Response of a scleractinian coral, Stylophora pistillata, to iron and nitrate enrichment

The purpose of this study was to determine whether the addition of iron alone or in combination with nitrate affects growth and photosynthesis of the scleractinian coral, Stylophora pistillata, and its symbiotic dinoflagellates. For this purpose, we used three series of two tanks for a 3-week enrichment with iron (Fe), nitrate (N) and nitrate + iron (NFe). Two other tanks were kept as a control (C). Stock solutions of FeCl3 and NaNO3 were diluted to final concentrations of 6 nM Fe and 2 muM N and continuously pumped from batch tanks into the experimental tanks with a peristaltic pump. Results obtained showed that iron addition induced a significant increase in the areal density of zooxanthellae (ANOVA, p = 0.0013; change from 6.3 +/- 0.7 x 10(5) in the control to 8.5 +/- 0.6 x 10(5) with iron). Maximal gross photosynthetic rates normalized per surface area also significantly increased following iron enrichment (ANOVA, p = 0.02; change from 1.23 +/- 0.08 for the control colonies to 1.81 +/- 0.24 mu mol O-2 cm(-2) h(-1) for the iron-enriched colonies). There was, however, no significant difference in the photosynthesis normalized on a per cell basis. Nitrate enrichment alone (2 muM) did not significantly change the zooxanthellae density or the rates of photosynthesis. Nutrient addition (both iron and nitrogen) increased the cell-specific density of the algae (CSD) compared to the control (G-test, p = 0.3 x 10(-9)), with an increase in the number of doublets and triplets. CSD was equal to 1.70 +/- 0.04 in the Fe-enriched colonies, 1.54 +/- 0.12 in the N- and NFe-enriched colonies and 1.37 +/- 0.02 in the control. Growth rates measured after 3 weeks in colonies enriched with Fe, N and NFe were 23%, 34% and 40% lower than those obtained in control colonies (ANOVA. p = 0.011). (C) 2001 Elsevier Science B.V. All rights reserved.

The phylogeography and connectivity of the latitudinally widespread scleractinian coral Plesiastrea versipora in the Western Pacific

Whereas terrestrial animal populations might show genetic connectivity within a continent, marine species, such as hermatypic corals, may have connectivity stretching to all corners of the planet. We quantified the genetic variability within and among populations of the widespread scleractinian coral, Plesiastrea versipora along the eastern Australian seaboard (4145 km) and the Ryukyu Archipelago (Japan, 681 km) using sequences of internal transcribed spacers (ITS1-2) from ribosomal DNA. Geographic patterns in genetic variability were deduced from a nested clade analysis (NCA) performed on a parsimony network haplotype. This analysis allowed the establishment of geographical associations in the distribution of haplotypes within the network cladogram, therefore allowing us to deduce phylogeographical patterns based under models of restricted gene flow, fragmentation and range expansion. No significant structure was found among Ryukyu Archipelago populations. The lack of an association between the positions of haplotypes in the cladogram with geographical location of these populations may be accounted for by a high level of gene flow of P. versipora within this region, probably due to the strong Kuroshio Current. In contrast, strong geographical associations were apparent among populations of P. versipora along the south-east coast of Australia. This pattern of restricted genetic connectivity among populations of P. versipora on the eastern seaboard of Australia seems to be associated with the present surface ocean current (the East Australian Current) on this side of the south-western Pacific Ocean.

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.

Increased mortality and photoinhibition in the symbiotic dinoflagellates of the Indo-Pacific coral Stylophora pistillata (Esper) after summer bleaching

Coral bleaching (the loss of symbiotic dinoflagellates from reef-building corals) is most frequently caused by high-light and temperature conditions. We exposed the explants of the hermatypic coral Stylophora pistillata to four combinations of light and temperature in late spring and also in late summer. During mid-summer, two NOAA bleaching warnings were issued for Heron Island reef (Southern Great Barrier Reef, Australia) when sea temperature exceeded the NOAA bleaching threshold, and a 'mild' (in terms of the whole coral community) bleaching event occurred, resulting in widespread S. pistillata bleaching and mortality. Symbiotic dinoflagellate biomass decreased by more than half from late spring to late summer (from 2.5x10(6) to 0.8x10(6) dinoflagellates cm(2) coral tissue), and those dinoflagellates that remained after summer became photoinhibited more readily (dark-adapted F (V) : F (M) decreased to (0.3 compared with 0.4 in spring), and died in greater numbers (up to 17% dinoflagellate mortality compared with 5% in the spring) when exposed to artificially elevated light and temperature. Adding exogenous antioxidants (D-mannitol and L-ascorbic acid) to the water surrounding the coral had no clear effect on either photoinhibition or symbiont mortality. These data show that light and temperature stress cause mortality of the dinoflagellate symbionts within the coral, and that susceptibility to light and temperature stress is strongly related to coral condition. Photoinhibitory mechanisms are clearly involved, and will increase through a positive feedback mechanism: symbiont loss promotes further symbiont loss as the light microenvironment becomes progressively harsher.

Temperature-induced bleaching of corals begins with impairment of the CO2 fixation mechanism in zooxanthellae

The early effects of heat stress on the photosynthesis of symbiotic dinoflagellates (zooxanthellae) within the tissues of a reef-building coral were examined using pulse-amplitude-modulated (PAM) chlorophyll fluorescence and photorespirometry. Exposure of Stylophora pistillata to 33 and 34 degrees C for 4 h resulted in (1) the development of strong non-photochemical quenching (qN) of the chlorophyll fluorescence signal, (2) marked decreases in photosynthetic oxygen evolution, and (3) decreases in optimal quantum yield (F-v/F-m) of photosystern II (PSII), Quantum yield decreased to a greater extent on the illuminated surfaces of coral branches than on lower (shaded) surfaces, and also when high irradiance intensities were combined with elevated temperature (33 degrees C as opposed to 28 degrees C), qN collapsed in heat-stressed samples when quenching analysis was conducted in the absence of oxygen, Collectively, these observations are interpreted as the initiation of photoprotective dissipation of excess absorbed energy as heat (qN) and O-2-dependent electron flow through the Mehler-Ascorbate-Peroxidase cycle (MAP-cycle) following the point at which the rate of light-driven electron transport exceeds the capacity of the Calvin cycle. A model for coral bleaching is proposed whereby the primary site of heat damage in S, pistillata is carboxylation within the Calvin cycle, as has been observed during heat damage in higher plants, Damage to PSII and a reduction in F-v/F-m (i.e. photoinhibition) are secondary effects following the overwhelming of photoprotective mechanisms by light. This secondary factor increases the effect of the primary variable, temperature. Potential restrictions of electron flow in heat-stressed zooxanthellae are discussed with respect to Calvin cycle enzymes and the unusual status of the dinoflagellate Rubisco, Significant features of our model are that (1) damage to PSII is not the initial step in the sequence of heat stress in zooxanthellae, acid (2) light plays a key secondary role in the initiation of the bleaching phenomena.

Sweeper polyps of the coral Goniopora tenuidens (Scleractinia : Poritidae)

The massive coral Goniopora tenuidens can develop elongated sweeper polyps. These are thought to be involved in aggressive interactions with neighbouring benthic organ isms, like the sweeper tentacles of other corals. The cnidoms of sweeper polyps and ordinary polyps of G. tenuidens from the Great Barrier Reef were compared. Sweeper polyps had significantly greater densities of elongate holotrichous isorhizas (34577 +/- 3839/mg; mean +/- SD, n = 6) than ordinary polyps (936 +/- 371/mg; p < 0.05), while ordinary polyps had significantly greater densities of spirocysts (75994 +/- 15992/mg) than sweeper polyps (19469 +/- 7808/mg; p < 0.05). This suggests that sweeper polyps of G. tenuidens, like the sweeper tentacles of other corals, are modified for aggression, and that they probably act through nematocyst discharge. However, the scattered distribution of sweeper polyps observed on colonies of G. tenuidens in the field suggests that sweeper polyps may have other functions.

PAM chlorophyll fluorometry: a new in situ technique for stress assessment in scleractinian corals, used to examine the effects of cyanide from cyanide fishing

Sodium cyanide is being used on reefs in the Asia-Pacific region to capture live fish for the aquarium industry, and to supply a rapidly growing, restaurant-based demand, The effects of cyanide on reef biota have not been fully explored. To investigate its effect on hard corals, we exposed small branch lips of Stylophora pistillata and Acropora aspera to cyanide concentrations estimated to occur during cyanide fishing. Pulse amplitude modulation (PAM) chlorophyll fluorescence techniques were used to examine photoinhibition and photosynthetic electron transport in the symbiotic algae (zooxanthellae) in the tissues of the corals, These measurements were made in situ and in real time using a recently developed submersible PAM fluorometer. In S. pistillata. exposure to cyanide resulted in an almost complete cessation in photosynthetic electron transport rate. Both species displayed marked decreases in the ratio of variable fluorescence (F-v) to maximal fluorescence (F-m) (dark-adapted F-v/F-m), following exposure to cyanide, signifying a decrease in photochemical efficiency. Dark-adapted F-v/F-m recovered to normal levels in similar to 6 d, although intense tissue discolouration, a phenomenon well-recognised as coral 'bleaching' was observed during this period, Bleaching was caused by loss of zooxanthellae from the coral tissues, a well-recognised sub-lethal stress response of corals. Using the technique of chlorophyll fluorescence quenching analysis, corals exposed to cyanide did not show light activation of Calvin cycle enzymes and developed high levels of non-photochemical quenching (q(N)), signifying the photoprotective dissipation of excess light as heat, These features are symptomatic of the known properties of cyanide as an inhibitor of enzymes of the Calvin cycle. The results of this in situ study show that an impairment of zooxanthellar photosynthesis is; the site of cyanide-mediated toxicity, and is the cue that causes corals to release their symbiotic zooxanthellac following cyanide exposure. This study demonstrates the efficacy of PBM fluorometry as a new tool for in situ stress assessment in zooxanthellate scleractinian corals. (C) 1999 Elsevier Science Ltd. All rights reserved.

Historical overfishing and the recent collapse of coastal ecosystems

Ecological extinction caused by overfishing precedes all other pervasive human disturbance to coastal ecosystems, including pollution, degradation of water quality, and anthropogenic climate change. Historical abundances of large consumer species were fantastically large in comparison with recent observations. Paleoecological, archaeological, and historical data show that time lags of decades to centuries occurred between the onset of overfishing and consequent changes in ecological communities, because unfished species of similar trophic level assumed the ecological roles of overfished species until they too were overfished or died of epidemic diseases related to overcrowding. Retrospective data not only help to clarify underlying causes and rates of ecological change, but they also demonstrate achievable goals for restoration and management of coastal ecosystems that could not even be contemplated based on the limited perspective of recent observations alone.

Kinetics of photoacclimation in corals

Traditional models describing the relationship between photosynthesis (P) and irradiance (I) do not account for photoacclimation to short-term variation in irradiance. Here we develop and test a model that predicts the rate of photosynthesis under fluctuating irradiances at the scale of days to weeks. Using oxygen respirometry, we measured the rates of change in the P-I model parameters P-max (maximum rate of gross photosynthesis) and I-k (sub-saturation irradiance) of the photo-symbiotic coral Turbinaria mesenterina (Lamarck) following large and small increases and decreases in growth irradiance. We analyse the behaviour of the dynamic P-I model in turbid-water conditions using a dataset of 3-month continuous irradiance as the input variable. In response to upward or downward changes in experimental growth irradiance, I-k values decreased or increased exponentially, reaching new and stable levels within 5-10 days. I-k responded 4 times stronger than P-max to changes in growth irradiance. The kinetics of I-k did not show hysteresis, and changed in similar ways when irradiance was increased or decreased in small or large amounts. This suggests that mechanisms associated with photo-protection during increases in irradiance, and the maximisation of photosynthetic efficiency during decreases in irradiance, are equally potent. On the scale of months, the dynamic P-I model did not predict higher rates of photosynthesis than the static P-I model, but buffered the variation in photosynthesis during periods of reduced irradiance. Fourier analysis indicated that the kinetics of I-k closely matches the main periodicities in daily irradiance (1-2 weeks). The recorded kinetics of photoacclimation in the Turbinaria-zooxanthella symbiosis is comparable to that of free-living phytoplankton and faster than that of higher plants.

Sediment yield and impacts from river catchments to the Great Barrier Reef lagoon

Land use intensification is estimated to result in an overall increase in sediment delivery to the Great Barrier Reef lagoon by a factor of approximately four. Modelling suggests that, following land use intensification, croplands cause the greatest increase of sediment yield and sediment concentration, whereas erosion of grazing land is the main contemporary source of sediments, primarily owing to the large spatial extent of this land use. The spatial pattern of sediment yield to the coast after land use intensification is strongly correlated with the pattern under natural conditions, although the greatest increase is estimated to have occurred in the wet-dry catchments. Sediment transport and resuspension processes have led to the development of a strongly sediment-partitioned shelf, with modern mud-rich sediments almost exclusively restricted to the inner and inner-middle shelf, northward-facing embayments and in the lee of headlands. Elevated sediment concentrations increase the potential transport rates of nutrients and other pollutants. Whether increased sediment supply to the coastal zone has impacted on reefs remains a point of contention. More sediment load data need to be collected and analysed in order to make detailed estimates of catchment yields and establish the possible sediment impact on the Great Barrier Reef.

Late Quaternary uplift and subsidence of the west coast of Tanna, south Vanuatu, southwest Pacific: U-Th ages of raised coral reefs in the Median Sedimentary Basin

Twelve Late Quaternary TIMS U-Th ages are reported here from 10 coral samples collected in situ from five transgressive coral/algal raised reefs (height: max. 113 m, min. 8 m) and two raised lagoonal deposits (height: max. 18 m, min. 8 m) along and near the west coast of Tanna, which lies in the Median Sedimentary Basin of South Vanuatu, southwest Pacific. These reefs and raised lagoonal deposits represent several age groups: (i) 215 ka (marine oxygen-isotope stage 7) penultimate interglacial (highest elevation and oldest); (ii) one lagoonal deposit of ca 127 ka (marine oxygen-isotope stage 5e); (iii) three last interglacial reefs with ages 102, 89 and 81 ka (representing marine oxygen-isotope stages 5c, 5b and 5a, respectively, of the latter part of the last interglacial); (iv) a lagoonal deposit with a 92 ka age (5b); and (v) a Holocene reef (age >5.7-5.0 ka) (lowest elevation and youngest). A ca 4.9 ka regressive reef (at elevation of 1.5 m above sea-level) is consistent with an island-wide 6.5 m uplift (probably largely coseismic), and a probable further island-wide uplift occurred in the late Holocene. The U-series ages taken together with the heights of transgressive reefs show that uplift since 215 ka was, on average, at similar to0.52 mm/y; although since 5 ka the uplift rate was, on average, similar to1.6 mm/y (the assumption being that a 1.5 m above sea-level reef has a coseismic origin). Elevation of transgressive reefs 5a, 5b and 5c and their ages indicates an island-wide subsidence during the period ?124-89 ka (i.e. Late Quaternary uplift/subsidence was jerky). Late Quaternary uplift/subsidence on the northwest coast of Tanna is considered to be due to irregular thicknesses of crust being subducted beneath Tanna.