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Coral Bleaching and Mortality Thresholds in the SE Gulf: Highest in the World

Bernhard M. Riegl, Sam J. Purkis, Ashraf S. Al-Cibahy, Suaad Al-Harthi, Edwin Grandcourt, Khalifa Al-Sulaiti, James Baldwin, and Alaa M. Abdel-Moati

6.1

Introduction

Bleaching is a stress reaction in corals, during which the symbiosis between corals and the algae (zooxanthellae) living in the coral cells breaks down. As a result, zooxanthellae are expelled, and the coral appears pale or even white (Fig. 6.1; Baker et al. 2008). The link between environmental variables and coral bleaching has been well-established in a variety of studies and synthesized in several places (Phinney et al. 2006; Baker et al. 2008; van Oppen and Lough 2009). Largescale and region-wide bleaching events, such as occur in the Gulf, have been clearly linked to unusually high temperatures and the accumulation of heat stress in corals. Other drivers, such as UV and water acidity can have compounding effects (Baker et al. 2008) and bleaching can also be caused by these factors alone, or other local drivers such as unusually cold temperatures (Saxby et al. 2003; Lajeunesse et al. 2007). However, it is heat stress that has been demonstrated as the most reliable predictor and dened time-integrated bleaching thresholds exist for various regions of the IndoPacic and the Caribbean (Berkelmans 2002b; Manzello et al. 2007; Berkelmans 2009). Since corals adapt to their local environment, bleaching thresholds vary within and among regions (Berkelmans
B.M. Riegl (*) S.J. Purkis National Coral Reef Institute, Nova Southeastern University, Dania Beach, FL, USA e-mail: rieglb@nova.edu; purkis@nova.edu A.S. Al-Cibahy S. Al-Harthi E. Grandcourt Environment Agency-Abu Dhabi, Abu Dhabi, United Arab Emirates e-mail: aalcibahy@ead.ae; salharthi@ead.ae; egrandcourt@ead.ae K. Al-Sulaiti J. Baldwin Qatar Gas, Ras Laffan Industrial City, Qatar e-mail: KAAlsulaiti@qatargas.com.qa; James.baldwin@qatargas.com.qa A.M. Abdel-Moati Ministry of Environment, Doha, Qatar e-mail: mamoati@moe.gov.qa

2002a, 2009; Manzello et al. 2007; Baker et al. 2008). As previous chapters in this book have demonstrated, Gulf corals exist in a uniquely extreme environment both with regards to extreme highs and lows, the regularly recurring long-time summertime highs (Chap. 4), however, unsurpassed among any region in the world. Corals do exist in other areas of the world in very hot environments, for example in the welldemonstrated case of tide-pools in Samoa (Birkeland et al. 2009) where temperatures can also reach 34.5C but only for few hours. Gulf corals are unique in being able to survive daily mean summertime temperatures in excess of 34 or even 35C for months (Chap. 4). Thus the question becomes just how much heat is necessary to bleach, or even kill these apparently uniquely adapted corals. Such information has much practical value, since it demonstrates just how much corals can acclimatize in a heating world (Sheppard 2003). During 2010, a major, region wide bleaching event occurred in the SE Gulf. While some previous region-wide beaching and mass mortality events (1996, 1998, 2002) are well-documented with regards to their effects on coral populations and communities (George and John 1999, 2000; Riegl 1999, 2001, 2002, 2003; Sheppard and Loughland 2002; Purkis and Riegl 2005; Burt et al. 2008; Riegl and Purkis 2009; Chap. 5), associated temperature data have been sporadic and coarse in resolution (Riegl 2002; Sheppard and Loughland 2002). During 2010, we had available both locally continuous temperature and coral monitoring data throughout the bleaching event which now allows the construction of bleaching threshold curves and estimates of differential impacts on all involved coral taxa.

6.2

Study Area and Methods

Study locations were situated in the SE Gulf, in the UAE and Qatar and are part of a routine monitoring program established over 2006/2007. Some of the sites are described in Chap. 4. For the present contribution, local bleaching thresholds were derived at Bu Tinah in the UAE from where an
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B.M. Riegl and S.J. Purkis (eds.), Coral Reefs of the Gulf: Adaptation to Climatic Extremes, Coral Reefs of the World 3, DOI 10.1007/978-94-007-3008-3_6, Springer Science+Business Media B.V. 2012

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Fig. 6.1 Bleached corals in Qatar, October 2010. (a) A blanched Plesiastrea versipora. Blanching is a stress state that occurs prior to bleaching and is detected by an overall paling of the corals color. (b) A partially bleached Cyphastrea microphthalma at the height of the 2010 bleaching event. Only a part of the coral is visibly bleached. (c) A partially bleached Porites harrisoni, (d) a completely bleached

Porites harrisoni not far from the specimen shown in (c) at the same time. Differences in clades of zooxanthellae, resulting in differential heat-susceptibility of the algae seem to be responsible (Baker et al. 2004). (e) A completely bleached Coscinarea columna. (f) A completely bleached Platygyra daedalea. All photographs from the same area and same time

almost complete temperature timeseries exists from 2007 through the end of 2010 (HoboTemp loggers recording hourly temperatures at 4 m depth) and at Fasht el Hurabi in Qatar from where a continuous temperature time series exists also from 2007 to 2010 (HoboTemp loggers recording at hourly intervals in 2007 and 2008, VEMCO Miniloggers recording at half hourly or hourly intervals in 2009 and 2010 at 5 m depth). Large-scale temperature records for the entire Gulf and world-wide, in order to evaluate the severity and extent of the thermal anomaly, were obtained from the Met Ofce, UK. For Gulf-wide analysis we employed the HadISST 11 gridded dataset, and for worldwide analysis

the HadISST and HadISST2 55 gridded products offered at www.hadobs.org (Rayner et al. 2003, 2006). Graphs were produced using the Erddap mapping facility (http://coastwatch.pfeg.noaa.gov/erddap/). Air temperature data for Sharjah airport were obtained from http://climexp.knmi.nl/. MODIS global SST data were obtained from http://neo.sci. gsfc.nasa.gov. Additionally, detailed information about coral mortality existed at both sites and in the general region from monitoring programs maintained in Qatar and the UAE. While obviously local variability exists in temperature among sites within the SE Gulf, the overall pattern is very similar (Chap. 4).

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We therefore investigated coral bleaching and mortality at several sites in order to detect any commonalities or differences in local population responses. Sites were at Ras Ghanada, Al Heel, Dalma and Bu Tina in Abu Dhabi and Fasht el Hurabi in Qatar. At these sites, phototransects were taken during and after the bleaching event and compared to similar transects taken earlier (Ras Ghanada; transect time series dating back to 2000, regular annual data since 2007). At Al Heel, and Dalma, phototransects exist dating back to 2007. Also at Fasht el Hurabi in Qatar, phototransects, as well as tagged coral colonies existed for the evaluation of bleaching. All coral monitoring took place at the end of the bleaching event, in the last week of September or the rst 2 weeks in October 2010. Thermal bleaching and mortality thresholds were derived according to Berkelmans (2002a, b, 2009) and Dunne (2002). At each location, the cumulative time (in days) at each temperature between 30C and 36C (average daily temperature, since either 24 or 48 daily readings were available) was calculated. It is known from previous bleaching events in the Gulf that temperatures need to exceed 35.5C for signicant mortality to occur (Riegl 2002). Bleaching thresholds were derived using Berkelmans corrected (2002b) equation: b c = Tn + [s / 5](Tb - Tn ), for Tb > Tt (6.1)

where LMST is long-term mean summer temperature (July, August, September) and Theating is the daily average temperature during bleaching. We did not have a long-term record available, since comparison of in-situ temperature records with remotely-sensed records did not coincide (see results). As closest proxy to a long-term record, we used the mean daily temperatures from 1 July to 30 September over the three non-bleaching years (2007, 2008, 2009) compared with the bleaching year (2010). The period 1 July-30 September encompasses peak temperatures (Fig. 6.9). Heating rate was calculated as HR = DHD / S days (Theating > LMST ) (6.3)

6.3

The 2010 Bleaching Event in the SE Gulf

Where bc is the predicted temperature value for the bleaching curve, s is a non-dimensional weighting factor that scales bleaching severity on a scale from 1 to 4 (we used 4 in this paper; 1 is mild bleaching, 4 the most severe), Tb is the temperature distribution in the coolest bleaching year (in our case, since only one bleaching event was captured, the temperature curve for 2010), and Tn is the temperature distribution in the warmest non-bleaching year. Tb and Tn are obtained by comparing graphed or tabulated values of the cumulative times spent at 30C or higher (see above). Tt is the threshold temperature, i.e. the highest temperature at which no bleaching occurs. Mortality curves were derived using Berkelmans (2009) method. These mortality curves are specic to the most susceptible genus, Acropora, which suffered almost 100% mortality (see below) in western Abu Dhabi (Dalma, Bu Tinah, Al Heel), and overall 6080% mortality in eastern Abu Dhabi (Ras Ghanada). We therefore followed Berkelmans (2009) in the assumption that TL50, the temperature value at which 50% mortality occurs in the indicator corals, is situated halfway between the bleaching threshold and the value causing 100% mortality. We also calculated degree heating days (DHD) and heating rate (HR), which Maynard et al. (2008a) have shown to be a useful stress indicators. DHD = S (Theating - LMST ) (6.2)

Coral bleaching was observed, depending on locality from August through October 2010. The event was comparable in severity with the 2002 bleaching event, when almost all corals bleached at least to some extent, but no wide-spread mass mortality was observed (Riegl 2003). Comparable data for Qatar are missing, since the 2002 event was not witnessed there, but presumably similar dynamics occurred. Depending on locality, the onset of bleaching occurred in the early to latter half of August and abated by mid September in eastern Abu Dhabi and in October in Qatar and western Abu Dhabi with colonies regaining color and no new bleaching observed. At all sites, at least some bleached corals persisted into October 2010. At Fasht el Hurabi, Qatar, most corals remained bleached until the third week of October (Figs. 6.1 and 6.2). At Ras Ghanada, most corals had regained color by the last week of September 2010. At all sites in Abu Dhabi, signicant mortality in Acropora was observed. At least at Ras Ghanada, this mortality was not uniquely due to bleaching but to a large extent, if not predominantly, to an outbreak of diseases, so-called white syndromes (Chap. 7) in the immediate aftermath of the bleaching. Many Acropora were able to recuperate from bleaching and coral diseases did not kill the entire local population. While some patches of Acropora escaped unscathed, much of the area suffered ~90% loss of Acropora tables of all sizes (recruits similarly affected as large colonies) (Fig. 6.2). At Ras Ghanada, an estimated 60% of the overall Acropora population was lost. Also at Al Heel and Dalma, Acropora was the worst-affected genus, with all colonies in the sampling sites suffering mortality (Fig. 6.2). At Bu Tinah and Fasht el Hurabi Acropora does not occur. Porites harrisoni, a dominant coral throughout the area, bleached heavily. At Ras Ghanada most colonies had regained most of their color by late September, with

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31 Iraq 30 29 Degrees latitude 28 27 26 25 24 23 46 Saudi Arabia


Qatar

Kuwait Iran

d
c,e

a United Arab Emirates

48

50

52 54 Degrees longitude

56

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Fig. 6.2 Taxon-specic bleaching levels during the 2010 event at selected sites in Abu Dhabi and Qatar, SE Gulf (locations detailed in inset map). (a, b) Relative frequency of taxa along point-intercept transects in Abu Dhabi and the bleached proportion. (c) Space cover in phototransects in Abu Dhabi and the bleached proportion. (d) Bleached

proportion of tagged experimental colonies in Qatar. (e) Space cover in phototransects and proportion of colonies that died in the period from the onset of bleaching to the monitoring period. The mortality value thus includes bleaching mortality and subsequent disease attacks (Chap. 7)

about 50% remaining blanched (i.e. somewhat paler than usual), at Al Heel and Dalma about half of the colonies remained bleached to various extents and at Fasht el Hurabi, Qatar, most colonies remained bleached white into the second week of October. Among the faviids, the genera Favia and Favites bleached less than Platygyra. Most Platygyra colonies still showed signs of bleaching (white, whitish or pale tissue patches as well as increased signs of recent mortality) and subsequent partial mortality. A clear gradation of bleaching recovery was observed from W to E, with the least impact and the most rapid regeneration at Ras Ghanada in the E, and the longest duration of the event at Fasht el Hurabi in Qatar, at the western extreme of the study area.

6.4 Temperature Regime During the 2010 Bleaching


6.4.1 Gulf-Wide Pattern

2010 was an overall hot year in the Gulf region. In September 2010, the Gulf was one of the warmest places in the worlds ocean (Fig. 6.3). A signicant positive temperature anomaly existed over most of the NW Indian Ocean, Red Sea and Gulf region that persisted throughout the bleaching months of September and October well into December 2010 (Fig. 6.4). The Gulf showed a positive anomaly between 1 and 3C over this period. The warm temperatures extended both over land and sea (HadISST and HadISST2 data; Rayner

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Fig. 6.3 Composite image of mean MODIS SST between Sep. 1 and Sep. 30, 2010. During this month and during the bleaching event, the Gulf was the warmest sea worldwide

Fig. 6.4 World-wide sea surface temperature anomalies during the 2010 Gulf bleaching (October, November 11 HadISST data) and shortly thereafter (November, December 55 HadISST2 data). Composite images courtesy The MetOfce (http://www.hadobs.org,

crown copyright). The anomalously warm temperatures in the Gulf region are clearly visible. The anomaly is dened as deviance from the monthly means in the period 19611990

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Fig. 6.5 Gulf-wide remotely-sensed sea surface temperature through the 2010 bleaching event (ad HadISST, eh MODIS). Coral monitoring sites for this study are indicated by white circles and correspond to those of Fig. 6.2. Sites where temperature was monitored as well are indicated by a lled white circle. Both HadISST and MODIS data suggest August as the hottest month and that bleaching was concentrated in

the central and SE Gulf. The Ras Ghanada study site experienced the lowest temperatures, which explains the observed lower bleaching impacts. The monthly averaged HadISST and the MODIS data underestimate peak local temperatures by about 1C. (HadISST data courtesy The Met Ofce, Crown copyright)

et al. (2003), obtained from the MetOfce website; http:// badc.nerc.ac.uk; crown copyright is acknowledged). Gulfwide, the strongest heating was restricted to its central and SE parts, as suggested by the HadISST and the MODIS datasets (Fig. 6.5). Highest 11 gridded temperatures were observed in August, with maximum heat in the area of the Abu Dhabi and Qatar offshore islands and banks. Average monthly SST (MODIS) in August/September was 33.5C in the northern Gulf, 32.78C in the eastern Gulf, but 34.45C in the central bleaching region. The eastern Abu Dhabi region (Ras Ghanada study site) received less heat. The HadISST dataset suggest that heat abated through September.

not the case at Fasht el Hurabi, where hourly maxima were lower in 2007 (35.8C).

6.5

Bleaching Threshold in the Gulf

6.4.2

Local Temperature Observations

2010 was overall a hot year with a long summer. Maximum locally-recorded hourly temperatures at BuTinah (Abu Dhabi) reached 36.4C and Fasht el Hurabi (Qatar) 36.1C. At Bu Tinah, these temperatures were recorded on 9th and 10th September, at Fasht el Hurabi daily from 12th to 15th August 2010 (Fig. 6.6). Average daily temperatures reached 35.48C at Bu Tinah on 3 September and 35.72C at Fasht el Hurabi on 15 August, which signicantly exceeded the long-term mean daily summer temperatures (Fasht el Hurabi 33.21C, Bu Tinah 33.78C) . DHD and DR values were higher in August than either in July or September, suggesting the onset of bleaching in August (Table 6.1). At Bu Tinah, short-term maximum temperatures were higher (36.6C) in 2007 but no bleaching was recorded. This was

Corals at Bu Tinah are exposed to higher temperature variability than corals at Fasht el Hurabi, as can also be seen from the summertime temperatures in Fig. 6.6. The shallow water body over the large Bu Tinah bank rapidly absorbs and loses more heat than the much smaller Fasht el Hurabi. As the cumulative temperature curves show, Gulf corals spend between 4 and 5 months every year at daily mean temperatures above 30C, about 2 months above 33C. These are the highest recorded temperature tolerances of corals anywhere. It is interesting to note that 2007 was almost as hot as 2010, yet no unequivocal bleaching records exist for the SE Gulf (but see ch. 4). However, coral reefs in Iran were subjected to signicant bleaching and subsequent mortality that year. The bleaching threshold curves for Bu Tinah and Fasht el Hurabi are similar (Fig. 6.7) and suggest that the maximum temperatures supported by Gulf corals are between 35.7 and 36.0C, above which bleaching is triggered. The annual cumulative temperature curves suggest that corals were subjected in 2010 to longer exposure to temperatures between 33 and 35C than in the other years (Table 6.2). The difference is much higher at Bu Tinah than at Fasht el Hurabi. Maximum temperatures in 2010 were similar to, or lower than, in 2007. This suggests that the total time exposure to sublethal temperatures in the range 3335C is the

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Fig. 6.6 Peak hourly temperatures observed in situ at temperature monitoring sites at (a) Bu Tinah (c) Fasht el Hurabi. Peak temperatures are circled red. (b, d) Deviation of mean daily temperatures in 2010 (red) from the mean daily temperatures 20072009 (blue). The in situ

temperature record shows more detail than the 11 gridded HadISST data (Fig. 6.4). Local temperatures are higher than in the 11 cell and the cooling trend observed in September in Fig. 6.4 did not correspond to all local data

Table 6.1 Degree heating days (DHD) and heating rate (HR) at sites in Abu Dhabi and Qatar. The peak in DHD in August suggests onset of bleaching in that month Fasht el Hurabi Bu Tinah DHD July 21.4 12.8 HR July 0.9 0.6 DHD August 74.3 36.1 HR August 2.4 1.2 DHD September 40.7 25.6 HR September 1.35 1.34

Fig. 6.7 Bleaching and mortality thresholds from Abu Dhabi and Qatar. Each datapoint represents the number of days spent at or above the temperature on the x-axis

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Table 6.2 Bleaching table and bleaching thresholds. Values in columns refer to the numbers of days spent at, or above the indicated temperature. Thus, the values of each higher temperature are included in those of the lower temperature. To obtain time spent at each temperature, the sum of all values of time spent at higher temperatures in the same line must be subtracted. Temperature data exist from 7 July 2010 to 31 December 2010, thus the time spent at temperatures <32C is under-represented. Bleaching thresholds were calculated using the method of Berkelmans (2002a, b). The values in columns are the cumulative days spent at or above the indicated temperature. Any time spent longer at these temperatures will lead to bleaching. Mortality thresholds (TL50) were calculated using the method of Berkelmans (2009). The TL50 is assumed to lie halfway between the bleaching threshold and the curve that produced 100% mortality. * = incomplete data record. Year Bu Tinah 2007 2008 2009 2010 Fasht el Hurabi 2007 2008 2009 2010 Bleaching Thresholds Fasht el Hurabi BuTinah Mean Mortality Thresholds Fasht el Hurabi BuTinah Mean >30C 79* 156 161 108 136 121 125 136 134 113 124 136 119 127 >31C 67* 130 135 108 119 105 105 115 113 110 112 115 113 114 >32C 56* 98 114 100 101 78 78 104 99 101 100 104 102 103 >33C 53* 67 67 91 76 57 22 93 79 86 83 93 89 91 >34C 38* 43 24 67 53 38 8 57 47 58 53 57 62 60 >35C 20* 2 1 23 27 4 0 33 26 18 22 33 21 27 >36C 0 0 0 0 0 0 0 0 0 0 0 0 0 0

determinant whether bleaching occurs or not, rather than the maximum temperatures. Although local variability is obvious in our dataset and has been demonstrated from many regions (Berkelmans 2002a, 2009; Manzello et al. 2007), we also evaluated a mean bleaching curve, which takes the average of the observations at Bu Tinah and Fasht el Hurabi. These mean values would suggest that, as a rule of thumb, Gulf corals are likely to bleach if they are exposed to more than 3 weeks at daily mean temperatures at or above 35C and between 8 and 9 weeks at or above 34C. Such a situation is only likely to occur in hot summers with heat wave conditions.

6.6

Discussion

Bleaching thresholds in the Gulf are the highest recorded in the world, and far exceed those recorded from the Great Barier Reef (Berkelmans 2002a, 2009), Galapagos (Podesta and Glynn 1997, 2001) and Caribbean (Manzello et al. 2007; Baker et al. 2008). This is clearly selected by the Gulf being in summer maybe the hottest place on earth supporting signicant coral growth (Fig. 6.3) forcing Gulf corals to have higher absolute temperature tolerance than anywhere else. Daily maximum temperatures survived by corals in lagoons in American Samoa can exceed 34C (Birkeland et al. 2009), however, only on an hourly basis, while in the

Gulf daily mean temperatures, peaking sometimes above 36C , can be above 34C for over a month. From nowhere else have similar long-term warm temperature extremes been recorded. On Australias Great Barrier Reef and in the Caribbean, bleaching thresholds are mostly anchored around maximum temperatures ~30C. While we fully appreciate that much local variability exists with regard to the onset of bleaching, and this is also shown in our dataset (Figs. 6.5 and 6.6), we nonetheless nd a mean bleaching curve, as we show in Fig. 6.8, very instructive. It suggests that a severe Gulf bleaching event of the type that causes severe Acropora die back, as treated in Chap. 5, requires about 3 weeks of exposure to >35C average daily temperatures. This agrees with ndings of Riegl (2002) for the 1996 and 1998 Acropora mass mortalities. Our ndings also correspond well with those of Manzello et al. (2007) who found that a combination of maximum SST, and the number of days spent above a certain threshold (30.5C in the Caribbean, 35C in the Gulf) are the most signicant determinants for the onset of a bleaching event. By varying parameter s in Eq. 6.1 (methods section), we can obtain some insight into the temperature loading required for milder bleaching to develop (Fig. 6.9). These results would suggest that, given previous heat-stress accumulation, about 1 week of average daily temperature means above 35C will cause mild symptoms of bleaching, while about 3 weeks of exposure will cause a severe bleaching event.

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Fig. 6.8 Bleaching thresholds derived for a variety of Caribbean and Great Barrier Reef locations by Berkelmans (2002a, b) and Manzello et al. (2007) compared to the mean bleaching threshold from the SE Gulf.

The overall tolerance envelope for the comparative areas is shown in grey. The differential between the curve and the bleaching threshold is directly related to the physiological capability of corals to adapt to heat

Fig. 6.9 Theoretical bleaching curves derived from Berkelmans (2002b) equation if the bleaching severity is altered while all other parameters are held equal. The value on the curve is parameter s of equation 6.1

While local differences exist with regards to stress-loading at lower temperatures (3032C), the pattern is very similar at the higher temperatures (3335C) that are responsible for the onset of bleaching (Berkelmans 2009). Thus, Gulf corals can support approximately 5C more heat than their relatives on the Great Barrier Reef and in the Caribbean. We also followed Berkelmans (2009) in developing mortality curves (Table 6.2). We nd that in the Gulf, mortality thresholds are close to the bleaching threshold (Table 6.2) but still exceed those known from any other region. Gulf corals are probably the most robust corals anywhere in the world with regards to bleaching, and mortality from bleaching.

The question arises whether the observed very high bleaching thresholds are already the result of acclimatization in response to rapidly recurring heat events (Maynard et al. 2008a, b; Riegl and Purkis 2009; Chap. 5). Riegl (2003) observed during the 2002 bleaching event that Acropora, which during the 1996 and 1998 events had bleached prior to and suffered more mortality than other taxa, locally (at Sir Abu Nuair) bleached later and to a lesser extent than other corals. Maynard et al. (2008a) observed that bleaching events indeed can lead to increased thermal tolerance. This may suggest a phenotypic shift in bleaching resistance in the surviving Gulf coral population. Temperatures in the Gulf region are indeed getting warmer and atmospheric temperatures in the region show more, and more closely

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Fig. 6.10 Air temperatures measured at Sharjah International Airport (2519 N, 5530 E). ( a ) Two mean annual temperature cycles based on the record from 1940 to 2010. Values are mean, 17% and 83% percentiles. ( b ) Anomalies with respect to the

annual cycle in ( a ). Known bleaching and coral mortality events (Purkis and Riegl 2005 ; Riegl and Purkis 2009 , Chap. 5 ) are shown by black dots . Another local bleaching event occurred in Abu Dhabi in 2011.

spaced, positive anomalies since the 1980s. This has been interpreted as a global warming signature (Nasrallah et al. 2004; Al-Rashidi et al. 2009) and may therefore have contributed to raising bleaching thresholds over the past decade due to selection caused by closely-spaced bleaching events (Fig. 6.10). The remarkable temperature tolerance of Gulf corals suggests that coral physiology is indeed capable of adapting to high temperatures and that some hope may exists that coral adaptation can track continuously increasing temperatures. If corals are not subjected to a host of other stressors, reasonable hope may exist that at least a subset of todays coral fauna may adapt to a heated world.
Acknowledgements Early phases of the monitoring project were funded by the WWF/EWF/Dolphin project and then sponsored by Abu Dhabi EAD, Qatar Gas, Qatar Ministry of Environment and NCRI at NSU. We thank R. Al-Mubarak, F. Launay, M. Chandler for support during that project.

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