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.     

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

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

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.     

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.     

Coral reefs modify their seawater carbon chemistry – implications for impacts of ocean acidification

Reviews suggest that that the biogeochemical threshold for sustained coral reef growth will be reached during this century due to ocean acidification caused by increased uptake of atmospheric CO₂. Projections of ocean acidification, however, are based on air‐sea fluxes in the open ocean, and not for shallow‐water systems such as coral reefs. Like the open ocean, reef waters are subject to the chemical forcing of increasing atmospheric pCO₂. However, for reefs with long water residence times, we illustrate that benthic carbon fluxes can drive spatial variation in pH, pCO₂ and aragonite saturation state (Ω_a) that can mask the effects of ocean acidification in some downstream habitats. We use a carbon flux model for photosynthesis, respiration, calcification and dissolution coupled with Lagrangian transport to examine how key groups of calcifiers (zooxanthellate corals) and primary producers (macroalgae) on coral reefs contribute to changes in the seawater carbonate system as a function of water residence time. Analyses based on flume data showed that the carbon fluxes of corals and macroalgae drive Ω_ain opposing directions. Areas dominated by corals elevate pCO₂ and reduce Ω_a, thereby compounding ocean acidification effects in downstream habitats, whereas algal beds draw CO₂ down and elevate Ω_a, potentially offsetting ocean acidification impacts at the local scale. Simulations for two CO₂ scenarios (600 and 900 ppm CO₂) suggested that a potential shift from coral to algal abundance under ocean acidification can lead to improved conditions for calcification in downstream habitats, depending on reef size, water residence time and circulation patterns. Although the carbon fluxes of benthic reef communities cannot significantly counter changes in carbon chemistry at the scale of oceans, they provide a significant mechanism of buffering ocean acidification impacts at the scale of habitat to reef.     

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.     

Ocean acidification and calcifying reef organisms: a mesocosm investigation

A long-term (10 months) controlled experiment was conducted to test the impact of increased partial pressure of carbon dioxide (pCO₂) on common calcifying coral reef organisms. The experiment was conducted in replicate continuous flow coral reef mesocosms flushed with unfiltered sea water from Kaneohe Bay, Oahu, Hawaii. Mesocosms were located in full sunlight and experienced diurnal and seasonal fluctuations in temperature and sea water chemistry characteristic of the adjacent reef flat. Treatment mesocosms were manipulated to simulate an increase in pCO₂ to levels expected in this century [midday pCO₂ levels exceeding control mesocosms by 365 ± 130 μatm (mean ± sd)]. Acidification had a profound impact on the development and growth of crustose coralline algae (CCA) populations. During the experiment, CCA developed 25% cover in the control mesocosms and only 4% in the acidified mesocosms, representing an 86% relative reduction. Free-living associations of CCA known as rhodoliths living in the control mesocosms grew at a rate of 0.6 g buoyant weight year⁻¹ while those in the acidified experimental treatment decreased in weight at a rate of 0.9 g buoyant weight year⁻¹, representing a 250% difference. CCA play an important role in the growth and stabilization of carbonate reefs, so future changes of this magnitude could greatly impact coral reefs throughout the world. Coral calcification decreased between 15% and 20% under acidified conditions. Linear extension decreased by 14% under acidified conditions in one experiment. Larvae of the coral Pocillopora damicornis were able to recruit under the acidified conditions. In addition, there was no significant difference in production of gametes by the coral Montipora capitata after 6 months of exposure to the treatments.     

Budget of Primary Production and Dinitrogen Fixation in a Highly Seasonal Red Sea Coral Reef

Biological dinitrogen (N₂) fixation (diazotrophy, BNF) relieves marine primary producers of nitrogen (N) limitation in a large part of the world oceans. N concentrations are particularly low in tropical regions where coral reefs are located, and N is therefore a key limiting nutrient for these productive ecosystems. In this context, the importance of diazotrophy for reef productivity is still not resolved, with studies up to now lacking organismal and seasonal resolution. Here, we present a budget of gross primary production (GPP) and BNF for a highly seasonal Red Sea fringing reef, based on ecophysiological and benthic cover measurements combined with geospatial analyses. Benthic GPP varied from 215 to 262 mmol C m^?2 reef d^?1, with hard corals making the largest contribution (41–76%). Diazotrophy was omnipresent in space and time, and benthic BNF varied from 0.16 to 0.92 mmol N m^?2 reef d^?1. Planktonic GPP and BNF rates were respectively approximately 60- and 20-fold lower than those of the benthos, emphasizing the importance of the benthic compartment in reef biogeochemical cycling. BNF showed higher sensitivity to seasonality than GPP, implying greater climatic control on reef BNF. Up to about 20% of net reef primary production could be supported by BNF during summer, suggesting a strong biogeochemical coupling between diazotrophy and the reef carbon cycle.     

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.     

Stock Structure, Mortality and Growth of The Decorated Goby, Istigobius decoratus (Gobiidae), at Lizard Island, Great Barrier Reef

Gobiids are an abundant component of coral reef ichthyofauna, yet little is known of their life histories. I examined population structure, mortality and growth of the decorated goby, Istigobius decoratus, a common gobiid of shallow patch reefs on the Great Barrier Reef. Presumed daily increments in sagittal otoliths were used as a proxy for age. The upper age estimate was 266 days suggesting at most an annual life cycle. Instantaneous natural mortality rate estimates were 5.92 year^–1 and 7.92 year^–1 using two estimators, both corresponding to less than 1% annual survivorship. Specimens ranged from 12 to 84 mm total length. Analysis of size-at-age data indicated linear growth at a rate of 0.33 mm day^–1. The linear relationship between size and age meant the population size structure mirrored the age structure with both skewed toward the smallest and youngest classes. High mortality over a 1-year longevity and linear growth suggest high population turnover and, therefore, that I. decoratus and ecologically similar species serve a potentially important role as prey species. This suite of traits is rarely reported for coral reef fishes, which is probably due to the limited attention paid to small-bodied species rather than the rarity of such a life history in these communities.     

Larvae of the coral eating crown‐of‐thorns starfish,Acanthaster planci in a warmer‐high CO2 ocean

Outbreaks of crown‐of‐thorns starfish (COTS), Acanthaster planci, contribute to major declines of coral reef ecosystems throughout the Indo‐Pacific. As the oceans warm and decrease in pH due to increased anthropogenic CO₂ production_, coral reefs are also susceptible to bleaching, disease and reduced calcification. The impacts of ocean acidification and warming may be exacerbated by COTS predation, but it is not known how this major predator will fare in a changing ocean. Because larval success is a key driver of population outbreaks, we investigated the sensitivities of larval A. planci to increased temperature (2–4 °C above ambient) and acidification (0.3–0.5 pH units below ambient) in flow‐through cross‐factorial experiments (3 temperature × 3 pH/pCO₂ levels). There was no effect of increased temperature or acidification on fertilization or very early development. Larvae reared in the optimal temperature (28 °C) were the largest across all pH treatments. Development to advanced larva was negatively affected by the high temperature treatment (30 °C) and by both experimental pH levels (pH 7.6, 7.8). Thus, planktonic life stages of A. planci may be negatively impacted by near‐future global change. Increased temperature and reduced pH had an additive negative effect on reducing larval size. The 30 °C treatment exceeded larval tolerance regardless of pH. As 30 °C sea surface temperatures may become the norm in low latitude tropical regions, poleward migration of A. planci may be expected as they follow optimal isotherms. In the absence of acclimation or adaptation, declines in low latitude populations may occur. Poleward migration will be facilitated by strong western boundary currents, with possible negative flow‐on effects on high latitude coral reefs. The contrasting responses of the larvae of A. planci and those of its coral prey to ocean acidification and warming are considered in context with potential future change in tropical reef ecosystems.     

Acclimation to ocean acidification during long‐term CO2 exposure in the cold‐water coral Lophelia pertusa

Ocean acidity has increased by 30% since preindustrial times due to the uptake of anthropogenic CO₂ and is projected to rise by another 120% before 2100 if CO₂ emissions continue at current rates. Ocean acidification is expected to have wide‐ranging impacts on marine life, including reduced growth and net erosion of coral reefs. Our present understanding of the impacts of ocean acidification on marine life, however, relies heavily on results from short‐term CO₂ perturbation studies. Here, we present results from the first long‐term CO₂ perturbation study on the dominant reef‐building cold‐water coral Lophelia pertusa and relate them to results from a short‐term study to compare the effect of exposure time on the coral's responses. Short‐term (1 week) high CO₂ exposure resulted in a decline of calcification by 26–29% for a pH decrease of 0.1 units and net dissolution of calcium carbonate. In contrast, L. pertusa was capable to acclimate to acidified conditions in long‐term (6 months) incubations, leading to even slightly enhanced rates of calcification. Net growth is sustained even in waters sub‐saturated with respect to aragonite. Acclimation to seawater acidification did not cause a measurable increase in metabolic rates. This is the first evidence of successful acclimation in a coral species to ocean acidification, emphasizing the general need for long‐term incubations in ocean acidification research. To conclude on the sensitivity of cold‐water coral reefs to future ocean acidification further ecophysiological studies are necessary which should also encompass the role of food availability and rising temperatures.     

Diel pCO2 variation among coral reefs and microhabitats at Lizard Island,Great Barrier Reef

Most laboratory experiments examining the effect of ocean acidification on marine organisms use stable pH/pCO₂ treatments based on average projections for the open ocean. However, pH/pCO₂ levels vary spatially and temporally in marine environments, and this variation can affect organism responses to pH/pCO₂. On coral reefs, diel pH/pCO₂ variability at the individual reef scale has been reported in a few studies, but variation among microhabitats within a reef remains poorly understood. This study determined the pH/pCO₂ variability of three different reefs, and three contrasting coral reef microhabitats (dominated by hard coral, soft coral, or open substrate) within each reef. Three SeaFET pH loggers were deployed simultaneously at the three microhabitats within a reef over a 9-day period. This was repeated at three different reefs around the Lizard Island lagoon. The loggers recorded pH_T and temperature every 5 min. Water samples were collected from each microhabitat during four points of the tidal cycle (high, low, rising, and falling) and analysed for total alkalinity and dissolved inorganic carbon. The data show a clear diel pCO₂ cycle, increasing overnight and decreasing during the day, in association with photosynthesis and respiration cycles. Diel pCO₂ differed more between reefs than between microhabitats within reefs. Variation between reefs was most likely influenced by water flow, with the more protected (low flow) reefs experiencing a greater range in pCO₂ (Δ 250 μatm) than the exposed (high flow) reefs (Δ 116 μatm). These results add to a growing body of the literature on the diel variation of pCO₂ of shallow, nearshore environments and suggest that when projecting future pCO₂ levels, it is important to consider reef metabolism as well as physical and hydrodynamic factors.

     

Reduced carbon emissions and fishing pressure are both necessary for equatorial coral reefs to keep up with rising seas

A rapid increase in sea-level rise is generating vertical accommodation space on modern coral reefs. Yet increases in sea-surface temperatures (SSTs) are reducing the capacity of coral reefs to keep up with sea-level rise. We use ensemble species distribution models of four coral species (Porites rus, Porites lobata, Acropora hyacinthus and Acropora digitifera) to gauge potential geographic differences in gross carbonate production. Net carbonate production was estimated by considering erosional rates of ocean acidification, increasing cyclone intensity, local pollution, fishing pressure and the projected burdens of increases in SSTs (under Representative Concentration Pathways (RCPs) 4.5, 6.0 and 8.5) through to the year 2100. Our models predict that only 4 ± 0.1% (~60 000 km²) of Indo-Pacific coral reefs are projected to keep up with sea-level rise by the year 2100 under RCP 8.5 – most of which will be located near the Equator. However, with drastic reductions in emissions (under RCPs 4.5 and 6.0 Wm⁻²), we predict that 15 ± 0.3% (~250 000 km²) (under RCP 4.5 Wm⁻²) and 12 ± 0.7% (~200 000 km²) (under RCP 6.0 Wm⁻²) of Indo-Pacific coral reefs, have the potential to keep up with sea-level rise by the year 2100. Yet the burdens of fishing pressure and its cascading effects are projected to be responsible for substantial reef erosion, nearly halving the number of reefs able to keep up with sea-level rise. If action is taken immediately and emissions are drastically reduced to RCPs 4.5 or 6.0 Wm⁻², and reef management reduces the burdens of local pollution and fishing pressure, then our model predicts that 21–27% (~350 000–470 000 km²) of Indo-Pacific coral reefs – most of which will be located near the Equator – would have the potential to keep up with sea-level rise by the year 2100.     

Phosphorus metabolism in coral reef communities: exchange between the water column and bottom biotopes

Exchange of phosphate between components of the reef bottom and the water column were studied on reefs around Heron Island (Great Barrier Reef), both in aquaria and in in situ enclosures, using radioactive phosphorus (³²P) as a tracer. Living corals, dead corals, coral rubble overgrown with periphyton, and soft sediments of coral sand were used in experiments. In all of these components of bottom reef biotopes, two opposite flows of inorganic phosphate were recorded and measured, i.e. the rate of PO₄-P uptake from water (A_c), and its release (A_e). At ambient PO₄-P concentrations in water of 0.1– 0.3 µmoll^–1, both flows varied in living corals and coral rubble between 10 and 70 µg P kg^–1 h^–1, 3–10 mg P m^–2 day^–1, and in coral sand between 10 and 30 µg P kg^–1 h^–1, or 2–7 mg P m^–2 day^–1. Under the latter concentration range (which is typical for coral reef areas), the reciprocal PO₄-P flows almost balanced each other, so that net uptake (A_t) was very low. Often it approached zero or was positive, showing that a net PO₄-P release had taken place. The uptake flow (A_c) in living coral was much more dependent on the PO₄-P content in overlying water than was the release flow (A_e). The influence of conditions of illumination upon the values of A_c and A_e was comparatively low. The data obtained are used to discuss problems of phosphorus balance and dynamics in coral reef ecosystems.     

Speciation in coral-reef fishes

Covering <0·1% of the ocean’s surface, coral reefs harbour about one‐third of all marine fishes or c. 5000 species. Allopatry (geographic isolation) is believed to be the primary mode of speciation, yet few biogeographic barriers exist between reefs, and most reef fishes have a pelagic larval stage capable of extensive dispersal. Under these circumstances, why are there so many species of reef fishes? Since most biogeographic barriers in the oceans are either spatially or temporally permeable on a relatively short time frame, the requirement of isolation during allopatric speciation is hard to satisfy. Evidence from empirical and theoretical studies, the biological characteristics of coral reefs, and a reanalysis of biogeographic barriers indicate that sympatric speciation is possible but not common at small spatial scales and that parapatric speciation is a common (and probably the prevalent) mode of diversification in coral‐reef fishes. Regardless of the speciation mode, previous hypotheses of accelerated diversification in the Pleistocene due to sea level fluctuations are not supported by phylogenetic analyses. Recent developments in the area of comparative genomics can fuel a new revolution in the way marine speciation is studied.     

Carbon storage and fluxes in ponderosa pine forests at different developmental stages

We compared carbon storage and fluxes in young and old ponderosa pine stands in Oregon, including plant and soil storage, net primary productivity, respiration fluxes, eddy flux estimates of net ecosystem exchange (NEE), and Biome‐BGC simulations of fluxes. The young forest (Y site) was previously an old‐growth ponderosa pine forest that had been clearcut in 1978, and the old forest (O site), which has never been logged, consists of two primary age classes (50 and 250 years old). Total ecosystem carbon content (vegetation, detritus and soil) of the O forest was about twice that of the Y site (21 vs. 10 kg C m^?2 ground), and significantly more of the total is stored in living vegetation at the O site (61% vs. 15%). Ecosystem respiration (R_e) was higher at the O site (1014 vs. 835 g C m^?2 year^?1), and it was largely from soils at both sites (77% of R_e). The biological data show that above‐ground net primary productivity (ANPP), NPP and net ecosystem production (NEP) were greater at the O site than the Y site. Monte Carlo estimates of NEP show that the young site is a source of CO₂ to the atmosphere, and is significantly lower than NEP(O) by c. 100 g C m^?2 year^?1. Eddy covariance measurements also show that the O site was a stronger sink for CO₂ than the Y site. Across a 15‐km swath in the region, ANPP ranged from 76 g C m^?2 year^?1 at the Y site to 236 g C m^?2 year^?1 (overall mean 158 ± 14 g C m^?2 year^?1). The lowest ANPP values were for the youngest and oldest stands, but there was a large range of ANPP for mature stands. Carbon, water and nitrogen cycle simulations with the Biome‐BGC model suggest that disturbance type and frequency, time since disturbance, age‐dependent changes in below‐ground allocation, and increasing atmospheric concentration of CO₂ all exert significant control on the net ecosystem exchange of carbon at the two sites. Model estimates of major carbon flux components agree with budget‐based observations to within ± 20%, with larger differences for NEP and for several storage terms. Simulations showed the period of regrowth required to replace carbon lost during and after a stand‐replacing fire (O) or a clearcut (Y) to be between 50 and 100 years. In both cases, simulations showed a shift from net carbon source to net sink (on an annual basis) 10–20 years after disturbance. These results suggest that the net ecosystem production of young stands may be low because heterotrophic respiration, particularly from soils, is higher than the NPP of the regrowth. The amount of carbon stored in long‐term pools (biomass and soils) in addition to short‐term fluxes has important implications for management of forests in the Pacific North‐west for carbon sequestration.     

Distribution and sediment production of large benthic foraminifers on reef flats of the Majuro Atoll,Marshall Islands

The distributions and population densities of large benthic foraminifers (LBFs) were investigated on reef flats of the Majuro Atoll, Marshall Islands. Annual sediment production by foraminifers was estimated based on population density data. Predominant LBFs were Calcarina and Amphistegina, and the population densities of these foraminifers varied with location and substratum type on reef flats. Both foraminifers primarily attached to macrophytes, particularly turf-forming algae, and were most abundant on an ocean reef flat (ORF) and in an inter-island channel near windward, sparsely populated islands. Calcarina density was higher on windward compared to leeward sides of ORFs, whereas Amphistegina density was similar on both sides of ORFs. These foraminifers were more common on the ocean side relative to the lagoon side of reef flats around a windward reef island, and both were rare or absent in nearshore zones around reef islands and on an ORF near windward, densely populated islands. Foraminiferal production rates varied with the degree to which habitats were subject to water motion and human influences. Highly productive sites (>10³ g CaCO₃ m⁻² year⁻¹) included an ORF and an inter-island channel near windward, sparsely populated islands, and a seaward area of a reef flat with no reef islands. Low-productivity sites (<10 g CaCO₃ m⁻² year⁻¹) included generally nearshore zones of lagoonal reef flats, leeward ORFs, and a windward ORF near densely populated islands. These results suggest that the distribution and production of LBFs were largely influenced by a combination of natural environmental factors, including water motion, water depth, elevation relative to the lowest tidal level at spring tide, and the distribution of suitable substratum. The presence of reef islands may limit the distribution and production of foraminifers by altering water circulation in nearshore environments. Furthermore, increased anthropogenic factors (population and activities) may adversely affect foraminiferal distribution and production.     

Coral reefs modify their seawater carbon chemistry – case study from a barrier reef (Moorea,French Polynesia)

Changes in the carbonate chemistry of coral reef waters are driven by carbon fluxes from two sources: concentrations of CO₂ in the atmospheric and source water, and the primary production/respiration and calcification/dissolution of the benthic community. Recent model analyses have shown that, depending on the composition of the reef community, the air‐sea flux of CO₂ driven by benthic community processes can exceed that due to increases in atmospheric CO₂ (ocean acidification). We field test this model and examine the role of three key members of benthic reef communities in modifying the chemistry of the ocean source water: corals, macroalgae, and sand. Building on data from previous carbon flux studies along a reef‐flat transect in Moorea (French Polynesia), we illustrate that the drawdown of total dissolved inorganic carbon (C_T) due to photosynthesis and calcification of reef communities can exceed the draw down of total alkalinity (A_T) due to calcification of corals and calcifying algae, leading to a net increase in aragonite saturation state (Ω_a). We use the model to test how changes in atmospheric CO₂ forcing and benthic community structure affect the overall calcification rates on the reef flat. Results show that between the preindustrial period and 1992, ocean acidification caused reef flat calcification rates to decline by an estimated 15%, but loss of coral cover caused calcification rates to decline by at least three times that amount. The results also show that the upstream–downstream patterns of carbonate chemistry were affected by the spatial patterns of benthic community structure. Changes in the ratio of photosynthesis to calcification can thus partially compensate for ocean acidification, at least on shallow reef flats. With no change in benthic community structure, however, ocean acidification depressed net calcification of the reef flat consistent with findings of previous studies.     

Differential thermal tolerance between algae and corals may trigger the proliferation of algae in coral reefs

Marine heatwaves can lead to rapid changes in entire communities, including in the case of shallow coral reefs the potential overgrowth of algae. Here we tested experimentally the differential thermal tolerance between algae and coral species from the Red Sea through the measurement of thermal performance curves and the assessment of thermal limits. Differences across functional groups (algae vs. corals) were apparent for two key thermal performance metrics. First, two reef‐associated algae species (Halimeda tuna and Turbinaria ornata) had higher lethal thermal limits than two coral species (Pocillopora verrucosa and Stylophora pistillata) conferring those species of algae with a clear advantage during heatwaves by surpassing the thermal threshold of coral survival. Second, the coral species had generally greater deactivation energies for net and gross primary production rates compared to the algae species, indicating greater thermal sensitivity in corals once the optimum temperature is exceeded. Our field surveys in the Red Sea reefs before and after the marine heatwave of 2015 show a change in benthic cover mainly in the southern reefs, where there was a decrease in coral cover and a concomitant increase in algae abundance, mainly turf algae. Our laboratory and field observations indicate that a proliferation of algae might be expected on Red Sea coral reefs with future ocean warming.