a weeks and samplings were made after 2, 5, and 9 weeks. Results confirmed that feeding enhances coral growth rate and increases both the dark and light calcification rates. These rates were 50-75% higher in fed corals (FC; 60±20 and 200±40 nmol Ca²⁺ cm⁻² h⁻¹ for dark and light calcification, respectively) compared to control corals (CC; 30±9 and 124±23 nmol Ca²⁺ cm⁻² h⁻¹). The dark calcification rates, however, were four times lower than the rates of light calcification (independent of trophic status). After 5 weeks, chlorophyll a (chl-a) concentrations were four to seven times higher in fed corals (7-21 μg cm⁻²) than in control corals (2-5 μg cm⁻²). The amount of protein was also significantly higher in fed corals (2.11-2.50 mg cm⁻²) than in control corals (1.08-1.52 mg cm⁻²). Rates of photosynthesis in fed corals were 2-10 times higher (1.24±0.75 μmol O₂ h⁻¹ cm⁻²) than those measured in control corals (0.20±0.08 μmol O₂ h⁻¹ cm⁻²).     

The impact of seawater saturation state and bicarbonate ion concentration on calcification by new recruits of two Atlantic corals

Rising concentrations of atmospheric CO₂ are changing the carbonate chemistry of the oceans, a process known as ocean acidification (OA). Absorption of this CO₂ by the surface oceans is increasing the amount of total dissolved inorganic carbon (DIC) and bicarbonate ion (HCO₃ ⁻) available for marine calcification yet is simultaneously lowering the seawater pH and carbonate ion concentration ([CO₃ ²⁻]), and thus the saturation state of seawater with respect to aragonite (Ω_ar). We investigated the relative importance of [HCO₃ ⁻] versus [CO₃ ²⁻] for early calcification by new recruits (primary polyps settled from zooxanthellate larvae) of two tropical coral species, Favia fragum and Porites astreoides. The polyps were reared over a range of Ω_ar values, which were manipulated by both acid-addition at constant pCO₂ (decreased total [HCO₃ ⁻] and [CO₃ ²⁻]) and by pCO₂ elevation at constant alkalinity (increased [HCO₃ ⁻], decreased [CO₃ ²⁻]). Calcification after 2 weeks was quantified by weighing the complete skeleton (corallite) accreted by each polyp over the course of the experiment. Both species exhibited the same negative response to decreasing [CO₃ ²⁻] whether Ω_ar was lowered by acid-addition or by pCO₂ elevation—calcification did not follow total DIC or [HCO₃ ⁻]. Nevertheless, the calcification response to decreasing [CO₃ ²⁻] was nonlinear. A statistically significant decrease in calcification was only detected between Ω_ar = <2.5 and Ω_ar = 1.1–1.5, where calcification of new recruits was reduced by 22–37% per 1.0 decrease in Ω_ar. Our results differ from many previous studies that report a linear coral calcification response to OA, and from those showing that calcification increases with increasing [HCO₃ ⁻]. Clearly, the coral calcification response to OA is variable and complex. A deeper understanding of the biomineralization mechanisms and environmental conditions underlying these variable responses is needed to support informed predictions about future OA impacts on corals and coral reefs.     

Ocean Acidification Accelerates Reef Bioerosion

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

Calcification in bleached and unbleached Montastraea faveolata: evaluating the role of oxygen and glycerol

All reef-building corals are symbiotic with dinoflagellates of the genus Symbiodinium, which influences many aspects of the host’s physiology including calcification. Coral calcification is a biologically controlled process performed by the host that takes place several membranes away from the site of photosynthesis performed by the symbiont. Although it is well established that light accelerates CaCO₃ deposition in reef-building corals (commonly referred to as light-enhanced calcification), the complete physiological mechanism behind the process is not fully understood. To better comprehend the coral calcification process, a series of laboratory experiments were conducted in the major Caribbean reef-building species Montastraea faveolata, to evaluate the effect of glycerol addition and/or the super-saturation of oxygen in the seawater. These manipulations were performed in bleached and unbleached corals, to separate the effect of photosynthesis from calcification. The results suggest that under normal physiological conditions, a 42% increase in seawater oxygen concentration promotes a twofold increase in dark-calcification rates relative to controls. On the other hand, the results obtained using bleached corals suggest that glycerol is required, as a metabolic fuel, in addition to an oxygenic environment in a symbiosis that has been disrupted. Also, respiration rates in symbiotic corals that were pre-incubated in light conditions showed a kinetic limitation, whereas corals that were pre-incubated in darkness were oxygen limited, clearly emphasizing the role of oxygen in this regard. These findings indicate that calcification in symbiotic corals is not strictly a “light-enhanced” or “dark-repressed” process, but rather, the products of photosynthesis have a critical role in calcification, which should be viewed as a “photosynthesis-driven” process. The results presented here are discussed in the context of the current knowledge of the coral calcification process.     

Net release of dissolved organic matter by the scleractinian coral Acropora pulchra

The net production of dissolved organic matter (DOM) and dissolved combined and free amino acids (DCAA and DFAA, respectively) by the hermatypic coral Acropora pulchra was measured in the submerged condition, and the production rates were normalized to the coral surface area, tissue biomass, and net photosynthetic rates by zooxanthellae. When normalized to the unit surface area, the production rates of dissolved organic carbon and nitrogen (DOC and DON, respectively) were 37 and 4.4 nmol cm^− 2 h^− 1, respectively. Comparing with the photosynthetic rate by zooxanthellae, which was measured by ¹³C-tracer accumulation in the soft tissue of the coral colony, the release rate of DOC corresponded to 5.4% of the daily net photosynthetic production. The tissue biomass of the coral colony was 178 µmol C cm^− 2 and 23 µmol N cm^− 2, indicating that the release of DOC and DON accounted for 0.021% h^− 1 and 0.019% h^− 1 of the tissue C and N, respectively. The C:N ratios of the released DOM (average 8.4) were not significantly different from those of the soft tissue of the coral colonies (average 7.7). While DFAA did almost not accumulate in the incubated seawater, DCAA was considerably released by the coral colonies at the rate of 2.1 nmol cm^− 2 h^− 1 on average. Calculating C and N contents of the hydrolyzable DCAA, it was revealed that about 20% and 50%–60% of the released bulk DOC and DON, respectively, were composed of DCAA.     

Coral growth with thermal stress and ocean acidification: lessons from the eastern tropical Pacific

The rapid growth of scleractinian corals is responsible for the persistence of coral reefs through time. Coral growth rates have declined over the past 30 years in the western Pacific, Indian, and North Atlantic Oceans. The spatial scale of this decline has led researchers to suggest that a global phenomenon like ocean acidification may be responsible. A multi-species inventory of coral growth from Pacific Panamá confirms that declines have occurred in some, but not all species. Linear extension declined significantly in the most important reef builder of the eastern tropical Pacific, Pocillopora damicornis, by nearly one-third from 1974 to 2006. The rate of decline in skeletal extension for P. damicornis from Pacific Panamá (0.9% year⁻¹) was nearly identical to massive Porites in the Indo-Pacific over the past 20–30 years (0.89–1.23% year⁻¹). The branching pocilloporid corals have shown an increased tolerance to recurrent thermal stress events in Panamá, but appear to be susceptible to acidification. In contrast, the massive pavonid corals have shown less tolerance to thermal stress, but may be less sensitive to acidification. These differing sensitivities will be a fundamental determinant of eastern tropical Pacific coral reef community structure with accelerating climate change that has implications for the future of reef communities worldwide.     

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.     

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.     

BICARBONATE STIMULATION OF CALCIFICATION AND PHOTOSYNTHESIS IN TWO HERMATYPIC CORALS1

A wide range of bicarbonate concentrations was used to monitor the kinetics of bicarbonate (HCO₃^?) use in both photosynthesis and calcification in two reef‐building corals, Porites porites and Acropora sp. Experiments carried out close to the P. porites collection site in Barbados showed that additions of NaHCO₃ to synthetic seawater proportionally increased the calcification rate of this coral until the concentration exceeded three times that of seawater (6 mM). Photosynthetic rates were also stimulated by HCO₃^? addition, but these became saturated at a lower concentration (4 mM). Similar experiments on aquarium‐acclimated colonies of Indo‐Pacific Acropora sp. showed that calcification and photosynthesis in this coral were enhanced to an even greater extent than P. porites, with calcification continuing to increase above 8 mM HCO₃^?, and photosynthesis saturating at 6 mM. Calcification rates of Acropora sp. were also monitored in the dark, and, although these were lower than in the light for a given HCO₃^? concentration, they still increased dramatically with HCO₃^? addition, showing that calcification in this coral is light stimulated but not light dependent.     

Ocean acidification effects on calcification and dissolution in tropical reef macroalgae

Net calcification rates for coral reef and other calcifiers have been shown to decline as ocean acidification (OA) occurs. However, the role of calcium carbonate dissolution in lowering net calcification rates is unclear. The objective of this study was to distinguish OA effects on calcification and dissolution rates in dominant calcifying macroalgae of the Florida Reef Tract, including two rhodophytes (Neogoniolithon strictum, Jania adhaerens) and two chlorophytes (Halimeda scabra, Udotea luna). Two experiments were conducted: (1) to assess the difference in gross (⁴⁵Ca uptake) versus net (total alkalinity anomaly) calcification rates in the light/dark and (2) to determine dark dissolution (⁴⁵CaCO₃), using pH levels predicted for the year 2100 and ambient pH. At low pH in the light, all species maintained gross calcification rates and most sustained net calcification rates relative to controls. Net calcification rates in the dark were ~84% lower than in the light. In contrast to the light, all species had lower net calcification rates in the dark at low pH with chlorophytes exhibiting net dissolution. These data are supported by the relationship (R² = 0.82) between increasing total alkalinity and loss of ⁴⁵Ca from pre-labelled ⁴⁵CaCO₃ thalli at low pH in the dark. Dark dissolution of ⁴⁵CaCO₃-labelled thalli was ~18% higher in chlorophytes than rhodophytes at ambient pH, and ~ twofold higher at low pH. Only Udotea, which exhibited dissolution in the light, also had lower daily calcification rates integrated over 24 h. Thus, if tropical macroalgae can maintain high calcification rates in the light, lower net calcification rates in the dark from dissolution may not compromise daily calcification rates. However, if organismal dissolution in the dark is additive to sedimentary carbonate losses, reef dissolution may be amplified under OA and contribute to erosion of the Florida Reef Tract and other reefs that exhibit net dissolution.

     

Space partitioning by stony corals soft corals and benthic algae on the coral reefs of the northern Gulf of Eilat (Red Sea)

The major faunistic and floristic components occupying space on the coral reefs of the northern Gulf of Eilat (Red Sea) are stony corals, soft corals and benthic algae. The percent living coverage of the three components and the relative abundance of the different species of each component were studied by line transects, on the reef flats and the upper forereef zones of nine localities. A wider and higher range of living coverage values of stony corals were recorded at the upper fore-reef zones (18.30–49.09%) compared with the reef flats (5.50–31.66%) at the different stations. The most abundant stony corals on the reef flats areCyphastrea microphthalma, Stylophora pistillata, Favia favus, Porites lutea, Platygyra lamellina and the hydrozoanMillepora dichotoma. The fire coralM. dichotoma dominates the upper fore-reef zone in most of the stations. The average percent living coverage of soft corals on the reef flats ranged between 0.20 and 17.06%, and on the upper fore-reef zones between 1.68 and 15.13%. Seventy percent of the total living coverage of the soft coral community is contributed by 2 to 3 species. They tend to form large monospecific carpets, such as those composed ofSinularia sp.,Sarcophyton glaucum andLobophytum pauciflorum. The common benthic algae on the coral reef studied occur as turfs or macroscopic noncalcareous algae. They play a significant role in occupying space, especially on the reef flats. The most abundant algae recorded in all localities are the turfsSphacelaria tribuloides, Jania sp. and the macroscopic non-calcareous algaeTurbinaria elatensis andColpomenia sinuosa. Comparison between reef flats and upper fore-reef zones, in terms of average living cover of stony corals, shows that the variation among the reef flats is grater than the variation among the upper fore-reef zones. However, there is no significant variation in the average living coverage of soft corals between these two zones. Annual living-coverage values of algae on the reef flats are significantly higher than those of the upper fore-reef zones. Extremely low tides occurring periodically but unpredictably at Eilat cause mass mortality of the benthic communities on the reef flats reopening new spaces for settlement. The coexistence of stony corals, soft corals and algae on the reef ecosystem is due to different biological properties of each component. Opportunistic life histories of certain stony corals and most algae enable quick colonization of newly opened spaces. Lack of predators, high tolerance against abiotic factors and ability to form large aggregates of colonies are suggested as possible factors supporting the existence of soft corals in shallow water. Biological factors such as competition, predation and grazing pressure play an increasingly important role in controlling space utilization by the components studied with the advancement of succession.