delta O-18, Sr/Ca and Mg/Ca records of Porites lutea corals from Leizhou Peninsula, northern South China Sea, and their applicability as paleoclimatic indicators

Combined seasonal to monthly resolution coral skeletal delta(18)O, Sr/Ca, and Mg/Ca records are reported for one modem and two late Holocene Porites lutea corals from a fringing reef at Leizhou Peninsula, the northern coast of the South China Sea (SCS). All the profiles for the period 1989-2000 reveal annual cycles well correlated with instrumental sea surface temperatures (SST), and display broad peaks in summer and narrow troughs in winter, reflecting seasonal growth rate variations. Calibration against instrumental SST yields the following equations: delta(18)O=-0.174(+/- 0.010)xSST(degrees C)-1.02(+/- 0.27) (MSWD=5.8), Sr/Ca-(mmol/mol)=-0.0424(+/- 0.0031)xSST(degrees C)+9.836(+/- 0.082) (MSWD=8.6), and Mg/Ca-(mmol/mol)=0.110(+/- 0.009)XSST(degrees C)+ 1.32(+/- 0.23) (MSWD=55). The scatter in the Mg/Ca-SST relationship is much larger than analytical uncertainties can account for, suggesting the presence of SST-unrelated components in the Mg/Ca variation. Calculated Sr/Ca-SST values for two later Holocene Porites lutea samples (U-series ages similar to 541 BC and similar to 487 AD, respectively) from the same reef suggest that SST in the SCS at similar to 541 BC was nearly as warm as in the 1990s (the warmest decade of the last century), but at similar to 487 AD, it was significantly cooler. This observation is consistent with climatic data reported in Chinese historic documents, confirming that the Sr/Ca-SST relationship is a reliable thermometer. Removing the SST component in the delta(18)O variation based on calculated Sr/Ca-SST values, the residual delta(18)O reflects the deviation of the Holocene seawater delta(18)O from the modem value, which is also a measure of the Holocene sea surface salinity (SSS) or the summer monsoon moisture level in mainland China. Such residual delta(18)O was close to zero at similar to 541 BC and -0.3 parts per thousand at similar to 487 AD, suggesting that it was as wet as in the 1990s at similar to 541 BC but significantly drier at similar to 487 AD in mainland China, which are also consistent with independent historic records. Calculated Mg/Ca-SST values for the two late Holocene corals are significantly lower than the Sr/Ca-SST values and are also in conflict with Chinese historic records, suggesting that coral Mg/Ca is not reliable proxy for SST. At comparable Sr/Ca ranges, fossil corals always display negative Mg/Ca offsets if compared with the modem coral of the same site. We interpret this observation as due to preferential loss of Mg during meteoric dissolution of cryptic Mg-calcite-bearing microbialites in the exposed fossil corals. Microbialites (MgO up to 17%, Sr only 100-300 ppm) are ubiquitous during reef-building processes and their presence in only a trace amount will have a significant impact on coral Mg/Ca ratios without detectable influence on coral Sr/Ca ratios. (c) 2004 Elsevier B.V. All rights reserved.

Mid-late Holocene monsoon climate retrieved from seasonal Sr/Ca and delta O-18 records of Porites lutea corals at Leizhou Peninsula, northern coast of South China Sea

South China Sea (SCS) is a major moisture source region, providing summer monsoon rainfall throughout Mainland China, which accounts for more than 80% total precipitation in the region. We report seasonal to monthly resolution Sr/Ca and delta(18)O data for five Holocene and one modem Porites corals, each covering a growth history of 9-13 years. The results reveal a general decreasing trend in sea surface temperature (SST) in the SCS from similar to 6800 to 1500 years ago, despite shorter climatic cycles. Compared with the mean Sr/Ca-SST in the 1990s (24.8 degrees C), 10-year mean Sr/Ca-SSTs were 0.9-0.5 degrees C higher between 6.8 and 5.0 thousand years before present (ky BP), dropped to the present level by similar to 2.5 ky BP, and reached a low of 22.6 degrees C (2.2 degrees C lower) by similar to 1.5 ky BP. The summer Sr/Ca-SST maxima, which are more reliable due to faster summer-time growth rates and higher sampling resolution, follow the same trend, i.e. being 1-2 degrees C higher between 6.8 and 5.0 ky BP, dropping to the present level by -2.5 ky BP, and reaching a low of 28.7 degrees C (0.7 degrees C lower) by similar to 1.5 ky BP. Such a decline in SST is accompanied by a similar decrease in the amount of monsoon moisture transported out of South China Sea, resulting in a general decrease in the seawater delta(18)O values, reflected by offsets of mean 6 180 relative to that in the 1990s. This observation is consistent with general weakening of the East Asian summer monsoon since early Holocene, in response to a continuous decline in solar radiation, which was also found in pollen, lake-level and loess/paleosol records throughout Mainland China. The climatic conditions similar to 2.5 and similar to 1.5 ky ago were also recorded in Chinese history. In contrast with the general cooling trend of the monsoon climate in East Asia, SST increased dramatically in recent time, with that in the 1990s being 2.2 degrees C warmer than that similar to 1.5 ky ago. This clearly indicates that the increase in the concentration of anthropogenic greenhouse gases played a dominant role in recent global warming, which reversed the natural climatic trend in East Asian monsoon regime. (c) 2004 Elsevier B.V. All rights reserved.

The effect of temperature on the size and population density of dinoflagellates in larvae of the reef coral Porites astreoides

Pre-settlement events play an important role in determining larval success in marine invertebrates with bentho-pelagic life histories, yet the consequences of these events typically are not well understood. The purpose of this study was to examine the pre-settlement impacts of different seawater temperatures on the size and population density of dinoflagellate symbionts in brooded larvae of the Caribbean coral Porites astreoides. Larvae were collected from P. astreoides at 14-20 m depth on Conch Reef (Florida) in June 2002, and incubated for 24 h at 15 temperatures spanning the range 25.1 degrees-30.0 degrees C in mean increments of 0.4 +/- 0.1 degrees C (+/- SD). The most striking feature of the larval responses was the magnitude of change in both parameters across this 5 degrees C temperature range within 24 h. In general, larvae were largest and had the highest population densities of Symbiodinium sp. between 26.4 degrees-27.7 degrees C, and were smallest and had the lowest population densities at 25.8 degrees C and 28.8 degrees C. Larval size and symbiont population density were elevated slightly (relative to the minimal values) at the temperature extremes of 25.1 degrees C and 30 degrees C. These data demonstrate that coral larvae are highly sensitive to seawater temperature during their pelagic phase, and respond through changes in size and the population densities of Symbiodinium sp. to ecologically relevant temperature signals within 24 h. The extent to which these changes are biologically meaningful will depend on the duration and frequency of exposure of coral larvae to spatio-temporal variability in seawater temperature, and whether the responses have cascading effects on larval success and their entry to the post-settlement and recruitment phase.

Characterizations of the major coral diseases of the Philippines: Ulcerative white spot disease and novel growth anomalies of Porites

Coral reefs are in decline worldwide and coral disease is a significant contributing factor. However, etiologies of coral diseases are still not well understood. In contrast with the Caribbean, extremely little is known about coral diseases in the Philippines. In 2005, off Southeast Negros Island, Philippines, I investigated relationships between environmental parameters and prevalence of the two most common coral diseases, ulcerative white spot (UWS) and massive Porites growth anomalies (MPGAs). Samples were collected along a disease prevalence gradient 40.5 km long. Principal component analyses showed prevalence of MPGAs was positively correlated with water column nitrogen, organic carbon of surface sediments, and colony density. UWS was positively correlated with water column phosphorus. This is the first quantitative evidence linking anthropogenically-impacted water and sediment to a higher prevalence of these diseases. Histological and cytological alterations were investigated by comparing tissues from two distinct types of MPGA lesions (types 1 and 2) and healthy coral using light and electron microscopy. Skeletal abnormalities and sloughing, swelling, thinning, and loss of tissues in MPGAs resembled tissues exposed to bacterial or fungal toxins. Both lesion types had decreases in symbiotic zooxanthellae, which supply nutrients to corals. Notable alterations included migrations of chromophore cells (amoebocytes) (1) nocturnally to outer epithelia to perform wound-healing, including plugging gaps and secreting melanin in degraded tissues, and (2) diurnally to the interior of the tissue possibly to prevent shading zooxanthellae in order to maximize photosynthate production. Depletion of melanin (active in wound healing) in type 2 lesions suggested type 2 tissues were overtaxed and less stable. MPGAs contained an abundance of endolithic fungi and virus-like particles, which may result from higher nutrient levels and play roles in disease development. Swollen cells and mucus frequently blocked gastrovascular canals (GVCs) in MPGAs. Type 1 lesions appeared to compensate for impeded flow of wastes and nutrients through these canals with proliferation of new GVCs, which were responsible for the observed thickened tissues. In contrast, type 2 tissues were thin and more degraded. Dysplasia and putative neoplasia were also observed in MPGAs which may result from the tissue regeneration capacity being overwhelmed.

Macroalgae Decrease Growth and Alter Microbial Community Structure of the Reef-Building Coral, Porites astreoides

With the continued and unprecedented decline of coral reefs worldwide, evaluating the factors that contribute to coral demise is of critical importance. As coral cover declines, macroalgae are becoming more common on tropical reefs. Interactions between these macroalgae and corals may alter the coral microbiome, which is thought to play an important role in colony health and survival. Together, such changes in benthic macroalgae and in the coral microbiome may result in a feedback mechanism that contributes to additional coral cover loss. To determine if macroalgae alter the coral microbiome, we conducted a field-based experiment in which the coral Porites astreoides was placed in competition with five species of macroalgae. Macroalgal contact increased variance in the coral-associated microbial community, and two algal species significantly altered microbial community composition. All macroalgae caused the disappearance of a γ-proteobacterium previously hypothesized to be an important mutualist of P. astreoides. Macroalgal contact also triggered: 1) increases or 2) decreases in microbial taxa already present in corals, 3) establishment of new taxa to the coral microbiome, and 4) vectoring and growth of microbial taxa from the macroalgae to the coral. Furthermore, macroalgal competition decreased coral growth rates by an average of 36.8%. Overall, this study found that competition between corals and certain species of macroalgae leads to an altered coral microbiome, providing a potential mechanism by which macroalgae-coral interactions reduce coral health and lead to coral loss on impacted reefs.

Development of Gene Expression Markers of Acute Heat-Light Stress in Reef-Building Corals of the Genus Porites

Coral reefs are declining worldwide due to increased incidence of climate-induced coral bleaching, which will have widespread biodiversity and economic impacts. A simple method to measure the sub-bleaching level of heat-light stress experienced by corals would greatly inform reef management practices by making it possible to assess the distribution of bleaching risks among individual reef sites. Gene expression analysis based on quantitative PCR (qPCR) can be used as a diagnostic tool to determine coral condition in situ. We evaluated the expression of 13 candidate genes during heat-light stress in a common Caribbean coral Porites astreoides, and observed strong and consistent changes in gene expression in two independent experiments. Furthermore, we found that the apparent return to baseline expression levels during a recovery phase was rapid, despite visible signs of colony bleaching. We show that the response to acute heat-light stress in P. astreoides can be monitored by measuring the difference in expression of only two genes: Hsp16 and actin. We demonstrate that this assay discriminates between corals sampled from two field sites experiencing different temperatures. We also show that the assay is applicable to an Indo-Pacific congener, P. lobata, and therefore could potentially be used to diagnose acute heat-light stress on coral reefs worldwide.

Characterizations of the Major Coral Diseases of the Philippines: Ulcerative White Spot Disease and Novel Growth Anomalies of Porites

Coral reefs are in decline worldwide and coral disease is a significant contributing factor. However, etiologies of coral diseases are still not well understood. In contrast with the Caribbean, extremely little is known about coral diseases in the Philippines. In 2005, off Southeast Negros Island, Philippines, I investigated relationships between environmental parameters and prevalence of the two most common coral diseases, ulcerative white spot (UWS) and massive Porites growth anomalies (MPGAs). Samples were collected along a disease prevalence gradient 40.5 km long. Principal component analyses showed prevalence of MPGAs was positively correlated with water column nitrogen, organic carbon of surface sediments, and colony density. UWS was positively correlated with water column phosphorus. This is the first quantitative evidence linking anthropogenically-impacted water and sediment to a higher prevalence of these diseases. Histological and cytological alterations were investigated by comparing tissues from two distinct types of MPGA lesions (types 1 and 2) and healthy coral using light and electron microscopy. Skeletal abnormalities and sloughing, swelling, thinning, and loss of tissues in MPGAs resembled tissues exposed to bacterial or fungal toxins. Both lesion types had decreases in symbiotic zooxanthellae, which supply nutrients to corals. Notable alterations included migrations of chromophore cells (amoebocytes) (1) nocturnally to outer epithelia to perform wound-healing, including plugging gaps and secreting melanin in degraded tissues, and (2) diurnally to the interior of the tissue possibly to prevent shading zooxanthellae in order to maximize photosynthate production. Depletion of melanin (active in wound healing) in type 2 lesions suggested type 2 tissues were overtaxed and less stable. MPGAs contained an abundance of endolithic fungi and virus-like particles, which may result from higher nutrient levels and play roles in disease development. Swollen cells and mucus frequently blocked gastrovascular canals (GVCs) in MPGAs. Type 1 lesions appeared to compensate for impeded flow of wastes and nutrients through these canals with proliferation of new GVCs, which were responsible for the observed thickened tissues. In contrast, type 2 tissues were thin and more degraded. Dysplasia and putative neoplasia were also observed in MPGAs which may result from the tissue regeneration capacity being overwhelmed.

Seawater carbonate chemistry and weight of two Atlantic corals Favia fragum and Porites astreoides during experiments, 2011

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]), and thus the saturation state of seawater with respect to aragonite. We investigated the relative importance of [HCO3] versus [CO3] 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 ?ar values, which were manipulated by both acid-addition at constant pCO2 (decreased total [HCO3] and [CO3]) and by pCO2 elevation at constant alkalinity (increased [HCO3], decreased [CO3]). 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] whether ?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] was nonlinear. A statistically significant decrease in calcification was only detected between Omega aragonite = <2.5 and Omega aragonite = 1.1-1.5, where calcification of new recruits was reduced by 22-37% per 1.0 decrease in Omega aragonite. 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.